In situ measurement of fine root water absorption in three temperate tree species—Temporal variability and control by soil and atmospheric factors

Miniature heat balance-sap flow gauges were used to measure water flows in small-diameter roots (3–4 mm) in the undisturbed soil of a mature beech–oak–spruce mixed stand. By relating sap flow to the surface area of all branch fine roots distal to the gauge, we were able to calculate real time water uptake rates per root surface area (J_s) for individual fine root systems of 0.5–1.0 m in length. Study aims were (i) to quantify root water uptake of mature trees under field conditions with respect to average rates, and diurnal and seasonal changes of J_s, and (ii) to investigate the relationship between uptake and soil moisture θ, atmospheric saturation deficit D, and radiation I. On most days, water uptake followed the diurnal course of D with a mid-day peak and low night flow. Neighbouring roots of the same species differed up to 10-fold in their daily totals of J_s (<100–2000 g m⁻² d⁻¹) indicating a large spatial heterogeneity in uptake. Beech, oak and spruce roots revealed different seasonal patterns of water uptake although they were extracting water from the same soil volume. Multiple regression analyses on the influence of D, I and θ on root water uptake showed that D was the single most influential environmental factor in beech and oak (variable selection in 77% and 79% of the investigated roots), whereas D was less important in spruce roots (50% variable selection). A comparison of root water uptake with synchronous leaf transpiration (porometer data) indicated that average water fluxes per surface area in the beech and oak trees were about 2.5 and 5.5 times smaller on the uptake side (roots) than on the loss side (leaves) given that all branch roots <2 mm were equally participating in uptake. Beech fine roots showed maximal uptake rates on mid-summer days in the range of 48–205 g m⁻² h⁻¹ (i.e. 0.7–3.2 mmol m⁻² s⁻¹), oak of 12–160 g m⁻² h⁻¹ (0.2–2.5 mmol m⁻² s⁻¹). Maximal transpiration rates ranged from 3 to 5 and from 5 to 6 mmol m⁻² s⁻¹ for sun canopy leaves of beech and oak, respectively. We conclude that instantaneous rates of root water uptake in beech, oak and spruce trees are above all controlled by atmospheric factors. The effects of different root conductivities, soil moisture, and soil hydraulic properties become increasingly important if time spans longer than a week are considered.     

Effects of P fertilisation and ectomycorrhizal fungal inoculation on early growth of eucalypt plantations in southern China

Eucalypt plantations in China have largely been established on soils that are low in phosphorus (P) and have few eucalypt-compatible ectomycorrhizal fungi. Effects of P application and ectomycorrhizal fungal inoculation on early tree growth in plantations of Eucalyptus urophylla Blake in Guangdong (Gaoyao) and E. globulus Labill. in Yunnan (Chuxiong) in southern China were investigated as part of a larger study. Application of superphosphate at establishment, in the presence of a basal fertiliser, increased early growth of E. urophylla and E. globulus. The optimum treatments for maximum stand volume at year 3 were 200 kg P ha^–1 which increased stand volume by 750% on the strongly acidic, P-deficient lateritic red oxisol at Gaoyao, and 40 kg P ha^–1 which increased stand volume by 55% on the mildly P-deficient red ultisol at Chuxiong, at 3 years. Superphosphate increased tree survival at Gaoyao as well as at Chuxiong. Nursery inoculation of eucalypt seedlings with ectomycorrhizal fungi significantly affected tree height and stand volume of the E. urophylla plantation, but the effect (positive or negative) was isolate-dependent and related to tree survival rate. A Laccaria isolate (CSIRO E4728) significantly increased stand volume by 27% at Gaoyao and a Scleroderma (MURU LH041) increased growth by 15% at Chuxiong at age 3 years. All isolates increased tree growth under P-limited soil conditions and only one isolate increased tree growth at marginal soil P. The results suggest that tree growth should be able to be optimised in plantations by the use of effective ectomycorrhizal fungi combined with a judicious fertilisation program at establishment.     

Vertical distribution and seasonal changes of fine and coarse root mass in Pinus kesiya Royle Ex.Gordon forest of three different ages

Seasonal changes and vertical distribution of fine (< 2 mm diameter) and coarse (2-10 mm diameter) root mass of Pinus kesiya and fine root and rhizome mass of herbaceous species, and root production were studied in the 6-, 15- and 23-year old Pinus kesiya forest stands at Shillong, in the Meghalaya state of north-east India. Maximum fine and coarse root mass of P. kesiya, and fine root and rhizome mass of the ground vegetation were recorded during the rainy season. The contribution of the tree fine roots in 0-10 cm soil layer declined from 51% in the 6-year old stand to about 33% in the older stands. The major proportion (63-88%) of herbaceous fine root and rhizome mass was concentrated in this soil layer in all the three stands. The majority (36-57%) of tree coarse roots were present in the 10-20 cm layer in all the stands. The biomass and necromass values in the case of fine roots were more or less equal in a given stand, but the coarse roots had 5 to 9 times more live than the dead mass. The proportion of herbaceous fine root mass to the total fine root mass declined from 54% in the 6-year old stand to 30-32% in the 15- and 23-year old stands. The mean total fine root mass (pine + herbaceous species) decreased from 417 g m^–2 in the 6-year old stand to 302 in 15-year and 322 g m^–2 in the 23-year old stand. Annual fine root production showed a marked decrease from 1055 g m^–2 in the 6-year old stand to 743 g m^–2 in the 23-year old stand, but coarse root production increased from 169 g m^–2 in the 6-year to 466 g m^–2  in the 23-year old stand; the total root production thus remained approximately constant.     

Boron requirements for Eucalyptus globulus seedlings

Although boron (B) deficiency limits the productivity of eucalypts in plantations in many parts of the world, the concentrations of foliar B used in the diagnosis of B deficiency vary greatly among studies. There has been a lack of reliable diagnosis standards for B deficiency in Eucalyptus species. Therefore, the present study investigated the relationship between internal and external B concentrations and growth of Eucalyptus globulus, the main commercial temperate eucalypt species. Seedlings were grown in a B-buffered solution culture (Amberlite IRA 743) from 0.03 to 8.35 M B. Boron deficiency symptoms appeared at day 5 in the nutrient solution containing less than 0.27 M B. The external critical B concentrations, estimated for the growth of shoots and roots, were 1.08 and 0.99 M B, respectively. The internal critical B concentration range in the youngest fully expanded leaf (YFEL) for shoot growth was 12–16 mg B kg^–1 dry weight. The internal critical B concentrations estimated in the present study have been successfully used in the diagnosis of B deficiency in E. globulus trees up to three years of age in south-east Asia.     

Assessing fine-root biomass and production in a Scots pine stand – comparison of soil core and root ingrowth core methods

Soil core and root ingrowth core methods for assessing fine-root (< 2 mm) biomass and production were compared in a 38-year-old Scots pine (Pinus sylvestris L) stand in eastern Finland. 140 soil cores and 114 ingrowth cores were taken from two mineral soil layers (0–10 cm and 10–30 cm) during 1985–1988. Seasonal changes in root biomass (including both Scots pine and understorey roots) and necromass were used for calculating fine-root production. The Scots pine fine-root biomass averaged annually 143 g/m² and 217 g/m² in the upper mineral soil layer, and 118 g/m² and 66 g/m² in the lower layer of soil cores and ingrowth cores, respectively. The fine-root necromass averaged annually 601 g/m² and 311 g/m² in the upper mineral soil layer, and 196 g/m² and 159 g/m² in the lower layer of soil cores and ingrowth cores, respectively. The annual fine-root production in a Scots pine stand in the 30 cm thick mineral soil layer, varied between 370–1630 g/m² in soil cores and between 210 – 490 g/m² in ingrowth cores during three years. The annual production calculated for Scots pine fine roots, varied between 330–950 g/m² in soil cores and between 110 – 610 g/m² in ingrowth cores. The horizontal and vertical variation in fine-root biomass was smaller in soil cores than in ingrowth cores. Roots in soil cores were in the natural dynamic state, while the roots in the ingrowth cores were still expanding both horizontally and vertically. The annual production of fine-root biomass in the Scots pine stand was less in root ingrowth cores than in soil cores. During the third year, the fine-root biomass production of Scots pine, when calculated by the ingrowth core method, was similar to that calculated by the soil core method. Both techniques have sources of error. In this research the sampling interval in the soil core method was 6–8 weeks, and thus root growth and death between sampling dates could not be accurately estimated. In the ingrowth core method, fine roots were still growing into the mesh bags. In Finnish conditions, after more than three growing seasons, roots in the ingrowth cores can be compared with those in the surrounding soil. The soil core method can be used for studying both the annual and seasonal biomass variations. For estimation of production, sampling should be done at short intervals. The ingrowth core method is more suitable for estimating the potential of annual fine-root production between different site types.     

The survival and development of inoculant ectomycorrhizal fungi on roots of outplanted Eucalyptus globulus Labill

The survival and development of two inoculant ectomycorrhizal fungi (Hebeloma westraliense Bough. Tom. and Mal. and Setchelliogaster sp. nov.) on roots of outplanted Eucalyptus globulus Labill. was examined at two expasture field sites in the south-west of Western Australia. Site 1 was a gravelly yellow duplex soil, and Site 2 was a yellow sandy earth. Plants were grown in steamed or unsteamed soil, in root bags designed as field containers for young growing trees. Three, 6 and 12 months after outplanting, plants were removed from these bags and assessed for dry weights of shoots and ectomycorrhizal colonization of roots.The inoculant ectomycorrhizal fungi (identified on the basis of the colour and morphology of their mycorrhizas) survived on roots of E. globulus for at least 12 months after outplanting at both field sites. At Site 1, however, colonization of new fine roots by the inoculant fungi was low (less than 20% of fine root length). Inoculation had no effect on the growth of E. globulus at this site. In contrast, at Site 2 the inoculant ectomycorrhizal fungi colonized up to 30–50% of new fine root length during the first 6 months after outplanting. There was a corresponding growth response to ectomycorrhizal inoculation at this site, with a close relationship (r²=0.82^**) between plant growth at 12 months and root colonization at 3 months. Plant growth at 12 months was related less closely with root colonization at 6 or 12 months. Root colonization by resident ectomycorrhizal fungi increased with time at both field sites. At Site 2, this increase appeared to be at the expense of colonization by the inoculant fungi, which was reduced to less than 10% of fine root length at 12 months. Steaming the soil had little effect on colonization by the inoculant ectomycorrhizal fungi at either field site, but decreased colonization by the resident ectomycorrhizal fungi.     

Root-promoting rhizobacteria in <Emphasis Type="Italic">Eucalyptus globulus</Emphasis> cuttings

Cloned Eucalyptus spp. plantations are based in greenhouse production of plants generated by vegetative propagation. Diverse studies have demonstrated that rhizospheric bacteria can stimulate plant growth, and more recently that they can increase rooting in vegetative material. Considering this potential, the objective of this study was to verify the effect of bacterial strains on rooting Eucalyptus globulus. A total of 132 bacterial strains isolated from the rhizosphere of E. globulus and Eucalyptus nitens were studied. The bacterial inoculums in a concentration of 4 × 10⁸ cfu/ml were applied to the rooting substrate at the cutting installation and 45 days after by irrigation. Rooting was evaluated on days 60 and 75 after cutting installation, considering the number of roots as well as their fibrosity and roots biomass. Of the 132 strains evaluated, 26 significantly increased cutting rooting in a range of 191.4–69.4% with respect to the control. Additionally, some strains stimulated the development of fine roots and incremented the roots biomass. The strains identificated that produced a rooting effect were: Bacillus firmus, Bacillus mycoides, Bacillus stearothermophilus, Bacillus subtilis, B. subtilis/amyloliquefaciens, Bacillus circulans, Brevibacillus brevis, Paenibacillus lautus and Stenotrophomona maltophilia. These first trials suggest the potential of these bacteria to be used in clonal production programs for E. globulus.     

Ontogeny of in vitro rooting processes in Eucalyptus globulus

Summary In the present work, histological changes observed at the base of Eucalyptus globulus shoots in in vitro culture are described. Shoots were placed on solidified Murashige and Skoog medium containing half the original salt concentration, the complete vitamin composition, 9.8 μM indolebutyric acid (IBA), and 30 gI⁻¹ agar, and were incubated in the dark for the first 7 d, followed by a 16-h photoperiod. In vitro-generated roots could be originated either from old vascular tissue or from newly formed xylem. The influence of the preexistent tissues on the neoformation process appeared to be varied. The medulla did not intervene directly, although there were abundant cellular divisions in response to the induction medium. On the other hand, the interruptions observed in the vascular cylinder of the stem suggested an influence of the interfascicular parenchyma, and therefore the medulla could have participated in the differentiation process. However, the cortical parenchyma showed most of the changes that lead to the formation of adventitious roots of E. globulus growing in vitro. Histological analysis suggests that vascular rays can also be formed in direct contact with the central cylinder of the stem, although they mainly originate from the cortical parenchyma.     

Carbohydrates as regulatory factors on the rooting of <Emphasis Type="Italic">Eucalyptus saligna</Emphasis> Smith and <Emphasis Type="Italic">Eucalyptus globulus</Emphasis> Labill

Comparisons between related species with different rooting capacities can provide insights into the mechanisms controlling adventitious root development. The availability of carbohydrates is often considered exclusively as an energetic requirement to drive root development; the major regulatory role in the process is often attributed to phytohormones, particularly auxin. The roles of light quantity (irradiance) and carbohydrate supply available to young aseptic donor-plants on the adventitious rooting response of Eucalyptus globulus (rooting recalcitrant) and Eucalyptus saligna (easy-to-root) were examined. The effects of the type of carbohydrate supply (sucrose or glucose) on the rooting response of cuttings was also evaluated. Light intensity supplied to mother-plants (30 or 60 mol m^–2 s^–1) had limited influence on the rooting response of both species, whereas dark periods were detrimental, particularly for E. globulus. In E. globulus, rooting was promoted by the absence of sucrose in donor-plant media. Presence of sucrose in donor plant medium promoted root number but did not affect rooting percentage of E. saligna. A positive effect of glucose on cutting rhizogenesis was found if this hexose was supplied during the root induction phase, followed by sucrose in the root formation step, especially for E. globulus. The same effect was not seen with fructose. The beneficial effect of glucose in the induction phase on root number was also evident under suboptimal auxin concentrations.     

Annual and seasonal changes in fine root biomass of a Quercus ilex L. forest

The biomass, production and mortality of fine roots (roots with diameter <2.5 mm) were studied in a typical Mediterranean holm oak (Quercus ilex L.) forest in NE Spain using the minirhizotron methodology. A total of 1212 roots were monitored between June of 1994 and March of 1997. Mean annual fine root biomass in the holm oak forest of Prades was 71±8 g m^–2 yr^–1. Mean annual production for the period analysed was 260+11 g m^–2 yr^–1. Mortality was similar to production, with a mean value of 253±3 g m^–2 yr^–1. Seasonal fine root biomass presented a cyclic behaviour, with higher values in autumn and winter and lower in spring and summer. Production was highest in winter, and mortality in spring. In summer, production and mortality values were the lowest for the year. Production values in autumn and spring were very similar. The vertical distribution of fine root biomass decreased with increasing depth except for the top 10–20 cm, where values were lower than immediately below. Production and mortality values were similar between 10 and 50 cm depth. In the 0–10 cm and the 50–60 cm depth intervals, both production and mortality were lower.     

Ecophysiological characterization of carnivorous plant roots: Oxygen fluxes,respiration, and water exudation

Various ecophysiological investigations on carnivorous plants in wet soils are presented. Radial oxygen loss from roots of Droseraceae to an anoxic medium was relatively low 0.02 – 0.07 mol(O₂) m^{– 2} s^–1 in the apical zone, while values of about one order of magnitude greater were found in both Sarracenia rubra roots and Genlisea violacea traps. Aerobic respiration rates were in the range of 1.6 – 5.6 mol kg^–1 (f.m.) s^–1 for apical root segments of seven carnivorous plant species and 0.4 – 1.1 mol kg^–1 (f.m.) s^–1 for Genlisea traps. The rate of anaerobic fermentation in roots of two Drosera species was only 5 – 14 % of the aerobic respiration. Neither 0.2 mM NaN₃ nor 0.5 mM KCN influenced respiration rate of roots and traps. In all species, the proportion of cyanide-resistant respiration was high and amounted to 65 – 89 % of the total value. Mean rates of water exudation from excised roots of 12 species ranged between 0.4 – 336 mm 3 kg^–1 (f.m.) s^–1 with the highest values being found in the Droseraceae. Exudation from roots was insensitive to respiration inhibitors. No significant difference was found between exudation rates from roots growing in situ in anoxic soil and those kept in an aerated aquatic medium. Carnivorous plant roots appear to be physiologically very active and well adapted to endure permanent soil anoxia.     

Development and function of Pisolithus and Scleroderma ectomycorrhizas formed in vivo with Allocasuarina,Casuarina and Eucalyptus

The effect of inoculating seedlings of Eucalyptus grandis, Allocasuarina littoralis and Casuarina equisetifolia with two isolates of Pisolithus and two isolates of Scleroderma from under eucalypts was examined in a glasshouse trial. Ectomycorrhizas formed extensively on Eucalyptus (23–46% fine roots ectomycorrhizal) and Allocasuarina (18–51% fine roots ectomycorrhizal). On Casuarina, the fungi were either unable to colonize the rhizosphere (one isolate of Pisolithus), or sheathed roots, resembling ectomycorrhizas, formed on 1–2% of the fine roots. Colonization of roots by one isolate of Scleroderma resulted in the death of Casuarina seedlings. Inoculation with fungi increased shoot dry weight by up to a factor of 32 (Eucalyptus), 4 (Allocasuarina) and 3 (Casuarina). Ectomycorrhizas formed in associations with Eucalyptus and Allocasuarina had fully differentiated mantles and Hartig nets in which the host and fungal cells were linked by an extensive fibrillar matrix. Sheathed roots in Casuarina lacked a Hartig net, and the epidermis showed a hypersensitive reaction resulting in wall thickening and cell death. The sheaths are described as mantles since the density and arrangement of the hyphae in the sheaths was similar to that in mantles of the eucalypt ectomycorrhizas. The intercellular carbohydrate matrix was not produced in the Casuarina mantle in association with Pisolithus, hence the mantle was not cemented to the root. These structures differ from poorly compatible associations described previously for Pisolithus and Eucalyptus. The anatomical data indicate that ectomycorrhizal assessment based on surface morphological features may be misleading in ecological studies because compatible and incompatible associations may not be distinguishable.     

Relationship between morphological and physiological responses to waterlogging and salinity in Sporobolus virginicus (L.) Kunth

The effects of waterlogging and salinity on morphological and physiological responses in the marsh grass Sporobolus virginicus (L.) Kunth were investigated in a 4×2 factorial experiment. Plants were subjected to four salinity levels (0, 100, 200 and 400 mol m^–3 NaCl) and two soil inundation conditions (drained and flooded) for 42 days. Flooding at 0 mol m^–3 NaCl caused initiation of adventitious surface roots, increased internal acration and plant height, induced alcohol dehydrogenase activity (ADH), and decreased belowground biomass and the number of culms per plant. Salinity increase from 0 to 400 mol m^–3 NaCl under drained conditions increased leaf and root proline concentrations and decreased photosynthesis, aboveground biomass, number of culms per plant and number of internodes per culm. Concurrent waterlogging and salinity induced ADH activity and adventitious surface roots but decreased plant height and aboveground biomass. Internal air space increased with waterlogging from 0 to 100 mol m^–3 NaCl but further increases in salinity to 400 mol m^–3 reduced air space. Combined waterlogging and salinity stresses, however, had no effect on photosynthesis or on the concentrations of proline in leaves or roots. These results are discussed in relation to the widespread colonization by S. virginicus of a wide range of coastal environments varying in soil salinity and in the frequency and intensity of waterlogging.     

Root distribution,standing crop biomass and belowground productivity in a semidesert in México

In a semidesert community in México (Zapotitlán de las Salinas, Puebla) the vertical distribution of roots and root biomass was estimated at 0–100 cm depth on two sampling dates, November 1995 (wet season) and January 1998 (dry season). Root productivity at 7 to 14.5 cm depth was estimated with the in-growth core technique every two months from March 1996 to February 1998. The relationship between environmental factors and seasonal root productivity was analyzed. Finally, we tested the effect of an irrigation equivalent to 20 mm of rain on root production. Seventy four percent of the total number of roots were found at 0-40 cm depth. Very fine roots (<1 mm diameter) were found throughout the soil profile (0-100 cm). In contrast, fine roots (1-3 mm diameter) were found only from 0–90 cm depth, and coarse roots (>3 mm diameter) from 0–60 cm depth. The root biomass was 971.5 g m^–2 (S.D. = 557.39), the very fine and fine roots representing 62.9% of the total. Total root productivity, as estimated with the ingrowth core technique, was 0.031 Mg ha^–1 over the dry season and 0.315 Mg ha^–1 over the wet season. Only very fine roots were obtained at all sampling dates. Rainfall was significantly correlated with very fine root production. The difference between fine root production in non-watered (0.054 g m^–2) and watered (0.429 g m^–2) treatments was significant. The last value was the same as that predicted for a rain of 20 mm, according to the exponential model describing the relation between the production of very fine roots and rainfall at the site.     

Development of root biomass in an Eucalyptus globulus plantation under different water and nutrient regimes

The distribution along the soil profile of Eucalyptus globulus root biomass was followed in a plantation in central Portugal at 1, 2 and 6 years after planting, using an excavation technique. The experimental design consisted of a control (C) and 3 treatments: application of solid fertilizers twice a year (F), irrigation without the application of fertilizers (I) and irrigation combined with liquid fertilizers (IL). Below- and above-ground biomass decreased as follows: IL>I>F>C. So, water stress limited growth more severely than nutrient stress. The roots rapidly colonized the top soil volume (0–20 cm depth) during the first year after planting. Fine root biomass 6 years after planting was 2.2, 1.8 and 1.6 times higher in IL treatment than it was respectively in control, and in F and I treatments. The distribution of fine roots along the soil profile 6 years after planting was more even in IL compared to the other treatments. However, fine roots in the top soil were more concentrated along the tree rows in the irrigated treatments than in the others. The proportion of below-ground biomass relative to the total tree biomass and the root/shoot ratio were higher in C than in the treatments at early growth stages. This pattern was not so clear 6 years after planting, due to the increased proportion of the tap root relative to total biomass, especially in the IL treatment.     

Root biomass changes after drainage and fertilization of a low-shrub pine bog

The stump and root systems of Scots pine (Pinus sylvestris) and field-layer vegetation were sampled before (1984) and three growing seasons after drainage and fertilization (1987) of a low-shrub pine bog. Average below-ground biomass of the field layer was 548 gDW m^–2 in 1984, with no significant treatment effects during experimentation. The stump-plus-root biomass of the pine stands was 1464 gDW m^–2 in the virgin state, and had increased to 1854 gDW m^–2 three years after the NPK-fertilizer treatment. The distribution over fractions also changed with this treatment. The fraction of fine roots ( < 1 mm) in stump-root biomass increased from 4% (56 gDW m^–2) to 11% (196 gDW m^–2), while the other compartments changed less. Total pine root length was 729 mm^–2 in 1984. Root length increased by 94% to 1380 mm^–2 on NPK-fertilized plots. Most of the fine pine roots were in the surface layer (0–10 cm), 79% in 1984 and 88% in 1987, and few pine roots were deeper than 20 cm. Maximum root length of fine pine roots ( < 1 mm) was estimated to be 2710 mm^–2 at about 800 gDW m^–2 (NPK treatment), and the corresponding maximum for small pine roots (=1–10 mm) was 227 mm^–2 at 809 gDW m^–2. Drainage stimulated net growth of fine roots, but this treatment also caused higher mortality rates of small roots. The fine roots responded to fertilization with higher net growth rate, and secondary growth of the large roots ( > 10 mm) was improved. The observed changes in root biomass and structure are explained as strategic adaptations to altered hydrological and nutritional circumstances in the root zone after drainage and fertilization.     

Effects of soil cadmium on Pinus sylvestris L. seedlings

The effects of Cd on the growth and distribution of Cd and mineral nutrients within plant tissues were investigated for Pinus sylvestris L. seedlings grown in mineral forest soil with increasing levels of Cd addition (0–100 mg kg^–1). Approximately 20% of added Cd was found to be extractable from sandy loam forest soil. Root growth was less affected by Cd than shoot growth, which showed a significant reduction in the 100 mg Cd kg^–1 treatment. Cadmium accumulated in roots up to 325 mg kg^–1. Decreased concentrations of K in needles and Ca in stems with increasing Cd levels suggest a disturbance of mineral nutrition as a result of Cd addition.     

Urban tree root systems and their survival near houses analyzed using ground penetrating radar and sap flow techniques

Root systems of two mature Field maple trees (Acer campestre L.) growing in both shaded and non-shaded sites, on clay soil in an urban environment, were analyzed by ground penetrating radar (GPR), light microscope and sap flow techniques. The ground surface above the root systems was covered by asphalt. However, a small piece of garden existed near the non-shaded tree, and root area of roots growing in this direction increased significantly, due to a presumed increase in available water and nutrients. However, no garden was present near the shaded tree, therefore roots remaining under the asphalt surface did not increase in area in any particular direction. Maximum rooting depth of shaded and exposed trees, as determined by GPR, was approximately 1.4 and 1.7 m, respectively. The trees utilized relatively large amounts of water for transpiration, i.e. 65–140 l per fine summer day and in average 10 m³ per growing season. However, transpiration expressed per root surface area (and/or whole root system enveloping area) was practically the same in both trees, i.e. 1 dm³ m^-2 d^-1 or almost 100 dm³ m^-2 per growing season. These figures represented about 50% of potential evapotranspiration when considering projected crown areas. Increased transpiration under long-term high evaporation demands may cause occasional local drying of soil around roots, associated with soil shrinking in clay, which can be followed by serious damage to buildings.     

Fine root dynamics and trace gas fluxes in two lowland tropical forest soils

Fine root dynamics have the potential to contribute significantly to ecosystem‐scale biogeochemical cycling, including the production and emission of greenhouse gases. This is particularly true in tropical forests which are often characterized as having large fine root biomass and rapid rates of root production and decomposition. We examined patterns in fine root dynamics on two soil types in a lowland moist Amazonian forest, and determined the effect of root decay on rates of C and N trace gas fluxes. Root production averaged 229 (±35) and 153 (±27) g m^?2 yr^?1 for years 1 and 2 of the study, respectively, and did not vary significantly with soil texture. Root decay was sensitive to soil texture with faster rates in the clay soil (k=?0.96 year^?1) than in the sandy loam soil (k=?0.61 year^?1), leading to greater standing stocks of dead roots in the sandy loam. Rates of nitrous oxide (N₂O) emissions were significantly greater in the clay soil (13±1 ng N cm^?2 h^?1) than in the sandy loam (1.4±0.2 ng N cm^?2 h^?1). Root mortality and decay following trenching doubled rates of N₂O emissions in the clay and tripled them in sandy loam over a 1‐year period. Trenching also increased nitric oxide fluxes, which were greater in the sandy loam than in the clay. We used trenching (clay only) and a mass balance approach to estimate the root contribution to soil respiration. In clay soil root respiration was 264–380 g C m^?2 yr^?1, accounting for 24% to 35% of the total soil CO₂ efflux. Estimates were similar using both approaches. In sandy loam, root respiration rates were slightly higher and more variable (521±206 g C m² yr^?1) and contributed 35% of the total soil respiration. Our results show that soil heterotrophs strongly dominate soil respiration in this forest, regardless of soil texture. Our results also suggest that fine root mortality and decomposition associated with disturbance and land‐use change can contribute significantly to increased rates of nitrogen trace gas emissions.