Leaf orientation,light interception and stomatal conductance of Eucalyptus globulus ssp. globulus leaves

Juvenile and adult leaves of the heteroblastic species Eucalyptus globulus Labill. ssp. globulus did not show active diurnal orientation toward or away from incident radiation. Juvenile leaves of a late-maturing sapling of a Tasmanian provenance were evenly distributed in all azimuth sectors. In contrast, an early-maturing sapling of the same age from Wilsons Promontory, Victoria had a predominance of adult leaf blades facing east and west. Mid-vein and blade angles of juvenile and adult leaves were non-random with an overall vertical declination of the leaves. Both leaf types intercepted a greater irradiance during the morning than at midday. Sub-horizontal juvenile leaves intercepted 22% more irradiance than vertical adult leaves during the middle of the day. The amphistomatal and isobilateral adult leaves intercepted sunlight equally on both leaf surfaces. Stomatal conductance was variable during the period of measurement but was similar for the Tasmanian juvenile (0.4 to 0.9 cm s(-1)) leaves and Wilsons Promontory adult (0.5 to 1.2 cm s(-1)) leaves. Greater light interception by the sub-horizontal juvenile leaves would confer a growth advantage to saplings and regenerating canopies. Reduced light interception and leaf temperature of vertical adult E. globulus leaves would assist in water conservation, particularly at high solar angles.     

Mycosphaerella species occurring on Eucalyptus globulus and Eucalyptus nitens plantations of Tasmania,Australia

The genus Mycosphaerella Johanson contains many pathogens capable of causing a severe impact on the growth of susceptible eucalypt species. The lack of knowledge about which species are present in Tasmania and their potential risk to the plantation industry prompted this study into the Mycosphaerella species occurring on Eucalyptus globulus and Eucalyptus nitens plantations in Tasmania. A total of 36 plantation and five road verge sites of E. globulus and E. nitens were sampled. Five Mycosphaerella species and three species from associated anamorph genera were isolated and identified in Tasmania; Mycosphaerella nubilosa, Mycosphaerella cryptica, Mycosphaerella tasmaniensis, Mycosphaerella grandis, Mycosphaerella vespa, Coniothyrium ovatum, Sonderhenia eucalypticola and Sonderhenia eucalyptorum. The most frequently isolated species with the highest incidence and severity of infection were M. cryptica and M. nubilosa. These two species appear to have the greatest potential to damage juvenile eucalypt plantations in Tasmania. A link between Mycosphaerella vespa and Coniothyrium ovatum is described for the first time.     

Root exposure effects on water relations of Eucalyptus nitens nursery stock

Bare-root seedlings of Eucalyptus nitens frequently exhibit water stress after planting resulting in leaf lamina damage and reduced leaf area. Two trials examined effects of root exposure and desiccation between lifting and transplanting on post-planting water relations, leaf retention and root growth. Plants with roots exposed on a glasshouse bench initially lost water rapidly. In one trial ₁ declined to around –2.0 MPa within 2.5 h, after which there was no further change with exposure up to 7.5 h. In the second trial, the initial decline in ₁ was more rapid, reaching below –2.0 MPa in the first hour, before remaining stable with continuing exposure up to 4.5 h. A further decline then continued to –4.0 MPa after 7.5 h.Two days after transplanting into potting mix, day – time leaf water potentials in all desiccation treatments had declined to near –2.0 MPa. Hydraulic resistivity, measured as leaf specific resistivity two days after transplanting, increased following exposure for greater than 2.5 h, but there was no further increase between 4.5 and 7.5 h. The increase in resistivity corresponded with leaf water potential declining below –2.0 MPa during exposure.In the second trial, increasing root exposure time resulted in decreased leaf area due to lamina necrosis. Root growth, measured three weeks after planting, was also reduced. and there was also a positive curvilinear relationship between leaf area remaining at three weeks and new root growth. The results are discussed in terms of hardiness and the management of E. nitens seedlings from nursery to plantation.     

蓝桉子代测定

本文对云南省一平浪蓝桉实生种子园的子代测定林进行初期评价。该子代测定采用α-循环设计,21个重复,每个重复有18个不完全区组,每个区组分为15个小区,每小区2株树。全部270个家系均为自由授粉的半同胞家系,按来源分为3个类型(原产地天然林、云南地方品种和种子园),11个地区。调查和分析的数据包括造林后2年时的树高、胸径、冠幅、干形、抗寒性和存活率等性状。分析结果如下:1. 不同的类型在生长表现上存在显著差异,来自种子园的类型为最好。2. 自不同地区的蓝桉在各性状上都表现出显著差异。在生长表现上,来自维克多利亚东部、西部和各种子园的子代比其它地区的好。云南当地蓝桉子代的耐寒性和两年时的保存率为最好,在生长、干形和抗病方面与塔斯马尼亚东部和南部的蓝桉子代表现相似。采用两株小区,明显比单株小区降低小区缺失率,提高分析的可靠性。表3参7。     

Nondestructive sampling of Eucalyptus globulus and E. nitens for wood properties. I. Basic density

A non-destructive sampling strategy for basic density, based on removing 12 mm bark-to-bark cores, was developed in E. globulus and E. nitens. Fifty trees of each species, aged 5 to 9 years, were sampled across a range of sites. Core samples were removed on both a north-south and an east-west axis from 6 fixed heights in the base of the tree (0.5 m, 0.7 m, …, 1.5 m). Whole-tree values were calculated from disc samples removed at eight percentage heights (0, 10, 20, …, 70%) and correlations between the cores and whole-tree values were used to determine the optimal sampling height. Core samples were found to be reliable predictors of whole-tree density, explaining between 84% and 89% of the variation between trees. Core sampling of E. globulus and E. nitens to estimate basic density of whole-trees and stands is feasible; cores from trees at all E. globulus sites gave high correlations with whole-tree values. For E. nitens, site differences were apparent, and it is recommended that a small destructive sampling program should be undertaken prior to commencing a major sampling program. Recommended optimal sampling heights are 1.1 m for E. globulus and 0.7 m for E. nitens. Core orientation was not important and density was not related to tree size. Six whole-tree samples or eight core samples are required for estimating the mean density of a stand at a specific site to an accuracy of ±20 kg m⁻³ with a 95% confidence interval. Received 17 September 1998     

Leaf growth of Eucalyptus globulus seedlings under water deficit

Eucalyptus globulus Labill. seedlings grown under field conditions in Portugal were watered either daily (control) or every 6 days (drought-treated). Relative to those of control plants, rates of leaf production and leaf biomass accumulation were reduced by almost half in drought-treated plants. However, whereas expansion of new leaves on control plants slowed toward the end of the 30 day experiment, expansion of leaves of the same age on drought-treated plants accelerated as a change in weather conditions resulted in midday plant water potentials above -3.0 MPa. In plants that were left unwatered until they wilted and were then watered daily, expansion of the fifth leaf pair from the apex was slower than that of the same pair of leaves of plants watered daily throughout; but it continued for about twice as long and resulted in the same final leaf area. Drought treatment also caused a substantial reduction in the rate of leaf production, which, in part, accounted for the effect of drought on leaf biomass production. In a greenhouse study, witholding water for 15 days had only a slight effect on the length or width of adaxial epidermal cells, and the effect was quickly reversed on rewatering.     

Nondestructive sampling of Eucalyptus globulus and E. nitens for wood properties; II. Fibre length and coarseness

Within-tree variation in fibre length and coarseness was studied in fifty trees of E. globulus and E. nitens to develop a non-destructive sampling strategy. Trees, aged 5 to 9 years, were sampled across a range of sites in southern Australia. Simulated core samples were removed at six fixed heights easily accessible from the ground (0.5, 0.7, ... 1.5 m) and at eight percentage heights (0, 10, 20, ... 70%). Whole-tree values, calculated from percentage height data, were correlated with the core data to determine the optimal sampling height. Core samples were found to be reliable predictors of whole-tree fibre length, but results were variable for fibre coarseness. Simulated cores taken from the recommended sampling heights explained 87% and 71% of variation in whole-tree fibre length for E. globulus and E. nitens respectively and 54% and 45% of the variation in whole-tree fibre coarseness. Fibre length at all fixed heights showed good correlations with whole-tree values at all sites for E. globulus. For E. nitens the correlations were slightly lower and variable across sites. Results for fibre coarseness varied across sampling heights and sites for both species. The recommended sampling height for fibre length is 1.5 m for both species, whilst for fibre coarseness, the recommended sampling heights are 0.9 and 1.1 m for E. globulus, and 0.9 and 1.3 m for E. nitens. Radial orientation of cores was not important and neither fibre length nor coarseness were related to tree size or basic density. To estimate stand mean fibre length to an accuracy of ±5% would require sampling 9 whole trees or taking cores from 13 trees for E. globulus and 4 whole trees or cores from 8 trees for E. nitens. For estimating stand mean fibre coarseness, 10 whole trees of E. globulus and 7 whole trees are needed for E. nitens. Core sampling for stand mean coarseness would require more trees: 13 to 21 for E. globulus and 11 to 16 trees for E. nitens. Received 17 September 1998     

Physiology and anatomy of lenticel-like structures on leaves of Eucalyptus nitens and Eucalyptus globulus seedlings

Intumescences or abnormal, non-pathogenic, blister-like protuberant growths, form on Eucalyptus globulus Labill. and, to a much lesser extent, Eucalyptus nitens (Deane and Maiden) Maiden leaves when plants are grown in a high relative humidity environment. We examined the histology of intumescences and their effects on leaf photosynthetic processes. Intumescences were induced by placing E. globulus and E. nitens seedlings in a relative humidity of 80% in a greenhouse for 5 days. Symptomatic and asymptomatic leaves of plants with intumescence development were compared with leaves of control plants. Light-saturated carbon dioxide (CO(2)) assimilation (A(max)) and responses of CO(2) assimilation (A) to varying intercellular CO(2) partial pressure (C(i)) were measured. Symptomatic and asymptomatic leaf samples were fixed and sectioned and cellular structure was examined. Intumescences greatly reduced the photosynthetic capacity of E. globulus leaves and were associated with reduced electron transport rate and ribulose bisphosphate (RuBP) regeneration capacity. Tissue necrotization and cellular collapse of the palisade mesophyll and deposition of phenolic compounds in the affected areas, probably reduced light penetration to photosynthesizing cells as well as reducing the amount of photosynthesizing tissue. Photosynthetic capacity of E. nitens was unaffected. The intumescences resembled simple lenticels, both morphologically and developmentally. To our knowledge, this is the first time that lenticel-like structures developed in response to environmental conditions have been described on leaves.     

Growth and water relations of seedlings of two subspecies of Eucalyptus globulus

Seedlings of Eucalyptus globulus Labill subsp. globulus grown in soil in pots in the greenhouse grew faster than seedlings of E. globulus subsp. bicostata, and responded better to added nutrients and water. However, water stress caused a greater reduction in the growth of shoots and roots, and in the root/shoot ratios of fertilized seedlings of subsp. globulus than in those of bicostata. More leaf surface wax was produced by seedlings grown in the presence of fertilizer and an adequate supply of water than by seedlings subjected to nutrient or water stress. Despite larger amounts of leaf surface wax, seedlings of subsp. bicostata had higher epidermal conductances than seedlings of subspecies globulus. However, epidermal conductances were reduced more by water stress and by fertilization in seedlings of subsp. bicostata than in subsp. globulus. Tissue osmotic potentials at full and zero turgor were reduced by water stress only in seedlings of subsp. bicostata and were increased by fertilizer only in seedlings of subsp. globulus. The results indicate that although seedlings of subsp. globulus have inherently higher growth rates, seedlings of subsp. bicostata are better adapted to drought.     

Leaf water relations of Eucalyptus cloeziana and Eucalyptus argophloia in response to water deficit

Leaf water relations responses to limited water supply were determined in 7-month-old plants of a dry inland provenance of Eucalyptus argophloia Blakely and in a humid coastal provenance (Gympie) and a dry inland provenance (Hungry Hills) of Eucalyptus cloeziana F. Muell. Each provenance of E. cloeziana exhibited a lower relative water content at the turgor loss point, a lower apoplastic water content, a smaller ratio of dry mass to turgid mass and a lower bulk modulus of elasticity than the single provenance of E. argophloia. Osmotic potential at full turgor and water potential at the turgor loss point were significantly lower in E. argophloia and the inland provenance of E. cloeziana than in the coastal provenance of E. cloeziana. There was limited osmotic adjustment in response to soil drying in E. cloeziana, but not in E. argophloia. Between-species differences in water relations parameters were larger than those between the E. cloeziana provenances. Both E. cloeziana provenances maintained turgor under moderate water stress through a combination of osmotic and elastic adjustments. Eucalyptus argophloia had more rigid cell walls and reached lower water potentials with less reduction in relative water content than either of the E. cloeziana provenances, thereby enabling it to extract water from dryer soils.     

Influence of light availability on leaf structure and growth of two Eucalyptus globulus ssp. globulus provenances

Light availability strongly affects leaf structure of the distinctive ontogenetic leaf forms of Eucalyptus globulus Labill. ssp. globulus. Late-maturing plants from St. Marys, Tasmania and early maturing plants from Wilsons Promontory, Victoria (hereafter referred to as Wilsons Prom.) were grown for 9 months in 100, 50 or 10% sunlight. Growth, biomass and leaf area were significantly reduced when plants were grown in 10% sunlight. Provenance differences were minimal despite retention of the juvenile leaf form by the Tasmanian plants throughout the study. The time taken for initiation of vegetative phase change by the Wilsons Prom. saplings increased with decreasing light availability, but the nodal position of change on the main stem remained the same. Both juvenile and adult leaves remained horizontal in low light conditions, but became vertical with high irradiance. Leaf dimensions changed with ontogenetic development, but were unaffected by light availability. Juvenile leaves retained a dorsiventral anatomy and adult Wilsons Prom. leaves retained an isobilateral structure despite a tenfold difference in light availability. Stomatal density and distribution showed ontogenetic and treatment differences. At all irradiances, juvenile leaves produced the smallest stomata and adult leaves the largest stomata. Amphistomy decreased with decreasing irradiance. Detrended, correspondence analysis ordination highlighted the structural changes influenced by ontogenetic development and light availability. Adult leaves had characteristics similar to the xeromorphic, sun-leaf type found in arid, high-light conditions. Although juvenile leaves had characteristics typical of mesomorphic leaves, several structural features suggest that these leaves are more sun-adapted than adult leaves.     

Growth, leaf morphology, water use and tissue water relations of Eucalyptus globulus clones in response to water deficit

Changes in leaf size, specific leaf area (SLA), transpiration and tissue water relations were studied in leaves of rooted cuttings of selected clones of Eucalyptus globulus Labill. subjected to well-watered or drought conditions in a greenhouse. Significant differences between clones were found in leaf expansion and transpiration. There was a significant clone x treatment interaction on SLA. Water stress significantly reduced osmotic potential at the turgor loss point (Pi0) and at full turgor (Pi100), and significantly increased relative water content at the turgor loss point and maximum bulk elastic modulus. Differences in tissue water relations between clones were significant only in the mild drought treatment. Among clones in the drought treatments, the highest leaf expansion and the highest increase in transpiration during the experiment were measured in those clones that showed an early and large decrease in Pi0 and Pi100.     

Where to shoot your pilodyn: within tree variation in basic density in plantation Eucalyptus globulus and E. nitens in Tasmania

Longitudinal patterns of within tree variation for basic density were determined for plantation Eucalyptus globulus (ages 5 and 10 years) and E. nitens (ages 5, 10 and 15 years) growing in three geographic areas in Tasmania. Each tree was sampled by taking discs from a combination of percentage heights (0, 10, 20 ___ 70%) and fixed height samples (0.5 m, 0.7 m ___ 1.5 m). At each of the fixed heights, a single pilodyn reading was taken from each of 4 aspects. Weighted whole tree density was calculated from the percentage height samples and used for correlation and regression analysis with the fixed height samples. Both species showed an initial drop in density between the felling cut (zero height) and 0.5 m, followed by a linear increase in density between 10% and 70% of tree height. Slope of fitted regressions were parallel within species but differed significantly between the species (1.14 kg/m³ and 1.62 kg/m³ per 1% increase in height above 10% for E. globulus and E. nitens, respectively). Density at all fixed heights was highly correlated with whole tree values for E. globulus, but results were variable across sites for E. nitens. Both species were found to contain trees which produced aberrant pilodyn readings. For E. globulus, the optimal sampling height was 1.3 m above ground and the mean pilodyn reading was found to predict whole tree density with an accuracy of ±21 kg/m³. For E. nitens, optimal sampling height was 1.5 m above ground. However, pilodyn readings around the stem were not very repeatable and correlations with whole tree density were lower, resulting in the accuracy of prediction of whole tree density being ±26 kg/m³.     

Nondestructive sampling of Eucalyptus globulus and E. nitens for wood properties. III. Predicted pulp yield using Near Infrared Reflectance Analysis

  Within-tree variation in kraft pulp yield, predicted using near infrared reflectance analysis, was studied in thirty trees of E. globulus and fifty trees of E. nitens to develop a non-destructive sampling strategy. Trees, aged 5 to 9 years, were sampled across a range of sites in southern Australia. Simulated core samples were removed at six fixed heights easily accessible from the ground (0.5, 0.7, ... 1.5 m) and at seven percentage heights (0, 20, 30, ... 70%). Whole-tree values, calculated from percentage height data, were correlated with the core data to determine the optimal sampling height. Core samples were found to be good predictors of whole-tree pulp yield for E. globulus, with simulated cores taken from the recommended sampling height (1.1 m) explaining more than 50% of variation in whole-tree pulp yield. Results for E. nitens were variable with large site differences apparent. On high quality sites, core samples from the recommended sampling height (0.9 m) were good predictors of whole-tree pulp yield, explaining around 60% of the variation. On poor quality sites, cores were poor predictors of whole-tree pulp yield. Radial orientation of cores was not important and predicted pulp yield was not related to tree size, basic density or fibre length. To estimate stand mean pulp yield to an accuracy of ±1% would require sampling 6 trees of E. globulus and 4 trees for E. nitens using either multiple discs or core samples. A single sampling height (1.1 m) is recommended for sampling for basic density, fibre length, fibre coarseness and predicted pulp yield in E. globulus. For E. nitens the recommended sampling height for basic density and fibre length is 0.7 m and 0.9 m is recommended for predicted pulp yield on good quality sites. Received 17 September 1998     

Changes in water relations of Eucalyptus nitens nursery stock during and after lifting and transplanting

Field symptoms of transplanting shock in Eucalyptus nitens open rooted (1/2 + 1/2) transplants suggest water stress. To identify stages in the development of symptoms, changes in water relations were monitored from undisturbed seedlings growing in a commercial nursery through lifting, transplanting into pots and re-establishment of normal water relations. Hydraulic resistance measured as leaf specific resistivity increased with each step in the transplanting process. Resistivity after lifting was double that measured on undisturbed plants in the nursery and subsequently doubled with each of the two successive stages, root trimming and planting out. Resistivity fell after the initial rise, returning to the values recorded in the nursery in two stages. An initial rapid decrease, which was not accompanied by any measurable root growth, was followed by a seven day period without a change in resistivity before a further gradual decrease as root volume increased with new root growth.All water relations measurements were taken on trees which had not suffered any permanent leaf damage as a result of transplanting. For a small group of trees which did suffer damage there was a significant relationship between root volume at planting and the retained healthy leaf area. Before shoot growth had recommenced the final ratio of turgid leaf area to initial root volume was comparable with plants immediately after lifting and before trimming. The results are discussed in terms of the changes in predawn leaf water potential, leaf water potential and the leaf damage or defoliation symptoms occuring in plantations.     

Leaf water relations and stomatal behavior of four allopatric Eucalyptus species planted in Mediterranean southwestern Australia

In 1986, four allopatric Eucalyptus species (E. camaldulensis Dehnh, E. saligna Smith, E. leucoxylon F. Muell and E. platypus Hook.) were planted together in a 480-mm rainfall zone, in 8-m wide contour belts as part of a plan to minimize waterlogging and secondary salinization. Throughout 1997, 1998 and 1999, there was significant inter-specific variation in predawn leaf water potential (Psi(pd)); however, maximum stomatal conductance (g(sm)) only differed significantly between species in mid to late summer. Relationships between g(sm) and Psi(pd) were significant and showed that stomata of E. camaldulensis were significantly more sensitive to Psi(pd), and presumably soil water potential, than stomata of E. leucoxylon or E. platypus. When applied to the Psi(pd) data, these relationships predicted that g(sm), and by inference transpiration, varied much less between species than Psi(pd). Diurnal measurements throughout the season confirmed this prediction, and showed that E. camaldulensis and E. saligna avoided drought by gaining access to deeper water, whereas E. leucoxylon and E. platypus maintained greater g(sm) at a given water stress than E. camaldulensis or E. saligna. Osmotic potentials measured after rehydration and water release curves of the leaves indicated that different mechanisms accounted for the apparent drought tolerance of E. leucoxylon and E. platypus. In summer, E. leucoxylon reduced osmotic potential at full and zero turgor by similar amounts compared with winter. In summer, E. platypus had a significantly lower bulk elastic modulus and relative water content at turgor loss point than E. camaldulensis, E. saligna or E. leucoxylon. This elastic adjustment resulted in a larger difference between osmotic potential at full and zero turgor in summer than in winter. The inherently low osmotic potential in E. leucoxylon and elastic adjustment in E. platypus resulted in turgor loss at a similar and significantly lower water potential than in E. camaldulensis or E. saligna. These results have implications for species selection for planting to manage groundwater recharge in areas prone to waterlogging and secondary salinization.     

Water and nutrient supply regimes and the water relations of juvenile leaves of Eucalyptus globulus

Pressure-volume analysis was used to study effects of irrigation and fertilization on the water relations of newly expanded juvenile leaves of Eucalyptus globulus Labill. seedlings growing in Portugal's Atlantic region. In May, at the beginning of the dry season, fertilization and irrigation treatments had no significant effects on the water relations parameters investigated. In September, at the end of the dry season, leaves from non-irrigated plants had a significantly higher apoplasmic water content and a higher dry weight/turgid weight ratio than leaves of similar physiological age from irrigated trees. The osmotic potential at full turgor and the water potential at the wilting point were lower in non-irrigated than in irrigated plants. Changes in osmotic potential at full turgor were negatively correlated with changes in dry weight/turgid weight ratio, suggesting that reductions in osmotic potential at full turgor were largely the result of decreases in cell size. Fertilization had no detectable effect on these variables or on leaf tissue bulk modulus of elasticity. Tissue elasticity was also unaffected by irrigation. Independently of water and nutrient supplies, leaf elasticity was higher and relative water content at the wilting point lower in leaves sampled in May than in leaves sampled in September. In non-irrigated plants, leaves sampled in September had a lower tissue elasticity and a lower osmotic potential at full turgor than leaves sampled in May, indicating that leaves produced at the end of the dry season generate lower water potentials as turgor is lost than leaves expanded early in the season.     

Anatomical and histochemical defence responses induced in juvenile leaves of Eucalyptus globulus and Eucalyptus nitens by Mycosphaerella infection

The responses of juvenile leaves of two Eucalyptus species, with contrasting susceptibility to infection by Mycosphaerella leaf disease, were compared. The anatomical changes, accumulation of phenolics, suberin, lignin and anthocyanin and the retention of chlorophyll were studied in leaf lesions of varying developmental stages caused by species of Mycosphaerella. Enhanced resistance of Eucalyptus nitens in southern Australia was attributed to the formation of an effective lignified and suberized necrophylactic periderm, to restrict pathogen spread. Leaves of E. nitens contained isobilateral palisade which resulted in both abaxial and adaxial cell division and the initiation of a strong reinforced cellular zone from an early lesion stage. Eucalyptus globulus formed a slower, distorted necrophylactic periderm through hypertrophic changes to existing mesophyll and limited cell divisions of the single adaxial palisade layer. Deposits of lignin and suberin did not occur until later in lesion development, which were not effective in preventing further disease development. From this study it is hypothesized that tolerance of eucalypts to Mycosphaerella pathogens may be associated with constitutive mesophyll density.     

Studies on Dual Mycorrhizas of Eucalyptus globulus and E

This paper represents the first part of the results from a glasshouse experiment designed to compare the competition of ECM and VAM fungi on root colonization and effects on growth of two Eucalyptus species　（E．globulus and E．urophylla）．One ECM fungus（Laccaria lateritia）　and three VAM fungi belonging to genera Glomus，Acaulospora and Scutellospora，along with field soil as well，were used to inoculate Eucalyptus seedlings alone or in combination．Both ECM and VAM fungi colonized and formed mycorrhizal associations on roots of inoculated seedlings，though infective rates of VAM and ECM tips per meter varied according to inoculant fungi and plant ages． There were some regressive interactions for colonization between the two fungi，as a general trend for ECM root colonization levels to increase with time at the expense of VAM colonization was observed during 16 weeks． The effect of phosphorus levels in soils on mycorrhizal formation was discussed．