Influence of ammonium on fine root development and rhizosphere pH of Douglas-fir seedlings in sand

Ammonium sulphate is a major component of the air pollutants deposited on forests in the Netherlands. Different amounts of NH₄ ⁺ were added to Douglas-fir seedlings grown in tall containers of sand, to study the influence of high concentrations of NH₄ ⁺ in the soil on the development of fine roots and the effects of nitrogen uptake on rhizosphere pH. At the end of this eight-month experiment part of the ammonium appeared to have nitrified into nitrate. High doses of ammonium negatively affected root length and root length per unit of dry matter (specific root length). Although Douglas fir shows a preferential ammonium uptake in nutrient solutions the increases in the pH of the rhizosphere in this experiment indicate that nitrogen was mostly taken up as nitrate. When the ammonium concentration in the soil is low, it cannot be taken up readily because of its low mobility in soil. Shoot growth was stimulated by high availability of nitrogen. The possible effects of high doses of ammonium on long-term forest vitality are discussed.     

Effects of ammonium sulphate application on the chemistry of bulk soil,rhizosphere, fine roots and fine-root distribution in a Picea abies (L.) karst. stand

The effect of ammonium sulphate application on the bulk and rhizosphere soil chemistry, elemental concentration of living fine roots (<2 mm in diameter), amounts of living and dead fine roots, root length density and specific root length density were investigated in a 28 year old Norway spruce stand in SW Sweden. The treatments started in 1988. Core samples of the LFH layer and mineral soil layers were sampled in control (C) and ammonium sulphate (NS) treatment plots in 1988, 1989 and 1990. Soil pH and NO₃-S and SO₄-S, Al, Ca, Mg, Mn and K concentrations were measured for both the bulk soil and rhizosphere soil.The pH-values of the bulk and rhizosphere soil decreased in 1989 and 1990 in NS plots compared to control plots, while the SO₄-S concentration increased. The Ca, Mg and K concentration increased in the NS treatment in almost all layers in the bulk and the rhizosphere soil. Ammonium ions may have replaced these elements in the soil organic matter. The NS treatment reduced Mg concentration in fine roots in all layers in 1990. The Al concentrations in the rhizosphere and bulk soil were higher in NS plots in all layers, except at 0–10 cm depth, both in 1989 and 1990. The Al content of living fine roots was higher in NS plots than C plots but the differences were not significant. The NS addition did not affect the P and K contents of fine roots in any soil layer, but the S concentrations of fine roots were significantly higher in NS plots in 1989 and 1990. The fine root necromass was higher in NS than in C in 1990, in the LFH layer, indicating a gradual decrease in the vitality of the fine roots. It was suggested that the NS treatment resulted in displacement of Mg and K from exchange sites in the LFH layer leading to leaching of these cations to the mineral soil. Further application of ammonium sulphate may damage the fine roots and consequently adversely affect the water and nutrient uptake of root systems.     

The fate of carbon and fertiliser nitrogen when dryland wheat is grown in monoliths of duplex soil

Triticum aestivum L. (cv. Gutha), a short-season wheat, was grown to maturity in large monoliths of duplex soil (sand over sandy-clay) in a daylight phytotron mimicking field conditions. Either ¹⁵N-labelled ammonium sulphate ((NH₄)₂SO₄) or urea was banded into the soil at a rate of 30 kg N ha^–1: even though roots were about 20% heavier when grown in the presence of (NH₄)₂SO₄ for 86 d (P<0.05), above-ground mass was not affected by the source of nitrogen. At four times through crop development up to grain-filling (50, 56, 70 and 86 d after sowing) shoots were labelled heavily with ¹⁴CO₂ with two purposes. First, to trace `instantaneous' assimilate movement over 24 h, revealing relative sink strengths throughout plants. This, in turn, allowed precise measurements of live root mass and the proportion of recent photoassimilates deposited in the rhizosphere. Although root systems were sparse, even in surface soil layers, they were strong sinks for photoassimilates early in development (0–50 d), supporting the conversion of inorganic applied nitrogen (N) to soil organic forms. In the presence of roots, up to 28% of ¹⁵N was immobilised, whereas only 12% of labelled ammonium sulphate was immobilised in unplanted plots in spite of a favourable moisture status in both treatments. The effect of plants on rates of ¹⁵N transformation is ascribed to recently imported photoassimilates sustaining rhizosphere metabolism. Not more than 15% of recently fixed carbon imported by roots was recovered from the rhizoplane, suggesting that a highly localised microbial biomass supported vigorous immobilisation of soil N. Thus, more than twice as much applied N was destined for soil organic fractions as for root material. By these processes, root- and soil-immobilised N become substantial stores of applied N and together with shoot N accounted for all the applied N under dryland conditions.     

Performance of two Picea abies (L.) Karst. stands at different stages of decline

Summary The development of root tips and apparent ectomycorrhizas was compared in the Fichtelgebirge (FRG) over one growing season in two 30-year-old Picea abies stands, both on soils derived from phyllite but showing varying symptoms of decline. Visual symptoms of tree decline reflected a lower relative and absolute mycorrhizal frequency, a lower number of ectomycorrhizas per m² leaf area and an uneven vertical distribution of root tips and ectomycorrhizas. The number of apparent ectomycorrhizas per ground area was correlated with the amount of magnesium, calcium, and ammonium, and the pH in the free-drainage soil solution, and with the molar calcium to aluminium ratio in mineral soil extracts. The foliage concentrations of magnesium and calcium were correlated with the numbers of apparent ectomycorrhizas per m² leaf or ground area. These observations were used to formulate testable hypotheses concerning the role of the root system and the soil environment in forest decline.     

Bulk soil pH and rhizosphere pH of peach trees in calcareous and alkaline soils as affected by the form of nitrogen fertilizers

One-year old nectarine trees [Prunus persica, Batsch var. nectarina (Ait.) Maxim.], cv Nectaross grafted on P.S.B2 peach seedlings [Prunus persica (L.) Batsch] were grown for five months in 4-litre pots filled with two alkaline soils, one of which was also calcareous. Soils were regularly subjected to fertigation with either ammonium sulphate or calcium nitrate providing a total of 550 mg N/tree. Trees were also grown in such soils receiving only deionized water, as controls. Rhizosphere pH, measured by the use of a microelectrode inserted in agar sheet containing a bromocresol purple as pH indicator and placed on selected roots, was decreased by about 2–3 units compared to the bulk soil pH in all treatments. This decrease was slightly less marked when plants were supplied with calcium nitrate rather than ammonium sulphate or control. Measurements conducted during the course of the experiment indicated that ammonium concentration was similar in the solution of soils receiving the two N fertilizers. During the experiment, soil solution nitrate-N averaged 115 mg L^–1 in soil fertilized with calcium nitrate, 68 mg L^–1 in those receiving ammonium sulphate and 1 mg L^–1 in control soils. At the end of the experiment nitrate concentrations were similar in soils receiving the two N sources and bulk soil pH was decreased by about 0.4 units by ammonium sulphate fertigation: these evidences suggest a rapid soil nitriflcation activity of added ammonium. Symptoms of interveinal chlorosis in apical leaves appeared during the course of the experiment in trees planted in the alkaline-calcareous soil when calcium nitrate was added. The slightly higher rhizosphere pH for calcium nitrate-fed plants may have contributed to this. The findings suggest that using ammonium sulphate in a liquid form (e.g. by fertigation) in high-pH soils leads to their acidification and the micronutrient availability may be improved.     

Ammonia-oxidizing bacteria on root biofilms and their possible contribution to N use efficiency of different rice cultivars

Ammonia-oxidizing bacteria (AOB) populations were studied on the root surface of different rice cultivars by PCR coupled with denaturing gradient gel electrophoresis (DGGE) and fluorescence in situ hybridization (FISH). PCR-DGGE of the ammonium monooxygenase gene (amoA) showed a generally greater diversity on root samples compared to rhizosphere and unplanted soil. Sequences affiliated with Nitrosomonas spp. tended to be associated with modern rice hybrid lines. Root-associated AOB observed by FISH were found within a discrete biofilm coating the root surface. Although the total abundance of AOB on root biofilms of different rice cultivars did not differ significantly, there were marked contrasts in their population structure, indicating selection of Nitrosomonas spp. on roots of a hybrid cultivar. Observations by FISH on the total bacterial community also suggested that different rice cultivars support different bacterial populations even under identical environmental conditions. The presence of active AOB in the root environment predicts that a significant proportion of the N taken up by certain rice cultivars is in the form of NO₃ ^–-N produced by the AOB. Measurement of plant growth of hydroponically grown plants showed a stronger response of hybrid cultivars to the co-provision of NH₄ ⁺ and NO₃ ^–. In soil-grown plants, N use efficiency in the hybrid was improved during ammonium fertilization compared to nitrate fertilization. Since ammonium-fertilized plants actually receive a mixture of NH₄ ⁺ and NO₃ ^– with ratios depending on root-associated nitrification activity, these results support the advantage of co-provision of ammonium and nitrate for the hybrid cultivar.     

Growth and morphology of eucalypt seedling-roots, in relation to soil strength arising from compaction

Information on the response of root growth and morphology to soil strength is useful for testing suitability of existing and new tillage methods and/or for selecting plants suitable for a specific site with or without tillage. Although there is extensive published information on the root growth-soil strength relationships for annual agricultural plants, such information is scarce for woody, perennial tree species. The purpose of this study is to examine growth and morphology of the root systems of 17-day-old eucalypt seedlings with respect to variation in soil strength. Soil strength in this study was varied by compaction of a well-aggregated clay soil to bulk densities of 0.7–1.0 Mg m^-3 whilst maintaining adequate water availability and aeration for plant growth. Lengths and tip-diameters of primary and lateral roots were measured on the excavated root systems of seedlings.With increase in bulk density and also soil strength (expressed as penetrometer resistance), total length of primary and lateral roots decreased. There were 71 and 31% reduction in the lengths of primary and lateral roots respectively with an increase in penetrometer resistance from 0.4 to 4.2 MPa. This indicated primary roots to be more sensitive to high soil strength than the lateral roots. Average length of lateral roots and diameters of both primary and lateral root tips increased with an increase in soil strength as well. There was greater abundance of lateral roots (no. of lateral roots per unit length of primary root) and root hairs with increased soil strength. The observed root behaviour to variable soil strength is discussed in the context of compensatory growth of roots and overall growth of plants.     

Response of citronella Java (Cymbopogon winterianus Jowitt) to VA mycorrhizal fungi and soil compaction in relation to P supply

Nutrient acquisition and growth of citronella Java (Cymbopogon winterianus Jowitt) was studied in a P-deficient sandy soil to determine the effects of mycorrhizal symbiosis and soil compaction. A pasteurized sandy loam soil was inoculated either with rhizosphere microorganisms excluding VAM fungi (non-mycorrhizal) or with the VAM fungus, Glomus intraradices Schenck and Smith (mycorrhizal) and supplied with 0, 50 or 100 mg P kg^-1 soil. The soil was compacted to a bulk density of 1.2 and 1.4 Mg m^-3 (dry soil basis). G. intraradices substantially increased root and shoot biomass, root length, nutrient (P, Zn and Cu) uptake per unit root length and nutrient concentrations in the plant, compared to inoculation with rhizosphere microorganisms when the soil was at the low bulk density and not amended with P. Little or no plant response to the VAM fungus was observed when the soil was supplied with 50 or 100 mg P kg^-1 soil and/or compacted to the highest bulk density. At higher soil compaction and P supply the VAM fungus significantly reduced root length. Non-mycorrhizal plants at higher soil compaction produced relatively thinner roots and had higher concentrations and uptake of P, Zn and Cu than at lower soil compaction, particularly under conditions of P deficiency. The quality of citronella Java oil measured in terms citronellal and d-citronellol concentration did not vary appreciably due to various soil treatments.     

Effects of ammonium sulphate application on the rhizosphere,fine-root and needle chemistry in aPicea abies (L.) Karst. stand

Rhizosphere, fine-root and needle chemistry were investigated in a 28 year old Norway spruce stand in SW Sweden. The uptake and allocation pattern of plant nutrients and aluminium in control plots (C) and plots repeatedly treated with ammonium sulphate (NS) were compared. Treatments started in 1988. Current year needles, one-year-old needles and cylindrical core samples of the LFH-layer and the mineral soil layers were sampled in 1988, 1989 and 1990. Compared to the control plots, pH decreased significantly in the rhizosphere soil in the NS plots in 1989 and 1990 while the SO₄-S concentration increased significantly. Aluminium concentration in the rhizosphere soil was generally higher in the NS plots in all soil layers, except at 0–10 cm depths, both in 1989 and 1990. Calcium, Mg and K concentrations also increased after treatment with ammonium sulphate. Ammonium ions may have replaced these elements in the soil organic matter. The NS treatment significantly reduced Mg concentrations in fine roots in all layers in 1990. A similar trend was found in the needles. Ca concentrations in fine roots were significantly lower in the NS plots in the LFH layer in 1990 and the same pattern was found in the current needles. The N and S concentrations of both fine roots and needles were significantly higher in the NS plots. It was suggested that NS treatment resulted in displacement of Mg, Ca and K from exchange sites in the LFH layer leading to leaching of these cations to the mineral soil. Further application of ammonium sulphate may damage the fine roots and consequently adversely affect the water and nutrient uptake of root systems.     

Fine-root morphology and uptake of 32P and 35S in a Norway spruce (Picea abies (L.) Karst.) stand subjected to various nutrient and water supplies

An investigation of fine (< 1 mm in diameter) and small (1–2 mm in diameter) roots in the organic soil layer was carried out in a Norway spruce forest stand with different treatments of water and nutrients, including control (C); ammonium sulphate application (NS); nitrogen-free fertilization (V); irrigation with liquid fertilization (a complete nutrient solution) (IF); NS followed by artificial drought (ND); V followed by artificial drought (VD). In order to evaluate the vitality and function of the fine roots, the following approaches were used: i) classification of fine roots, based on morphological characteristics; ii) nutrient uptake bioassay, using ³²P-phosphate and ³⁵S-sulphate; iii) nutrient concentration in fine roots and its relation to nutrient uptake. The NS treatment showed effects on the fine and small roots, with a decrease in amount of living roots, and a decrease in the total amount of fine and small roots. The VD treatment resulted in increased amounts of living small roots, while the ND treatment showed the opposite, as compared with the V and NS treatments, respectively. The uptake of P was negatively related to the P supply, with a higher P uptake for C and NS fine roots than for IF and V fine roots. The specific root length (SRL, m g^-1 DW) decreased for NS fine roots and increased for IF fine roots, indicating a further increase in uptake for NS roots and a decreased uptake for IF roots if calculated on a root length basis. So far, the NS and IF treatments maintain a considerable increase in above-ground biomass with a significantly reduced root biomass and standing crop.     

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.     

Effect of exogenous ammonium gluconate on growth,ion flux and antioxidant enzymes of maize (Zea Mays L.) seedlings under NaCl stress

• Ammonium gluconate (AG) provides both an organic carbon source and a nitrogen source, which can positively improve soil fertility and delay soil degradation.
• We investigated the underlying mechanisms of both NH₄⁺‐ and C₆H₁₁O₇^?‐mediated resistance to high salt concentrations in maize (Zea mays L.), and how they relate to antioxidant cellular machinery, root system architecture, root activity and lignin content in roots.
• Seedlings treated with AG maintained lower Na⁺ content, higher chlorophyll content, higher CAT and POD activity, compared with those without AG and ammonium carbonate (AC). The total size of the root system, primary root length and number of lateral roots detected on the primary root treated with AG decreased compared with those not treated with AG at the same NaCl concentration. However, average root diameter and root activity when treated with AG were significantly higher than roots without AG at the same NaCl concentration. Furthermore, total size of the root system, primary root length and number of lateral roots detected on primary rootsof seedlings treated with AG were higher than those treated with AC at the same NaCl concentration.
• These results suggested that AG may be a good organic fertiliser under salt stress by decreasing Na⁺ content and increasing chlorophyll content, activity of antioxidant enzymes, root diameter and root activity in maize seedlings.

     

The formation,morphology and anatomy of cluster root of Lupinus albus L. as dependent on soil type and phosphorus supply

The effect of phosphorus supply on the formation, morphology and anatomy of cluster roots of Lupinus albus L. cv Ultra grown in a loam and two sandy soils was examined relative to its effect on total root length, shoot weight and the phosphorus concentration of the shoots. The loam soil was most conducive to the formation of cluster roots. Cluster roots growing in the sandy soils developed to a lesser extent on plants of an equivalent phosphorus status, suggesting that some biotic or abiotic factors independent of phosphorus supply were also operating. The presence of mature cluster rootlets on a length of lateral root increased the root surface area by 14–22 times of an equal length of lateral roots not bearing cluster rootlets. The application of phosphorus decreased cluster-root length, whereas total root length showed a steady increase. There was an inverse relationship between cluster-root production and phosphorus concentration in shoots ranging from 2 to 8.5 mg g^–1 with the critical phosphorus level for maximum shoot growth being around 2.5 mg g^–1. Cluster roots formed in solution culture were not well developed in comparison with those grown in the loam soil or nutrient solution with added loam soil. The organisation of the cluster rootlet was similar to that of the lateral roots. Mature rootlets lacked an apical meristem and a vascular cambium with a reduced root cap and cortical tissue.     

Secretion activity of white lupin’s cluster roots influences bacterial abundance,function and community structure

White lupin (Lupinus albus L. cv. Amiga) reacts to phosphate deficiency by producing cluster roots which exude large amounts of organic acids. The detailed knowledge of the excretion physiology of the different root parts makes it a good model plant to study plant-bacteria interaction. Since the effect of the organic acid exudation by cluster roots on the rhizosphere microflora is still poorly understood, we investigated the abundance, diversity and functions of bacteria associated with the cluster roots of white lupin, with special emphasis on the influence of root proximity (comparing root, rhizosphere soil and bulk soil fractions) and cluster root growth stages, which are characterized by different excretion activities. Plants were grown for five weeks in microcosms, in the presence of low phosphate concentrations, on acidic sand inoculated with a soil suspension from a lupin field. Plate counts showed that bacterial abundance decreased at the stage where the cluster root excretes high amounts of citrate and protons. In vitro tests on isolates showed that the frequencies of auxin producers were highest in juvenile and mature cluster roots and significantly decreased in senescent cluster roots. However, no significant difference in the frequency of auxin producers was found between cluster and non cluster roots. The diversity and structure of bacterial communities were investigated by DGGE of 16S rDNA and 16S rRNA. The diversity and community structure were mostly influenced by root proximity and, to a lesser extent, by cluster root stage. The richness of bacterial communities decreased with root proximity, whereas the proportion of active populations increased. The high citrate and proton excretion occurring at the mature stage of cluster roots had a strong impact on the structure and richness of the bacterial communities, both in the root and in the rhizosphere soil.     

Plant growth, nutrient acquisition and mycorrhizal symbioses of a waterlogging tolerant legume (Lotus glaber Mill.) in a saline-sodic soil

Seedlings of Lotus glaberMill., were grown in a native saline-sodic soil in a greenhouse for 50 days and then subjected to waterlogging for an additional period of 40 days. The effect of soil waterlogging was evaluated by measuring plant growth allocation, mineral nutrition and soil chemical properties. Rhizobiumnodules and mycorrhizal colonisation in L. glaberroots were measured before and after waterlogging. Compared to control plants, waterlogged plants had decreased root/shoot ratio, lower number of stems per plant, lower specific root length and less allocation of P and N to roots. Waterlogged plants showed increased N and P concentrations in plant tissues, larger root crown diameter and longer internodes. Available N and P and organic P, pH and amorphous iron increased in waterlogged soil, but total N, EC and exchangeable sodium were not changed. Soil waterlogging decreased root length colonised by arbuscular mycorrhizal (AM) fungi, arbuscular colonisation and number of entry points per unit of root length colonised. Waterlogging also increased vesicle colonisation and Rhizobium nodules on roots. AM fungal spore density was lower at the end of the experiment in non-waterlogged soil but was not reduced under waterlogging. The results indicate that L. glaber can grow, become nodulated by Rhizobium and colonised by mycorrhizas under waterlogged condition. The responses of L. glaber may be related its ability to form aerenchyma.     

Effects of extracts of Alnus glutinosa seeds on growth of Frankia strain ArI3 under static and fermentor culture conditions

The amount of root mortality caused by root pathogens such as Phytophthora nicotianae (syn. Phytophthora parasitica) has typically been inferred from the net change in root length density in sequential soil cores. Because such measurements give information only on net changes in root populations, the actual rate of root turnover is often underestimated. We used minirhizotrons to track the fate of a large number of individual fine roots of mature field-grown citrus trees over a 6-month period. This method enabled us to examine the effect of P. nicotianae population levels on fine-root mortality. Seasonal and genotypic variation in patterns of citrus fine root mortality were associated with variation in population levels of P. nicotianae. Fine root lifespans were shorter when populations of P. nicotianae were high. Fine roots of the Phytophthora-susceptible rootstock, rough lemon (Citrus jamibhiri), had shorter median lifespans and supported larger populations of P. nicotianae than the fine roots of the more tolerant rootstock, Volkamer lemon (Citrus volkameriana). Rates of root mortality were either relatively constant for roots of all ages, or increased with age; the latter pattern was most pronounced for Volkamer lemon roots. Differences in the age-dependence of root mortality may, therefore, play a role in genotypic differences in tolerance of Phytophthora root rot by these two rootstocks. H Lambers Section editor     

米槠次生林和天然林细根养分及化学计量特征

探讨人为干扰对森林养分利用和生物地球化学循环特征的影响,对亚热带米槠(Castanopsis carlesii)次生林和天然林细根化学计量特征及其随土壤深度(0～80cm)的变化趋势进行了研究。结果表明,混合线性模型表明林分和土层深度对细根化学计量特征影响的主效应显著,但是交互作用不显著;米槠次生林细根N和P含量均显著低于天然林,而细根C浓度、C:N、C:P显著高于天然林,天然林1～2mm细根C浓度显著低于次生林;天然林和次生林细根N、P含量均随土层深度增加而呈显著下降趋势,C:N、C:P、N:P均随土层深度增加而呈显著上升趋势,且两林分下降趋势无显著差异;天然林和次生林细根N、P含量及N:P分别与土壤全N、全P含量和N:P存在线性关系,而细根N、P含量和N:P随土层深度变化均与直径显著相关,与细根比根长(SRL)无关。天然林经过人为干扰后,细根化学计量特征随土层深度变化规律虽未发生改变,但细根N和P浓度显著降低。     

Fine Root Production and Turnover in a Norway Spruce Stand in Northern Sweden: Effects of Nitrogen and Water Manipulation

Fine root length production, biomass production, and turnover in forest floor and mineral soil (0–30 cm) layers were studied in relation to irrigated (I) and irrigated-fertilized (IL) treatments in a Norway spruce stand in northern Sweden over a 2-year period. Fine roots (<1 mm) of both spruce and understory vegetation were studied. Minirhizotrons were used to estimate fine root length production and turnover, and soil cores were used to estimate standing biomass. Turnover was estimated as both the inverse of root longevity (RTL) and the ratio of annual root length production to observed root length (RTR). RTR values of spruce roots in the forest floor in I and IL plots were 0.6 and 0.5 y⁻¹, respectively, whereas the corresponding values for RTL were 0.8 and 0.9 y⁻¹. In mineral soil, corresponding values for I, IL, and control (C) plots were 1.2, 1.2, and 0.9 y⁻¹ (RTR) and 0.9, 1.1, and 1 y⁻¹ (RTL). RTR and RTL values of understory vegetation roots were 1 and 1.1 y⁻¹, respectively. Spruce root length production in both the forest floor and the mineral soil in I plots was higher than in IL plots. The IL-treated plots gave the highest estimates of spruce fine root biomass production in the forest floor, but, for the mineral soil, the estimates obtained for the I plots were the highest. The understory vegetation fine root production in the I and IL plots was similar for both the forest floor and the mineral soil and higher (for both layers) than in C plots. Nitrogen (N) turnover in the forest floor and mineral soil layers (summed) via spruce roots in IL, I, and C plots amounted to 2.4, 2.1, and 1.3 g N m⁻² y⁻¹, and the corresponding values for field vegetation roots were 0.6, 0.5, and 0.3 g N m⁻² y⁻¹. It was concluded that fertilization increases standing root biomass, root production, and N turnover of spruce roots in both the forest floor and mineral soil. Data on understory vegetation roots are required for estimating carbon budgets in model studies.     

The role of soil conditions in fine root ecomorphology in Norway spruce (Picea abies (L.) Karst.)

The present study is an attempt to investigate the pattern of morphological variability of the short roots of Norway spruce (Picea abies (L.) Karst.) growing in different soils. Five root parameters – diameter, length and dry weight of the root tip, root density (dry weight per water-saturated volume) and specific root area (absorbing area of dry weight unit) were studied with respect to 11 soil characteristics using CANOCO RDA analysis. The investigation was conducted in seven study areas in Estonia differing in site quality class and soil type. Ten root samples per study area were collected randomly from the forest floor and from the 20 cm soil surface layer. Eleven soil parameters were included in the study: humus content, specific soil surface area, field capacity, soil bulk density, pH (KCl and H₂O dilution's), N and Ca concentrations, Ca/Al and C/N ratios, and the decomposition rate of fine roots (<2 mm dia.). Root morphological characteristics most strongly related to the measured soil characteristics in the different sites were specific root area, root density and diameter of the short roots, the means varying from 29 to 42 m² kg⁻¹, from 310 to 540 kg m⁻³ and from 0.26 to 0.32 mm, respectively; root density being most sensitive. The most favourable site and soil types resulting in fine roots with morphological characteristics for optimizing nutrient uptake (e.g. low short root density and high specific root area) were Umbric Luvisol (Oxalis), Dystric Gleysol (Oxalis) and Gleyic Luvisol (Hepatica). These soil types correspond to highly productive natural forest stands of Norway spruce in Estonia. All measured soil variables explained 28% of total variance of the root characteristics. The most important variables related to root morphology were the humus content, field capacity and specific soil surface area. This revised version was published online in June 2006 with corrections to the Cover Date.     

Fertilization in a mature Scots pine (Pinus sylvestris L.) stand—effects on fine roots

The effects of liquid and solid fertilizers on fine-root development were studied in a 130-year-old Scots pine (Pinus sylvestris L.) stand. Ingrowth cores,viz. initially root-free mesh bags with sieved mineral soil taken outside the plots and driven to a depth of 30 cm from the soil surface, were subsequently resampled and the amount of fine roots was estimated. The total accretion of both fine-root length and dry weight was comparatively high in the liquid fertilization plot. The most substantial net accretion, however, during the 1984 period was in the control plot. The results of the study is that the ramification pattern of the fine roots was strongly influenced by fertilization. The average number of root tips per unit length was 9.9 cm⁻¹ in the control plot cm⁻¹, compared with both the solid (A and B) and liquid fertilization plots (2.3, 3.2 and 3.3 cm⁻¹, respectively) due to a greater occurrence of mycorrhizal aggregates (‘ball mycorrhiza’). The effects of fertilization on the mineral nutrient concentrations in the fine roots were limited—the strongest effects were in the liquid-fertilized area. The observed increase in the concentration of most mineral nutrients in the latter experimental area, however, may be due to a change in the growth pattern of the fine roots rather than an effect of the fertilizers themselves.