Dependence of the aboveground respiration of hinoki cypress (Chamaecyparis obtusa) on tree size

Nighttime respiration was measured at monthly intervals over one year on the aboveground parts of five sample trees in an 8-year-old hinoki cypress (Chamaecyparis obtusa (Sieb. et Zucc.) Endl.) stand, by an enclosed standing-tree method. The respiration rate rose rapidly from early spring to a maximum in June, and decreased abruptly in July and then gradually toward autumn and winter. The seasonal change in the respiration rate was synchronized with stem volume increment rather than with monthly mean air temperature. The respiration rate, r, of individual trees increased with increasing tree dimensions, such as stem volume, v(S), and stem girth at the base of the live crown, G(B). The dependence of respiration rate on tree size was successfully represented by a power function. The r - v(S) dependence was rather stronger than the r - G(B) (2) dependence, especially toward the end of the growing season (from July to September). The observed respiration rate was almost the same as the respiration rate corrected for the monthly mean air temperature. The annual respiration of individual trees was directly proportional to their phytomass or to its increment. Although the annual respiration of individual trees decreased proportionally to the square root of the leaf mass, it decreased abruptly in the range close to the smallest sample tree. Combining the monthly relationship between respiration rate and stem volume with the tree size distribution in the stand, the stand aboveground annual respiration was estimated to be 20.4 Mg CO(2) ha(-1) year(-1) (= 12.5 Mg dry mass ha(-1) year(-1)) for an aboveground biomass of 17.4 Mg ha(-1) with an annual increment of 6.51 Mg ha(-1) year(-1), i.e., the stand aboveground annual respiration amounted to the equivalent of 72% of the biomass or to almost twice the biomass increment.     

Patterns of root respiration rates and morphological traits in 13 tree species in a tropical forest

The root systems of forest trees are composed of different diameters and heterogeneous physiological traits. However, the pattern of root respiration rates from finer and coarser roots across various tropical species remains unknown. To clarify how respiration is related to the morphological traits of roots, we evaluated specific root respiration and its relationships to mean root diameter (D) of various diameter and root tissue density (RTD; root mass per unit root volume; gcm(-3)) and specific root length (SRL; root length per unit root mass; mg(-1)) of the fine roots among and within 14 trees of 13 species from a primary tropical rainforest in the Pasoh Forest Reserve in Peninsular Malaysia. Coarse root (2-269mm) respiration rates increased with decreasing D, resulting in significant relationships between root respiration and diameter across species. A model based on a radial gradient of respiration rates of coarse roots simulated the exponential decrease in respiration with diameter. The respiration rate of fine roots (<2mm) was much higher and more variable than those of larger diameter roots. For fine roots, the mean respiration rates for each species increased with decreasing D. The respiration rates of fine roots declined markedly with increasing RTD and increased with increasing SRL, which explained a significant portion of the variation in the respiration among the 14 trees from 13 species examined. Our results indicate that coarse root respiration in tree species follows a basic relationship with D across species and that most of the variation in fine root respiration among species is explained by D, RTD and SRL. We found that the relationship between root respiration and morphological traits provides a quantitative basis for separating fine roots from coarse roots and that the pattern holds across different species.     

Quantification of two-year-old hybrid poplar root systems: morphology, biomass, and (14)C distribution

Root morphology, biomass, and (14)C distribution were studied in two 2-year-old Populus trichocarpa x P. deltoides hybrids, which originated from hardwood cuttings, to determine the pattern of root distribution in a plantation and to refine methods for root recovery. The trees were labeled with (14)CO(2) and harvested after a 72-hour chase period. Roots attached to each labeled tree were analyzed for morphological traits at the time of harvest. Detached roots from within a 1-m(3) volume of soil surrounding each tree were separated from the soil and sorted on the basis of rooting depth and root diameter. Lateral roots > 2 mm in diameter had a largely horizontal orientation at their point of origin from the cutting and extended horizontally up to 4 m from the cutting. This resulted in considerable overlap of root systems in the plantation. Results from (14)C labeling indicated that 24 +/- 4% (+/- SD) of the carbon exported from branches-labeled within two weeks after branch budset-was translocated to the root system. Dilution of the root (14)C label indicated that from 0 (> 5 mm diameter roots) to 75% (< 2 mm diameter roots) of the roots recovered from within the 1-m(3) volume of soil surrounding a harvested tree originated from other trees. Total root biomass was 6 +/- 1 Mg ha(-1) for both hybrids. Sixty percent of the root biomass was recovered directly from excavation, 16% from coarse-sieving excavated soil, and 24% from re-sorting sieved soil. The study indicated that root growth of hybrid poplars may be rapid and extensive and that detailed sorting of soil subsamples substantially improves the recovery of fine roots < 2 mm in diameter.     

Changes in the relationship between tree size and aboveground respiration in field-grown hinoki cypress (Chamaecyparis obtusa) trees over three years

Respiration measurements of aerial parts of 18-year-old hinoki cypress (Chamaecyparis obtusa (Sieb. et Zucc.) Endl.) trees were made under field conditions over three years to study changing relationships with tree age between respiration and phytomass, phytomass increment, and leaf mass. The relationship between annual respiration (r(a)) and phytomass (w(T)) was approximated by a proportional function (r(a) = aw(T)), where the proportional constant (a) decreased year by year. The effect of time on the relationship between annual respiration and phytomass of each sample tree was fitted by a power function. Respiration of the tree suppressed by the canopy decreased year by year, but respiration of the other trees increased slightly with age. The relationship between annual respiration and leaf mass was also approximated by a generalized power function. Excluding the suppressed tree, the relationship between annual respiration (r(a)) and the annual increment of aboveground phytomass (Deltaw(T)) was described by a proportional function (r(a) = 2.27Deltaw(T)), where the proportional constant, 2.27, was independent of sample tree and year, indicating that about 2.3 times of the annual aboveground phytomass increment equivalent was respired annually. For any tree, the time constant relationships between annual respiration and leaf mass and phytomass increment for different-sized trees were similar to the corresponding time continuum relationships. In contrast, the time continuum relationship between annual respiration and phytomass differed from the time constant relationship, indicating that respiration of less active woody tissue contributed significantly to aboveground respiration. Based on the relationship between tree size and annual respiration, annual aboveground stand respiration was estimated to be 25.0, 26.9, and 25.8 Mg(dm) ha(-1) year(-1) for the three consecutive years, respectively, and the corresponding aboveground stand biomass was 60.0, 69.0, and 76.8 Mg(dm) ha(-1).     

Coarse and fine root respiration in aspen (Populus tremuloides)

Coarse and fine root respiration rates of aspen (Populus tremuloides Michx.) were measured at 5, 15 and 25 degrees C. Coarse roots ranged from 0.65 to 4.45 cm in diameter, whereas fine roots were less than 5 mm in diameter. To discriminate between maintenance and growth respiration, root respiration rates were measured during aboveground growing periods and dormant periods. An additional measurement of coarse root respiration was made during spring leaf flush, to evaluate the effect of mobilization of resources for leaf expansion on root respiration. Fine roots respired at much higher rates than coarse roots, with a mean rate at 15 degrees C of 1290 micromol CO2 m-3 s-1 during the growing period, and 660 micromol CO2 m-3 s-1 during the dormant period. The temperature response of fine root respiration rate was nonlinear: mean Q10 was 3.90 for measurements made at 5-15 degrees C and 2.19 for measurements made at 15-25 degrees C. Coarse root respiration rates measured at 15 degrees C in late fall (dormant season) were higher (370 micromol CO2 m-3 s-1) than rates from roots collected at leaf flush and early summer (200 micromol CO2 m-3 s-1). The higher respiration rates in late fall, which were accompanied by decreased total nonstructural carbohydrate (TNC) concentrations, suggest that respiration rates in late fall included growth expenditures, reflecting recent radial growth. Neither bud flush nor shoot growth of the trees caused an increase in coarse root respiration or a decrease in TNC concentrations, suggesting a limited role of coarse roots as reserve storage organs for spring shoot growth, and a lack of synchronization between above- and belowground growth. Pooling the data from the coarse and fine roots showed a positive correlation between nitrogen concentration and respiration rate.     

14C Allocation in tree-soil systems

We studied whole-tree C allocation with special emphasis on the quantification of C allocation to roots and root respiration. To document seasonal patterns of C allocation, 2-year-old hybrid poplar trees greater than 3 m tall were labeled with (14)CO(2) in a large Plexiglas chamber in the field, in July and September. Climate and CO(2) concentration were controlled to track ambient conditions during labeling. Individual tree canopy CO(2) assimilation averaged 3.8 micromol CO(2) m(-2) s(-1) (12.9 g C day(-1) tree(-1)) in July and 6.2 micromol CO(2) m(-2) s(-1) (9.8 g C day(-1) tree(-1)) in September. Aboveground dark respiration was 12% of net daytime C fixation in July and 15% in September. Specific activity of root-soil respiration peaked 2 days after labeling and stabilized to less than 5% of maximum 2 weeks later. Low specific activity of root-soil respiration and a labeled pool of root C demonstrated that current photosynthate was the primary source of C for root growth and maintenance during the growing season. Root respiration averaged 20% of total soil respiration in both July and September based on the proportion of labeled C respired to labeled C fixed. In July, 80% of the recovered (14)C was found above ground and closely resembled the weight distribution of the growing shoot. By September, 51% of the recovered (14)C was in the root system and closely resembled the weight distribution of different size classes of roots. The finding that the distribution of biomass and (14)C were similar verified that the C introduced during labeling followed normal seasonal translocation pathways. Results are compared to smaller scale labeling studies and the suitability of the approach for studying long-term C fluxes is discussed.     

Relating coarse root respiration to root diameter in clonal Eucalyptus stands in the Republic of the Congo

Root respiration is an important component of the carbon balance of a forest ecosystem. We measured CO2 efflux of excised fine roots and intact coarse roots in 3-, 4- and 13-year-old Eucalyptus stands in the region of Pointe-Noire, Republic of the Congo. A transportable and adaptable closed chamber gas exchange system directly measured CO2 efflux of roots from 0.5 to 32 mm in diameter. Fluxes were corrected for measurement system leaks and normalized to a reference temperature of 30 degrees C. Mean fine root respiration rates at the reference temperature varied between 8.5 and 10.8 micromol CO2 kg(-1) s(-1) depending on the stand. Coarse root respiration was strongly negatively correlated to root diameter. We propose a model based on a radial gradient of respiratory activity within the root to simulate the exponential decrease in respiration with diameter. Although many sources of uncertainty in the measurements remain, as discussed in this paper, these results provide a basis for scaling up organ-level root respiration measurements to the tree and stand levels.     

Foliage, fine-root, woody-tissue and stand respiration in Pinus radiata in relation to nitrogen status

We measured respiration of 20-year-old Pinus radiata D. Don trees growing in control (C), irrigated (I), and irrigated + fertilized (IL) stands in the Biology of Forest Growth experimental plantation near Canberra, Australia. Respiration was measured on fully expanded foliage, live branches, boles, and fine and coarse roots to determine the relationship between CO(2) efflux, tissue temperature, and biomass or nitrogen (N) content of individual tissues. Efflux of CO(2) from foliage (dark respiration at night) and fine roots was linearly related to biomass and N content, but N was a better predictor of CO(2) efflux than biomass. Respiration (assumed to be maintenance) per unit N at 15 degrees C and a CO(2) concentration of 400 micro mol mol(-1) was 1.71 micro mol s(-1) mol(-1) N for foliage and 11.2 micro mol s(-1) mol(-1) N for fine roots. Efflux of CO(2) from stems, coarse roots and branches was linearly related to sapwood volume (stems) or total volume (branches + coarse roots) and growth, with rates for maintenance respiration at 15 degrees C ranging from 18 to 104 micro mol m(-3) s(-1). Among woody components, branches in the upper canopy and small diameter coarse roots had the highest respiration rates. Stem maintenance respiration per unit sapwood volume did not differ among treatments. Annual C flux was estimated by summing (1) dry matter production and respiration of aboveground components, (2) annual soil CO(2) efflux minus aboveground litterfall, and (3) the annual increment in coarse root biomass. Annual C flux was 24.4, 25.3 and 34.4 Mg ha(-1) year(-1) for the C, I and IL treatments, respectively. Total belowground C allocation, estimated as the sum of (2) and (3) above, was equal to the sum of root respiration and estimated root production in the IL treatment, whereas in the nutrient-limited C and I treatments, total belowground C allocation was greater than the sum of root respiration and estimated root production, suggesting higher fine root turnover or increased allocation to mycorrhizae and root exudation. Carbon use efficiency, the ratio of net primary production to assimilation, was similar among treatments for aboveground tissues (0.43-0.50). Therefore, the proportion of assimilation used for construction and maintenance respiration on an annual basis was also similar among treatments.     

Fine root respiration in the mangrove Rhizophora mangle over variation in forest stature and nutrient availability

Root respiration uses a significant proportion of photosynthetically fixed carbon (C) and is a globally important source of C liberated from soils. Mangroves, which are an important and productive forest resource in many tropical and subtropical countries, sustain a high ratio of root to shoot biomass which may indicate that root respiration is a particularly important component in mangrove forest carbon budgets. Mangroves are often exposed to nutrient pollution from coastal waters. Here we assessed the magnitude of fine root respiration in mangrove forests in Belize and investigated how root respiration is influenced by nutrient additions. Respiration rates of excised fine roots of the mangrove, Rhizophora mangle L., were low (4.01 +/- 0.16 nmol CO(2) g(-1) s(-1)) compared to those measured in temperate tree species at similar temperatures. In an experiment where trees where fertilized with nitrogen (N) or phosphorus (P) in low productivity dwarf forests (1-2 m height) and more productive, taller (4- 7 m height) seaward fringing forests, respiration of fine roots did not vary consistently with fertilization treatments or with forest stature. Fine roots of taller fringe trees had higher concentrations of both N and P compared to dwarf trees. Fertilization with P enhanced fine root P concentrations in both dwarf and fringe trees, but reduced root N concentrations compared to controls. Fertilization with N had no effect on root N or P concentrations. Unlike photosynthetic C gain and growth, which is strongly limited by P availability in dwarf forests at this site, fine root respiration (expressed on a mass basis) was variable, but showed no significant enhancements with nutrient additions. Variation in fine root production and standing biomass are, therefore, likely to be more important factors determining C efflux from mangrove sediments than variations in fine root respiration per unit mass.     

Maintenance and growth respiration of the aboveground parts of young field-grown hinoki cypress (Chamaecyparis obtusa)

Aboveground respiration of five 8-year-old trees of field-grown hinoki cypress (Chamaecyparis obtusa (Sieb. et Zucc.) Endl.) was nondestructively measured at monthly intervals over 1 year with an enclosed standing tree method. The relationship between monthly specific respiration rate and monthly mean relative growth rate at the individual tree level was described by a linear equation. During the dormant season, respiration was used mainly for maintenance purposes, whereas during the growing season, more than 40% of the respiration was used for growth purposes, i.e., 60 to 70% in May. We conclude that annual maintenance and growth respiration of a tree are directly proportional to the aboveground phytomass and its annual increment, respectively. The maintenance coefficient was estimated to be 0.504 +/- 0.039 (SE) kg kg(-1) year(-1), indicating that the amount respired for maintaining already existing phytomass was equivalent to about half of the existing phytomass. The growth coefficient was estimated to be 0.772 +/- 0.043 (SE) kg kg(-1), indicating that the amount respired for constructing new phytomass was equivalent to about three-fourths of the annual phytomass increment. The annual stand maintenance and growth respiration were, respectively, 8.8 Mg ha(-1) year(-1) for an aboveground biomass of 17.4 Mg ha(-1) and 5.0 Mg ha(-1) year(-1) for an annual stand aboveground biomass increment of 6.5 Mg ha(-1) year(-1). About two-thirds of the total respiration was used to maintain already existing biomass, and about one-third was used to construct new biomass.     

Long-term respiration in relation to growth and maintenance processes of the aboveground parts of a hinoki forest tree

Over a three-year period, CO(2) exchange rates were measured continuously on the aboveground parts of a 13-year-old hinoki (Chamaecyparis obtusa (Sieb. et Zucc.) Endl.) tree in the field, using an open gas-exchange system tracking ambient temperature. The relationship between daily aerial respiratory consumption and daily gross production, which was analyzed monthly, closely resembled McCree's equation. The value of the coefficient of growth respiration ranged between 0.0 in winter and 0.16 in summer and increased with increasing monthly mean temperature. A clockwise loop was observed for monthly change in the relationship between the coefficient of growth respiration and temperature. Maintenance respiration could be formulated as a power function of aboveground dry weight. The exponent of the equation ranged from 0.3 to 1.1. A value of 1.1 in May and June, when trees were growing most actively, indicated that maintenance respiration was directly proportional to aboveground weight. In March, April, July, and August, maintenance respiration was not proportional to aboveground weight, but it was closely proportional to surface area. The exponent value exhibited seasonal change with a clockwise loop in response to monthly average temperature. During the dormant season, respiration was used only for maintenance purposes, whereas during the growing season both growth and maintenance respiration occurred. Annual growth and maintenance respiration increased with increasing tree age. The average annual contribution to total respiratory consumption was 21% for growth respiration and 79% for maintenance respiration.     

Elevational change in woody tissue CO2 efflux in a tropical mountain rain forest in southern Ecuador

Much uncertainty exists about the magnitude of woody tissue respiration and its environmental control in highly diverse tropical moist forests. In a tropical mountain rain forest in southern Ecuador, we measured the apparent diurnal gas exchange of stems and coarse roots (diameter 1-4 cm) of trees from representative families along an elevational transect with plots at 1050, 1890 and 3050 m a.s.l. Mean air temperatures were 20.8, 17.2 and 10.6 degrees C, respectively. Stem and root CO(2) efflux of 13 to 21 trees per stand from dominant families were investigated with an open gas exchange system while stand microclimate was continuously monitored. Substantial variation in respiratory activity among and within species was found at all sites. Mean daily CO(2) release rates from stems declined 6.6-fold from 1.38 micromol m(-2) s(-1) at 1050 m to 0.21 micromol m(-2) s(-1) at 3050 m. Mean daily CO(2) release from coarse roots decreased from 0.35 to 0.20 micromol m(-2) s(-1) with altitude, but the differences were not significant. There was, thus, a remarkable shift from a high ratio of stem to coarse root respiration rates at the lowest elevation to an apparent equivalence of stem and coarse root CO(2) efflux rates at the highest elevation. We conclude that stem respiration, but not root respiration, greatly decreases with elevation in this transect, coinciding with a substantial decrease in relative stem diameter increment and a large increase in fine and coarse root biomass production with elevation.     

Root architecture in relation to tree-soil-crop interactions and shoot pruning in agroforestry

Desirable root architecture for trees differs between sequential and simultaneous agroforestry systems. In sequential systems extensive tree root development may enhance nutrient capture and transfer to subsequent crops via organic pools. In simultaneous systems tree root development in the crop root zone leads to competition for resources.Fractal branching models provide relationships between proximal root diameter, close to the tree stem, and total root length or surface area. The main assumption is that a root branching proportionality factor is independent of root diameter. This was tested in a survey of 18 multipurpose trees growing on an acid soil in Lampung (Indonesia). The assumption appeared valid for all trees tested, for stems as well as roots. The proportionality factor showed a larger variability in roots than in stems and the effects of this variabilily should be further investigated. A simple index of tree root shallowness is proposed as indicator of tree root competitiveness, based on superficial roots and stem diameter.Pruning trees is a major way to benefit from tree products and at the same time reduce above-ground competition between trees and crops. It may have negative effects, however, on root distribution and enhance below-ground competition. In an experiment with five tree species, a lower height of stem pruning led to a larger number of superficial roots of smaller diameter, but had no effect on shoot:root ratios or the relative importance of the tap root.     

Variation in sugar maple root respiration with root diameter and soil depth

Root respiration may account for as much as 60% of total soil respiration. Therefore, factors that regulate the metabolic activity of roots and associated microbes are an important component of terrestrial carbon budgets. Root systems are often sampled by diameter and depth classes to enable researchers to process samples in a systematic and timely fashion. We recently discovered that small, lateral roots at the distal end of the root system have much greater tissue N concentrations than larger roots, and this led to the hypothesis that the smallest roots have significantly higher rates of respiration than larger roots. This study was designed to determine if root respiration is related to root diameter or the location of roots in the soil profile. We examined relationships among root respiration rates and N concentration in four diameter classes from three soil depths in two sugar maple (Acer saccharum Marsh.) forests in Michigan. Root respiration declined as root diameter increased and was lower at deeper soil depths than at the soil surface. Surface roots (0-10 cm depth) respired at rates up to 40% greater than deeper roots, and respiration rates for roots < 0.5 mm in diameter were 2.4 to 3.4 times higher than those for roots in larger diameter classes. Root N concentration explained 70% of the observed variation in respiration across sites and size and depth classes. Differences in respiration among root diameter classes and soil depths appeared to be consistent with hypothesized effects of variation in root function on metabolic activity. Among roots, very fine roots in zones of high nutrient availability had the highest respiration rates. Large roots and roots from depths of low nutrient availability had low respiration rates consistent with structural and transport functions rather than with active nutrient uptake and assimilation. These results suggest that broadly defined root classes, e.g., fine roots are equivalent to all roots < 2.0 mm in diameter, do not accurately reflect the functional categories typically associated with fine roots. Tissue N concentration or N content (mass x concentration N) may be a better indicator of root function than root diameter.     

Fine root dynamics in a developing Populus deltoides plantation

A closely spaced (1 x 1 m) cottonwood (Populus deltoides Bartr.) plantation was established to evaluate the effects of nutrient availability on fine root dynamics. Slow-release fertilizer (17:6:12 N,P,K plus micronutrients) was applied to 225-m(2) plots at 0, 50, 100 and 200 kg N ha(-1), and plots were monitored for two growing seasons. Fine root production, mortality, live root standing crop and life span were analyzed based on monthly minirhizotron observations. Fine root biomass was measured in soil cores. Fine root dynamics were controlled more by temporal, depth and root diameter factors than by fertilization. Cumulative fine root production and mortality showed strong seasonal patterns; production was greatest in the middle of the growing season and mortality was greatest after the growing season. Small diameter roots at shallow soil depths cycled more rapidly than larger or deeper roots. The strongest treatment effects were found in the most rapidly cycling roots. The standing crop of live roots increased with fertilizer treatment according to both minirhizotron and soil coring methods. However, production and mortality had unique treatment response patterns. Although cumulative mortality decreased in response to increased fertilization, cumulative production was intermediate at 0 kg N ha(-1), lowest with 50 kg N ha (-1), and highest with 200 kg N ha(-1). Aboveground growth responded positively to fertilization up to an application rate of 50 kg N ha(-1), but no further increases in growth were observed despite a threefold increase in application rate. Median fine root life span varied from 307 to over 700 days and increased with depth, diameter and nutrient availability.     

Nitrogen forms affect root structure and water uptake in the hybrid poplar

The study analyses the effects of two different forms of nitrogen fertilisation (nitrate and ammonium) on root structure and water uptake of two hybrid poplar (Populus maximowiczii x P. balsamifera) clones in a field experiment. Water uptake was studied using sap flow gauges on individual proximal roots and coarse root structure was examined by excavating 18 whole-root systems. Finer roots were scanned and analyzed for architecture. Nitrogen forms did not affect coarse-root system development, but had a significant effect on fine-root development. Nitrate-treated trees presented higher fine:coarse root ratios and higher specific root lengths than control or ammonium treated trees. These allocation differences affected the water uptake capacity of the plants as reflected by the higher sapflow rate in the nitrate treatment. The diameter of proximal roots at the tree base predicted well the total root biomass and length. The diameter of smaller lateral roots also predicted the lateral root mass, length, surface area and the number of tips. The effect of nitrogen fertilisation on the fine root structure translated into an effect on the functioning of the fine roots forming a link between form (architecture) and function (water uptake).     

Influence of controlled water supply on shoot and root development of young peach trees

Three controlled water supply treatments were applied to 1-year-old peach trees grown in root observation boxes. The treatments were: I(0), growth medium maintained at 50% field capacity; I(1), water supplied when daily net tree stem diameter change was negative or zero for 1 day; I(3) as for I(1) except that water was applied after net daily stem diameter change was negative or zero for 3 consecutive days. Trees in treatment I(0) had the greatest mean daily first-order shoot growth rates, and trees in treatment I(3) had the lowest shoot growth rates. Because leaf production rate (apparent plastochron) of first-order shoots was unaffected by treatment, differences in shoot length were due to differences in internode extension and not to the number of internodes. Trees in treatment I(0) had a greater number of second-order shoot axes than trees in treatment I(1) or I(3). Furthermore, an increase in the rate of growth of the first-order shoot axis was associated with an increased tendency for branching (i.e., the development of sylleptic second-order shoots). Increased leaf length was also associated with more frequent watering. Trees in treatment I(0) had the greatest root lengths and dry weights, and this was attributed to a greater number of first-and second-order (lateral) root axes compared with trees in the I(1) and I(3) treatments. The extension rate and apical diameter of first-order roots were reduced by the I(3) treatment. The density of second-order roots along primary root axes was not affected by any of the treatments.     

Root distribution of Pinus pinaster, P. radiata, Eucalyptus globulus and E. kochii and associated soil chemistry in agricultural land adjacent to tree lines

We quantified the extent and distribution of roots of four commonly planted tree species (Eucalyptus globulus Labill., Pinus radiata D. Don, P. pinaster Aiton and E. kochii Maiden & Blakely subsp. plenissima C.A. Gardner) in agricultural land adjacent to tree lines, and examined the effect of soil type and root pruning on root morphology. Root distribution in soil adjacent to tree lines was mapped by a trench profile method at 13 sites on the south coast of Western Australia. Soil samples were collected to determine water content and fertility. The lateral extent of tree roots ranged from 10 m for E. kochii to 44 m for P. pinaster. This equated to between 1.5 and 2.5 times tree height (H) for E. globulus and Pinus spp. to 4H for E. kochii. Root density declined logarithmically with distance from the trees and was greatest for P. pinaster and least for E. globulus (P < 0.001). The rate of decrease in root density with distance from the trees was greatest for the Pinus spp. and least for E. kochii (P < 0.05). Root density was generally greatest in the top 0.5 m of the soil profile and decreased with increasing depth. This decrease was relatively gradual in the deep sands, but abrupt in clay subsoil. Root dry mass in the sandy top soil beyond 0.5H ranged between 1.0 and 55.5 Mg km(treeline) (-1) for 6-year-old E. kochii and 50-year-old P. pinaster, respectively. Soil water content generally increased with distance from the trees (P < 0.001). There was no evidence of reduced soil fertility in the top 1.4 m of the soil profile adjacent to the trees. Two to four years after trees had been root pruned, both the lateral extent and vertical distribution of roots were similar for pruned and unpruned trees. The density of roots < 2 mm in diameter was greater for root-pruned trees than for unpruned trees (P < 0.05). We conclude that the study species can compete with agricultural crops based on the lateral extent of their roots and the occurrence of greatest root density within 0.5 m of the soil surface.     

Carbon dioxide efflux from roots of calamodin and apple

The specific rate of CO(2) efflux (respiration) from roots of intact fruiting calamodin plants (Citrus madurensis Lour.) showed no diel trend, and did not respond significantly to short-term (2 day) changes in shoot irradiance. Mean root respiration rate was about 8.4 nmol CO(2) g(-1) s(-1) at 20 degrees C, and increased with temperature with a Q(10) of about 2. In calamodin plants, the proportion of total root length that was white averaged 6.0 mm m(-1). Respiration of roots of apple plants (Malus domestica Borkh.), planted in spring as rootstocks and grown at high irradiance and N supply, declined from about 5.3 to 2.8 nmol CO(2) g(-1) s(-1) between 46 and 138 days after bud burst. At 50% irradiance, root respiration was reduced more than 25% at 46 and 92 days after bud burst, but was not significantly affected later in the season. Reducing shoot irradiance reduced the proportion of total root length that was white, e.g., from 217 to 146 mm m(-1) at 46 days after bud burst. For plants previously grown at low irradiance, increasing shoot irradiance for 6 days increased the rate of root respiration by 5 to 10%. For plants previously grown at high irradiance, reducing shoot irradiance for 6 days reduced root respiration by about 20% early in the season, but had no significant effect later in the season. For plants grown with low-N supply (5% of the high-N), root respiration was reduced early in the season, but was not significantly affected later. Reducing the N supply increased slightly the proportion of total root length that was white. For plants previously grown with low-N, increasing the N supply for 6 days reduced further the rate of root respiration. For plants previously grown with high-N, reducing the N supply for 6 days did not significantly affect the rate of root respiration. Specific respiration rates of root systems excised from mature trees growing outdoors peaked in June, at about 2.4 nmol CO(2) g(-1) s(-1), and then declined for the remainder of the growing season.     

Distribution and tensile strength of Hornbeam (Carpinus betulus) roots growing on slopes of Caspian Forests, Iran

Biomechanical characteristics of the root system of hornbeam (Carpinus betulus) were assessed by measuring Root Area Ratio (RAR) values and tensile strength of root specimens of eight hornbeam trees growing on hilly terrain of Northern Iran. RAR values of the roots were obtained using profile trenching method at soil depth of the top 0.1 m. In total 123 root specimens were analyzed for tensile strength. Results indicate that in general, RAR decreases with depth, following a power function. The RAR values in up and down slopes have no significant statistical differences. In most cases, the maximum RAR values were located in soil depth of the top 0.1 m, with maximum rooting depth at about 0.75 m. The minimum and maximum RAR values along the profiles were 0.004% and 6.431% for down slope and 0.004% and 3.995% for up slope, respectively. The number of roots in the up and down slope trenches was not significantly different. In the same manner as for RAR, number of roots distributing with depth was satisfactorily approximated a power function. The penetration depths of above 90 percent of the roots were at soil depths of 50 cm and 60 cm for up and down slopes, respectively. Results of Spearman's bivariate correlation showed no significant correlation between the RAR value with tree diameter and gradient of slope and number of roots. The mean value of root tensile strength was 31.51 ± 1.05 MPa and root tensile strength decreased with the increase in root diameter, following a power law equation. Using ANCOVA, we found intraspecies variation of tensile strength.