Non-destructive measurement of wheat roots in large undisturbed and repacked clay soil cores

Summary Non-destructive observations of root growth and distribution can be obtained from counting root intercepts with observation tubers inserted in the root zone. This paper describes the technique of inserting clear acrylic tubes horizontally into large undisturbed and repacked soil cores. Counts of roots intersecting scribed lines on the sides of the tubes were made with a fibrescope. Comparison was made between observation root tubes of different diameter (25 and 38.5 mm).The r² values for the relationships between root intercept counts and destructively determined values of root length density (RLD) ranged from 0.78 to 0.96. The larger diameter tubes had higher r² values. Theoretical calibration of the technique does not appear to be possible since analysis indicated that fewer roots intersected the scribed lines on the observation tube than would have been expected from a non-disturbed, randomly distributed root system. It is not known if this discrepancy is due to non-randomness or to an artifact associated with the insertion of the observation tube. Roots were not more prolific at the edge of the soil cores. Comparison of values of root length per unit soil surface area, rates of downward root growth and water uptake rates were within the ranges previously reported for wheat roots of field crops grown on clay soils. Observed root growth and distribution was found to be sensitive to four soil and water treatments imposed. It is concluded that the technique will allow quantitative analysis of root growth and distribution in undisturbed soil cores.     

Automated image analyses for separating plant roots from soil debris elutrated from soil cores

Historically, destructive root sampling has been labor intensive and requires manual separation of extraneous organic debris recovered along with the hydropneumatic elutriation method of separating plant roots from soils. Quantification of root system demographics by public domain National Institute of Health (NIH-Image) and Root Image Processing Laboratory (RIPL) image processing algorithms has eliminated much of the labor-intensive manual separation. This was accomplished by determining the best length to diameter ratio for each object during image analyses. Objects with a length to diameter ratio less than a given threshold are considered non-root materials and are rejected automatically by computer algorithms. Iterative analyses of length to diameter ratios showed that a 15:1 ratio was best for separating images of maize (Zea mays L.) roots from associated organic debris. Using this threshold ratio for a set of 24 soil cores, a highly significant correlation (r² = 0.89) was obtained between computer image processed total root length per core and actual root length. A linear relationship (r² = 0.80) was observed between root lengths determined by NIH-Image analyses and lengths determined independently by the RIPL imaging system, using the same maize root + debris samples. This correlation demonstrates that computer image processing provides opportunities for comparing root length parameters between different laboratories for samples containing debris.     

Most probable numbers of Azospirillum associated with the roots of inoculated pearl millet

Summary Inoculation of pearl millet (Pennisetum americanum (L.) Leeke) with Azospirillum significantly increased the numbers of this organism in the rhizosphere, rhizoplane, washed and crushed roots and surface sterilized and crushed roots. The maximum number of organisms plant^–1 were localized in the rhizosphere. The numbers of Azospirillum on the roots of inoculated plants grown under sterilized conditions were much higher than in the field grown plants. In both cases populations outside the roots were higher than in the surface sterilized roots. The highest numbers per unit root weight were recorded between 60–75 days of growth. N₂-ase activity throughout the growth cycle was very low and was not related to the populations of Azospirillum on the roots. Root exudates and extracts of pearl millet showed a stimulatory effect on the growth of Azospirillum suggesting their possible involvement in the colonization of this organism on the roots of inoculated plants.     

Use of the ureide technique to describe the nitrogen economy of field-grown soybeans

Nitrogen fixation was estimated in `Bragg,' `Forrest,' and `Bethel' soybean (Glycine max [L.] Merrill) from seven locations northwest of New South Wales, Australia, by relating ureide and nitrate contents of plant parts sampled at regular intervals during growth to standard curves derived under controlled nitrate regimes. Estimates were combined with data on crop growth and mineral N contents of soils to (a) determine the total requirements for N by the crops, (b) determine the contributions of N₂ fixation to crop growth, and (c) relate symbiotic dependence ([N₂ fixed/total plant N] × 100) of the crops to levels of mineral N in the soil at sowing. At two locations, Myall Vale and Glenara, levels of ureides in the shoot axes and roots of unnodulated seedlings were surprisingly high at the first time of sampling, perhaps reflecting effects of uptake of ammonium-N by the soybeans or breakdown and remobilization of cotyledonary protein. Ureides in plant parts declined significantly by the second (V5 to V7 growth stage) sampling. Subsequently, ureide contents increased whereas levels of nitrate in plant parts decreased. The relative abundance of ureides ([ureide-N/ureide-N + nitrate-N] × 100) in the shoot axes and nodulated roots of both crops increased linearly from almost zero during mid-vegetative growth (V5 to V7) to virtually 100% during late reproductive growth (R4 to R5, Myall Vale and R6, Glenara). The data suggest a steady transition in soybeans at both locations from dependence upon mineral N for early growth to complete reliance upon fixed N during late reproductive growth. Estimates of seasonal N₂ fixation for soybeans at the seven locations ranged from 73 to 288 kilograms per hectare N (shoot axes ureides) and from 147 to 337 kilograms per hectare N ha (nodulated roots ureides).     

Soil biota responses to long-term atmospheric CO2 enrichment in two California annual grasslands

Root, arbuscular-mycorrhizal (AM), soil faunal (protozoa and microarthropods), and microbial responses to field exposure to CO₂ for six growing seasons were measured in spring 1997 in two adjacent grassland communities. The grasslands showed contrasting root responses to CO₂ enrichment: whereas root length was not affected in the sandstone grassland, it was greater in the serpentine grassland, as was specific root length. AM fungal hyphal lengths were greater in the sandstone, but were unaffected in the serpentine community. This lent support to the hypothesis that there may be a tradeoff in resource allocation to more fine roots or greater mycorrhizal extraradical hyphal length. AM root infection was greater in both communities at elevated CO₂, as was the proportion of roots containing arbuscules. Our data on total hyphal lengths, culturable and active fungi, bacteria, and protozoa supported the hypothesis that the fungal food chain was more strongly stimulated than the bacterial chain. This study is one of the first to test these hypotheses in natural multi-species communities in the field. Received: 23 October 1998 / Accepted: 19 February 1999     

Estimation of tree root lengths using fractal branching rules: a comparison with soil coring for Grevillea robusta

Previous theoretical research has suggested that lengths of tree roots can be estimated on the basis of their branching characteristics, if branching has a fractal pattern that is independent of root diameter. This theory and its underlying assumptions was tested for Grevillea robusta trees at a site in Kenya by comparing estimates of root length from conventional soil coring and the output of a fractal branching algorithm. The trees were in a 4-year-old stand established on a 3 × 4 m planting grid. Root lengths (L _r) in four units of the planting grid were estimated by soil coring. Branching characteristics determined by examination of 32 excavated roots from 16 trees were: The number of branches at each branching point; the length of links between branching points (L _l); the diameter of root tips; and parameters which describe the change in diameter at each branching point. Each was found to be independent of root size. These data were used to parameterise a branching algorithm, which was then used to estimate numbers of root links in the four grid units (n _l) from root diameters at the bases of the four trees at the corners of each unit. Root lengths, from L _r = n₁ L₁, severely underestimated L _r. This discrepancy probably resulted from inaccuracy in the parameterisation of the branching algorithm, as output from the algorithm was very sensitive to small changes in parameter values. Use of fractal branching rules alone to estimate roots length does not appear possible unless the algorithm is calibrated to adjust for errors in parameter estimation. Calibration can be achieved by calculation of an 'effective link length', L ^eff ₁, from L _r/n _l, where L _r is measured by a reference method such as soil coring.     

Influence of rocks on soil temperature,soil water potential,and rooting patterns for desert succulents

Summary At a site in the Sonoran Desert, subterranean rocks and exposed boulders affected soil water potential as well as root morphology and distribution. For Agave deserti, the number of lateral roots per unit length of main root was 11 times higher under rocks and six times higher alongside rocks than in rock-free regions. Total root length per unit soil volume for Echinocereus engelmannii averaged 3-fold higher within 1 cm of boulders than 5 cm away, where the soil was drier. The total length of lateral roots per unit length of main root for Ferocactus acanthodes was 4.2 m m^–1 under rocks but only 0.8 m m^–1 in rock-free regions. The number of lateral roots per unit length of main root for Opuntia acanthocarpa was 7-fold higher alongside rocks than in rock-free regions and even higher under rocks. For transplanted and watered A. deserti, the number of new main roots produced per 1–2 month interval averaged 13 for five plants on the north side of boulders, 8 on the south side, 11 for five plants with half of their roots under rocks, 2 for those with half of their roots over rocks, and 3 for the control plants without rocks. Laboratory experiments showed that the soil water potential under rocks for 10 and 30 mm waterings stayed above –0.5 MPa for 13 and 19 d longer, respectively, than for regions away from rocks. The shortwave absorptance of granitic rocks from the field site was 0.82, the thermal conductivity coefficient was 1.50 W m^–1 °C^–1, and the volumetric heat capacity was 1.75 MJ m^–3 °C^–1. Field measurements indicated that 5-cm-thick buried rocks decreased the diel variation in soil temperatures on their undersurface by only 0.4° C compared with soil. Thus, the primary influence of rocks at the field site on root proliferation and branching for the four species was apparently caused by influences on soil water content.     

The growth,activity and distribution of the fruit tree root system

Summary The paper reviews information, much of it obtained from studies using the East Malling root observation laboratories, on the growth and development of the fruit tree root system. The production of new white root varies from year-to-year, generally being highest in the early years. As trees age, woody roots constitute an increasing fraction of total root length although the contribution made by new root growth to the total root length of established trees is also affected by soil management, being higher for trees under grass than under herbicide. Soil management also affects the balance of short (lateral) to long (extension) roots; under grass there are more lateral roots.Calculation of the rate of water uptake per unit root length needed at various times in the year to meet transpirational demand, suggests that woody roots, which recent experimental work has shown to be capable of absorbing water, must be responsible for much of total water supply.Measurements of VA mycorrhizal infection in field-grown trees indicated, for part of the season, higher per cent infection in trees grown under irrigated grass than under herbicide management. It is suggested that this, which is associated with raised leaf phosphorus levels, may be due at least partly to higher numbers of lateral roots, the root type which becomes infected. The growth and functioning of the root system under field conditions depend upon the production and integration of a range of root types.     

基于土芯法的亚热带常绿阔叶林细根空间变异与取样数量估计

树木细根具有高度空间异质性,确定合理的细根取样策略是林木细根研究的前提。通过在福建省三明米槠天然常绿阔叶林内随机钻取96个土芯,分析细根生物量和形态特征的空间变异特征,并估计各指标所需的取样数量。结果表明:(1)随着径级增加,细根各指标变异系数增大,相应的取样数量增加;(2)随着土壤深度增加,单位面积细根生物量变异程度和相应的取样数量均增加。在置信水平为95%、精度为80%的条件下,直径为0-1 mm和1-2 mm的细根,分别采集16和42个样品可以满足测定单位面积细根生物量,采集17和31个样品可以满足测定单位面积细根长度,采集25和33个样品可以满足测定单位面积细根表面积。Shapiro-Wilk检验表明,除表层土壤0-1 mm细根单位面积生物量符合正态分布外,其余细根生物量和形态指标数据均不符合正态分布。研究结果为亚热带常绿阔叶林细根的合理取样提供了科学依据。     

Root growth as influenced by aggregate size

Summary This study examined the effects of aggregate size on root impedance and developed an equation to describe the root pressure necessary to avoid deflection around an aggregate. This critical root pressure was predicted to increase with increasing aggregate size, decreasing root diameter, and decreasing deflection angle. In growth chamber experiments, maize (Zea mays L.) seedlings were grown in A horizon material of Groseclose silt loam (Clayey, mixed, mesic, Typic Hapludult). The soil had been moist sieved into different aggregate sizes (0–1, 1–2, 2–3, and 3–6 mm diameter). The larger aggregates did constitute a slight root impedance as roots were deflected around them. Diameters of roots grown in 3–6 mm aggregates increased significantly, whereas root lengths were not always signficantly decreased. The smaller aggregates did not impede root growth and were readily displaced by roots. Large aggregates were more of an impedance to lateral roots than to main axes.     

Effects of liming and mycorrhizal colonization on soil phosphate depletion and phosphate uptake by maize (Zea mays L.) and soybean (Glycine max L.) grown in two tropical acid soils

The effects of liming and inoculation with the arbuscular mycorrhizal fungus, Glomus intraradices Schenck and Smith on the uptake of phosphate (P) by maize (Zea mays L.) and soybean (Glycine max [L.] Merr.) and on depletion of inorganic phosphate fractions in rhizosphere soil (Al-P, Fe-P, and Ca-P) were studied in flat plastic containers using two acid soils, an Oxisol and an Ultisol, from Indonesia. The bulk soil pH was adjusted in both soils to 4.7, 5.6, and 6.4 by liming with different amounts of CaCO₃.In both soils, liming increased shoot dry weight, total root length, and mycorrhizal colonization of roots in the two plant species. Mycorrhizal inoculation significantly increased root dry weight in some cases, but much more markedly increased shoot dry weight and P concentration in shoot and roots, and also the calculated P uptake per unit root length. In the rhizosphere soil of mycorrhizal and non-mycorrhizal plants, the depletion of Al-P, Fe-P, and Ca-P depended in some cases on the soil pH. At all pH levels, the extent of P depletion in the rhizosphere soil was greater in mycorrhizal than in non-mycorrhizal plants. Despite these quantitative differences in exploitation of soil P, mycorrhizal roots used the same inorganic P sources as non-mycorrhizal roots. These results do not suggest that mycorrhizal roots have specific properties for P solubilization. Rather, the efficient P uptake from soil solution by the roots determines the effectiveness of the use of the different soil P sources. The results indicate also that both liming and mycorrhizal colonization are important for enhancing P uptake and plant growth in tropical acid soils.     

A comparison of penetrometer pressures and the pressures exerted by roots

Summary Previous work is reviewed in which the ratio of the pressures required for soil penetration by roots and penetrometers are compared. It appears that this ratio can vary from about 2 to 8 depending on conditions. However, there is very little experimental evidence and most of the work has been inferential.Direct measurements are reported for the stresses exerted by a 1 mm diameter penetrometer probe and by the roots of pea seedlings when penetrating Urrbrae fine sandy loam. Six soil conditions were used: (non-weathered remoulded soil cores + artificially weathered remoulded soil cores + undisturbed field clods) × (confined + unconfined cores or clods). The confinement treatment was to test for any effects of additional restraint to cylindrical root expansion. The weathering and field clod treatments were to test the hypothesis that root elongation is facilitated by tensile failure ahead of the root tip.The principal conclusions are as follows. The laboratory weathering treatment reduced the soil tensile strength by 25%. This resulted in a small but significant reduction in the pressure for root penetration into confined cores. Compared with remoulded non-weathered cores, field clods had a 2 to 3 fold greater penetrometer resistance and a 50% lower tensile strength. The force required for root penetration into unconfined field clods was only 10% greater than for unconfined non-weathered cores. For the former (which is closest to field conditions) the penetrometer had to exert a pressure 5.1 times greater than a root tip in order to penetrate the soil. Penetrometer penetration pressure was independent of probe diameter in the 1–2 mm range in the soil used. Core confinement restricts root radial expansion and modifies the penetration force of metal probes and plant roots.On the basis of the new results it is tentatively concluded that soil tensile failure can facilitate penetration by roots.     

Mapping quantitative trait loci associated with root penetration ability in rice (Oryza sativa L.)

Root penetration ability is an important factor for rice drought resistance in areas with soils subject to both compaction and periodic water deficits. However, breeding for root penetration ability is inhibited by the difficulties associated with measuring root traits. Our objective was to identify restriction fragment length polymorphisms (RFLPs) associated with root penetration ability. Using wax-petrolatum layers as a proxy for compacted soil, we counted the number of vertical root axes penetrating through the layer, the total number of vertical root axes and the number of tillers per plant of 202 recombinant inbred (RI) lines over three replications. As a measure of root penetration ability, we used a root penetration index defined as the percent of the total number of vertical root axes that penetrated through a wax-petrolatum layer. The RI population exhibited a wide range in the number of penetrating roots axes (10–115 roots), the total number of roots axes (74–226 roots), tillers per plant (6–18), and in the root penetration index (0.11–0.71). Single-marker and interval quantitative trait analyses were conducted to identify RFLP loci associated with the number of penetrating roots, total root number, root penetration index, and tiller number. Four quantitative trait loci (QTLs) were associated with the number of penetrated roots, 19 with the total root number, six QTLs with the root penetration index and ten with tiller number. Individually, these QTLs accounted for a maximum of 8% of the variation in the number of penetrating roots, 19% of the variation in total root number, 13% of the variation in root penetration index and 14% of the variation in tiller number as estimated from regressions. The multimarker regression model accounting for the greatest proportion of the variation in the root penetration index was a three-marker model that accounted for 34% of the variation. Two-marker models accounted for 13% of the variation in the number of penetrated roots, 25% of the variation in total root number, and 21% of the variation in tiller number. This is the first research paper to apply RFLP quantitative trait analysis to dissect genetic loci associated with the total number of roots, root penetration ability and tiller number.Contribution from the Department of Plant and Soil Science, College of Agricultural Sciences and Natural Resources, Texas Tech University Lubbock, TX 79409, USA. Journal Number T-4-385     

Using a fine root extraction device to quantify small diameter corn roots (≥0.025 mm) in field soils

The study of fine roots growing under field conditions is limited by the techniques currently available for separating these roots from soil. This study had two objectives: to measure the total root length of field grown corn (Zea mays L.) by root diameter class, and to develop an inexpensive and efficient root washing device that would effectively capture all of the roots in a field soil sample. An inexpensive Fine Root Extraction Device (FRED) was constructed from readily available materials and was successful at extracting all roots, including very fine diameter roots (0.025 mm), from field soil samples. Greater than 99.7% of marked roots introduced to the FRED were recaptured by the device. Soil samples from three depths, and on three dates, from field grown corn were placed in the FRED. We found that more than 56% of total root length occurred in roots whose diameters were smaller than 0.175 mm, and more than 35% of root length occurred in roots smaller than 0.125 mm in diameter. Corn roots of the diameters described here have not been reported in field soils prior to this study. Root researchers who fail to measure these very fine roots will significantly underestimate root length density. Widespread use of the FRED should improve our understanding of root distribution in field soils.     

A comparison between minirhizotron and monolith sampling methods for measuring root growth of maize (Zea mays L.)

Transparent plastic minirhizotron tubes have been used to evaluate spatial and temporal growth activities of plant root systems. Root number was estimated from video recordings of roots intersecting minirhizotron tubes and of washed roots extracted from monoliths of the same soil profiles at the physiological maturity stage of a maize (Zea mays L.) crop. Root length was measured by the line intercept (LI) and computer image processing (CIP) methods from the monolith samples.There was a slight significant correlation (r=0.28, p<0.005) between the number of roots measured by minirhizotron and root lengths measured by the LI method, however, no correlation was found with the CIP method. Using a single regression line, root number was underestimated by the minirhizotron method at depths between 0–7.6 cm. A correlation was found between root length estimated by LI and CIP. The slope of estimated RLD was significant with depth for these two methods. Root length density (RLD) measured by CIP showed a more erratic decline with distance from the plant row and soil surface than the LI method.     

Variation in phosphorus uptake efficiency by genotypes of cowpea (Vigna unguiculata) due to differences in root and root hair length and induced rhizosphere processes

The lengths of roots and root hairs and the extent of root-induced processes affect phosphorus (P) uptake efficiency by plants. To assess the influence of variation in the lengths of roots and root hairs and rhizosphere processes on the efficiency of soil phosphorus (P) uptake, a pot experiment with a low-P soil and eight selected genotypes of cowpea (Vigna unguiculata (L) WALP) was conducted. Root length, root diameter and root hair length were measured to estimate the soil volume exploited by roots and root hairs. The total soil P was considered as a pool of Olsen-P, extractable with 0.5 M NaHCO₃ at pH 8.5, and a pool of non-Olsen-P. Model calculations were made to estimate P uptake originated from Olsen-P in the root hair zone and the Olsen-P moving by diffusion into the root hair cylinder and non-Olsen-P uptake. The mean uptake rate of P and the mean rate of non-Olsen-P depletion were also estimated. The genotypes differed significantly in lengths of roots and root hairs, and in P uptake, P uptake rates and growth. From 6 to 85% of total P uptake in the soil volume exploited by roots and root hairs was absorbed from the pool of non-Olsen-P. This indicates a considerable activity of root-induced rhizosphere processes. Hence the large differences show that traits for more P uptake-efficient plants exist in the tested cowpea genotypes. This opens the possibility to breed for more P uptake-efficient varieties as a way to bring more sparingly soluble soil P into cycling in crop production and obtain capitalisation of soil P reserves.     

INFLUENCE OF SUPPLEMENTAL INORGANIC NUTRIENTS ON GROWTH,SURVIVORSHIP, AND MYCORRHIZAL RELATIONSHIPS OF SCHIZACHYRIUM SCOPARIUM (POACEAE) GROWN IN FUMIGATED AND UNFUMIGATED SOIL

Little bluestem grass Schizachyrium scoparium ([Michx.] Nash) plants were grown under field conditions for 2 years in soils fumigated with methyl bromide and chloropicrin, or in unfumigated soil, and treated with supplemental inorganic nutrients (bases calcium and magnesium) phosphorus, nitrogen, and potassium. Most differences in measured plant responses were due to interactions between fumigation and nutrient treatments. These included biomass production, root mass per unit length (μg/cm), root lengths, flowering culm production, percent colonization, colonized root length, and spore production in rhizosphere soil. Plants generally responded to mycorrhizal fungal colonization by reducing total root length and producing thicker roots. Treatment of plants with bases appeared to profoundly affect the mycorrhizal association by reducing sporulation of vesicular-arbuscular mycorrhizal fungi and increasing colonization. When fumigated or unfumigated soils were considered separately, base-treated plants produced more biomass than other treatments. Base-treated plants grown on unfumigated soil had more flowering culms and longer colonized root lengths than all other plants. Percent colonization by mycorrhizal fungi and colonized root length were positively correlated with phosphorus/nitrogen ratios, but the ratio was not correlated with plant biomass production. This suggests that phosphorus is not a limiting nutrient in our soil and investment in a mycorrhizal association may not result in enhanced plant growth. The base-nutrient effects may indicate a need to reevaluate earlier studies of macro nutrient effects that did not take into account the role played by calcium and magnesium in assessing fungus-host plant interactions.     

落叶松人工林细根动态与土壤资源有效性关系研究

树木细根在森林生态系统C和养分循环中具有重要的作用。由于温带土壤资源有效性具有明显的季节变化, 导致细根生物量、根长密度 (Rootlengthdensity, RLD) 和比根长 (Specificrootlength, SRL) 的季节性变化。以 17年生落叶松 (Larixgmelini) 人工林为研究对象, 采用根钻法从 5月到 10月连续取样, 研究了不同土层细根 (直径≤ 2mm) 生物量、RLD和SRL的季节动态, 以及这些根系指标动态与土壤水分、温度和N有效性的关系。结果表明 :1) 落叶松细根年平均生物量 (活根 +死根 ) 为 189.1g·m-2 ·a-1, 其中 5 0 %分布在表层 (0~ 10cm), 33%分布在亚表层 (11~ 2 0cm), 17%分布在底层 (2 1~ 30cm) 。活根和死根生物量在 5~ 7月以及 9月较高, 8月和 10月较低。从春季 (5月 ) 到秋季 (10月 ), 随着活细根生物量的减少, 死细根生物量增加 ;2 ) 土壤表层 (0~ 10cm) 具有较高的RLD和SRL, 而底层 (2 1~ 30cm) 最低。春季 (5月 ) 总RLD和SRL最高, 分别为 10 6 2 1.4 5m·m-3 和 14.83m·g-1, 到秋季 (9月 ) 树木生长结束后达到最低值, 分别为 2 198.2 0m·m-3 和 3.77m·g-1;3) 细根生物量、RLD和SRL与土壤水分、温度和有效N存在不同程度的相关性。从单因子分析来看, 土壤水分和有效N对细根的影响明显大于温度, 对活根的影响大于死根。由于土壤资源有效性的季节变化, 使得C的地下分配格局发生改变。各土层细根与有效性资源之间的相关性反映了细根功能季节性差异。细根 (生物量、RLD和SRL) 的季节动态 (5 8%~ 73%的变异 ) 主要由土壤资源有效性的季节变化引起。     

The effect of soil strength on the growth of pigeonpea radicles and seedlings

The effect of soil strength on the growth of pigeonpea radicles and seedlings was investigated in cores of three clay soils prepared at different water contents and bulk densities in the laboratory.Radicle elongation directly into soil cores was reduced from 50–70 mm d^-1 at strengths less than 0.5 MPa to 0 mm d^-1 at 3.5–3.7 MPa. The response to soil strength was affected by the water content of the soil, presumably as a result of reduced oxygen availability in wetter soil. This effect was apparent in soils wet to air-filled porosities less than 0.15 m³ m^-3.Radicles were more sensitive to high soil strength (>1.5 MPa) than were seedling roots which encountered the same conditions at 60 mm in the profile. Radicle growth ceased at 3.5 MPa which reduced seedling root growth by only 60%.Despite a 60% reduction in root length in the high strength zone, seedling roots compensated in zones of loose soil above and below the compacted layer, and total root length and shoot growth were unaffected. There was no evidence of a root signal response which results in reduced shoot growth in some species in response to high soil strength.The proliferation of roots in surface layers and the delayed penetration of the root system to depth in compacted soil are likely to expose seedlings to a greater risk of water-deficit in the field, particularly under dryland conditions where plants rely on stored subsoil water for growth.     

Mycorrhizas and C and N transformations in the rhizospheres of Pinus sylvestris,Picea abies and Betula pendula seedlings

Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies L.) and silver birch (Betula pendula L.) seedlings were grown in a greenhouse for four months in three different soils. The soils were from a field afforestation site on former agricultural land: soil from a pine site, soil from a spruce site and soil from a birch site. Pots without seedlings were included. The aim was to discover, independent of the effects of the different quality of aboveground litter and microclimate under the tree species, whether the roots change the microbial activities and chemical characteristics of the soil, whether the changes are dependent on the tree species, and whether the changes vary in different soils. Pine, spruce and birch had, on average, five, one and six meters of roots, respectively. Birch had by far the highest number of short root tips, on average 11 450 per seedling, compared to 1900 and 450 in pine and spruce seedlings, respectively. The majority of the short roots of pine and spruce were brown sheathed mycorrhizas, and those of birch were mycorrhizas in an early stage of development. The seedlings caused no major changes in either the soil pH or the concentrations of nutrients in the soils, but did affect the microbial characteristics of the soils. The effect of the tree species did not differ in different soils. Microbial biomass C and N, C mineralization rate and the concentration of ergosterol were all higher under birch and pine than under spruce and in plantless soils. Nitrate concentrations were lowest under pine and birch, but rates of net N mineralization, nitrification and denitrification did not differ under different seedlings. The stimulative effect of pine and especially birch on soil microbes was possibly due to them having more roots and releasing more root exudates to soil. There were, however, indications that not only the length/mass of roots determined the changes in microbial activities, but also differences in root activities per unit of root or in the quality of root exudates.