Interactions of Vesicular-Arbuscular Mycorrhizal Fungi,Phosphorus, and Heterodera glycines on Soybean

Effects of vesicular-arbuscular mycorrhizal (VAM) fungi and soil phosphorus (P) fertility on parasitism of soybean cultivars Bragg and Wright by soybean cyst nematode (SCN) were investigated in field micropiot and greenhouse experiments. VAM fungi increased height of both cultivars and yield of Wright in microplot studies in 1986 and 1987. Conversely, yield of mycorrhizal and nonmycorrhizal plants of both cultivars was suppressed by SCN. Soil population densities of SCN were unaffected by VAM fungi in 1986 but were greater in microplots infested with VAM fungi than in control microplots in 1987. Growth of Wright soybean was stimulated by VAM fungi and suppressed by SCN in greenhouse experiments. The effect of VAM fungi on SCN varied with time. Numbers of SCN in roots and soil were decreased by VAM fungi by as much as 73% at the highest SCN inoculum level through 49 days after planting. Later, however, SCN numbers were usually comparable on mycorrhizal and nonmycorrhizal plants. Soil P fertility generally had no effect on SCN. Results of a split-root experiment indicated that VAM fungal suppression of SCN was not systemic.     

Genetic diversity for mycorrhizal symbiosis and phosphate transporters in rice

Phosphorus(P) is a major plant nutrient and developing crops with higher P-use efficiency is an important breeding goal.In this context we have conducted a comparative study of irrigated and rainfed rice varieties to assess genotypic differences in colonization with arbuscular mycorrhizal(AM) fungi and expression of different P transporter genes.Plants were grown in three different soil samples from a rice farm in the Philippines.The data show that AM symbiosis in all varieties was established after 4 weeks of growth under aerobic conditions and that,in soil derived from a rice paddy,natural AM populations recovered within6 weeks.The analysis of AM marker genes(AM1,AM3,AM14) and P transporter genes for the direct Pi uptake(PT2,PT6) and AM-mediated pathway(PT11,PT13) were largely in agreement with the observed root AM colonization providing a useful tool for diversity studies.Interestingly,delayed AM colonization was observed in the aus-type rice varieties which might be due to their different root structure and might confer an advantage for weed competition in the field.The data further showed that P-starvation induced root growth and expression of the high-affinity P transporter PT6 was highest in the irrigated variety IR66 which also maintained grain yield under P-deficient field conditions.     

The Vesicular Arbuscular Mycorrhiza Associated with Three Cultivars of Rice (Oryza sativa L.)

The present study deals with the occurrence of vesicular arbuscular mycorrhizal fungi in three cultivars of rice in Barak valley. Three cultivars of rice were Pankaj, Malati and Ranjit. The results revealed the association of VAM fungi in all the cultivars of rice. The association was maximum in Pankaj cultivar followed by Malati, and Ranjit, respectively, in all the three sampling phases. All the three cultivars of rice crop showed maximum soil spore population and number of VAM fungal species at the harvesting phase (135 DAS) and minimum at the phase of maturation (90 DAS). Glomus species were found dominating followed by Acaulospora species. Glomus microcarpum, Glomus claroideum, Glomus mosseae and Acaulospora scrobiculata were found in all the three fields.     

Distribution of vesicular-arbuscular mycorrhizal fungi in the natural ecosystem

Natural occurrence of vesicular-arbuscular mycorrhizal (VAM) fungi in Haryana soils showed that VAM sporulation was more intensive in the rhizosphere of nonlegumes than of legumes. Maximum number of spores (342 spores per 50 g of soil) was observed in the rhizosphere of mustard, followed by chickpea, wheat, pearl millet and pigeonpea. Four VAM generaviz. Glomus, Gigaspora, Sclerocystis andAcaulospora, were present there. Soil pH, total soil P, available P, type of soil, soil moisture and cropping season all variables influenced the VA mycorrhizal population in the natural ecosystem. Numbers of VAM spores highly correlated with the presence of total soil P and soil pH indirectly affected the VAM population through the total soil P. The spore population was abundant in sandy soils as compared to loamy sands. Drier soils had higher number of VAM spores. In summer, the VAM population in soil was less as compared to winter season.     

The interaction of rockphosphate,Bradyrhizobium, vesicular-arbuscular mycorrhizae and phosphate-solubilizing microbes on soybean grown in a sub-Himalayan mollisol

A factorial design 2³ × 4 with two levels of Mussorie rockphosphate (RP) with or without vesicular-arbuscular mycorrhizal (VAM) fungi and Bradyrhizobium japonicum, and four treatments of phosphate-solubilizing microbes (PSM) Pseudomonas striata, Bacillus polymyxa, Aspergillus awamori was employed using Patharchatta sandy loam soil (Typic Hapludoll). The observations included mycorrhization, nodulation, grain and straw yield, N and P uptake, available soil P and the PSM population in the soil after crop harvest. Inoculation with endophytes alone caused about 70% root colonization. Addition of rockphosphate or inoculation with PSM, except B. polymyxa, stimulated root infection of native as well as introduced VAM endophytes. Application of RP or inoculation with Bradyrhizobium japonicum, mycorrhizal fungi or phosphate-solubilizing microorganisms significantly increased nodulation, N uptake, available soil P and the PSM population in the soil after the crop harvest. The grain and straw yields did not increase following RP addition or mycorrhizal inoculation but increased significantly after inoculation wit Bradyrhizobium or PSM. In general, the application of RP, Bradyrhizobium, VAM and PSM in combinations of any two or three resulted in significant increases in nodulation, plant growth, grain yield and uptake of N and P. Among the four factor interactions, rockphosphate, Bradyrhizobium and P. striata in the absence of VAM resulted in maximal nodulation, grain and straw yields and N uptake by soybean. The highest P uptake by soybean grain was recorded with Bradyrhizobium and A. awamori in the absence of rockphosphate and VAM. Generally, available soil P and PSM population after crop harvest were not significantly increased by the treatment combinations giving the maximal uptake of nutrients. However, they increased significantly in response to PSM, which produced no significant increase in total uptake of nutrients.Research paper no. 7498     

冬季覆盖作物秸秆还田对双季稻田根际土壤微生物群落功能多样性的影响

微生物群落功能多样性是土壤质量变化重要的指标,不同作物类型的秸秆还田措施对土壤微生物群落功能多样性具有明显的影响。以位于双季稻主产区不同冬季覆盖作物-双季稻种植模式大田定位试验田为研究对象,以冬闲-双季稻种植模式为对照(CK),应用Biolog-GN技术开展黑麦草-双季稻(Ry)、紫云英-双季稻(Mv)、油菜-双季稻(Ra)和马铃薯-双季稻(Po)种植模式条件下不同冬季覆盖作物秸秆还田后对双季稻田根际土壤微生物功能多样性影响的研究。研究结果表明,早稻和晚稻成熟期,与CK处理相比,冬季覆盖作物秸秆还田处理增加了稻田土壤碳源平均颜色变化率(AWCD),以Po处理AWCD均为最高,均显著高于Ry和CK处理。不同冬季覆盖作物秸秆还田处理土壤微生物代谢多样性指数表现出明显的差异,早稻成熟期,Po处理的Richness、Shannon和McIntosh指数均为最高,其次为Ry、Mv和Ra处理,CK处理最低;晚稻成熟期,各处理的Richness、Shannon和McIntosh指数大小顺序均表现为PoRaMvRyCK。土壤微生物碳源利用的主成分分析结果表明,各冬季覆盖作物秸秆还田处理根际土壤微生物利用的主要碳源为氨基酸类和糖类物质,不同处理间碳源利用类型有差异。冬季覆盖作物秸秆还田措施有利于提高双季稻田根际土壤微生物对碳源的利用能力、物种丰富度和均匀度。     

Seasonality of VAM infection in three populations of Agrostis capillaris (Gramineae) on soil with or without heavy metal enrichment

Three populations of the perennial grass Agrostis capillaris, growing on limestone derived clay with and without natural enrichment of the heavy metals cadmium, lead, and zinc, and on a sandy soil polluted by a metal smelter have been investigated with regard to the percentage and seasonality of infection with vesicular-arbuscular mycorrhizal (VAM) fungi and its impact on mineral nutrition.In all populations VAM infection was lowest during winter, and highest during late summer and autumn. The population at the metal smelter site was less infected by VAM fungi than both other populations. The concentration of mineral nutrients for the three populations was clearly related to the soil concentration, but hardly modified by the degree of VAM infection.     

Influence of temperature on colonization of spring barleys by vesicular arbuscular mycorrhizal fungi

The effects of three soil temperatures on growth of spring barleys (Hordeum vulgare L.) and on their root colonization by vesicular arbuscular mycorrhizal (VAM) fungi from agricultural soils in Montana (USA) or Syria at different inoculum concentrations were tested in soil incubators in the greenhouse. The number of mycorrhizal plants as well as the proportion and intensity of roots colonized increased with higher soil temperatures. VAM fungi from Montana, primarily Glomus macrocarpum, were cold tolerant at 11°C while those from Syria, primarily G. hoi, were heat tolerant at 26°C. Inoculum potential of Montana VAM fungi was higher than Syrian VAM fungi in cool soils. Harmal, selected from Syrian barley land races, had the highest colonization by mycorrhizal fungi of the cultivars tested.Journal Series Paper: J-2532 Montana Agricultural Experiment Station.     

Phosphorus cycling in rainfed lowland rice ecosystems on sandy soils

Phosphorus cycling in rainfed lowland rice ecosystems is poorly understood. Soil drying and grazing of rice straw during the long dry season, the growth of volunteer pastures during the early wet season, and intermittent loss of soil-water saturation while the rice crop is growing are important distinguishing characteristics of the rainfed lowlands in relation to P cycling. We studied P cycling in an acid sandy rainfed lowland soil that covers about 30% of the rice growing area of Cambodia. Soils with similar properties in comparable rainfed sub- ecosystems occur in Laos and northeast Thailand. We developed a general schema of P pools and fluxes in the crop and soil for rice-based cropping systems in the rainfed lowlands of Cambodia. The schema was derived from a number of field experiments carried out over five consecutive cropping seasons to quantify the residual value of P fertiliser, P mass balances, soil P fractions, the effect on subsequent rice crops of crop residues and volunteer pastures incorporated into the soils, and the dynamics of P turnover in the soil. With a single rice crop yielding 2.5–3 t ha⁻¹, application of 8–10 kg P ha⁻¹ maintained yields and a small positive P balance in the soil. However, the soil P balance was sensitive to the proportion of rice straw returned to the soil. Volunteer pastures growing during the early wet season accumulated significant amounts of P, and increased their P uptake when soils were previously fertilised with P. These pastures recycled 3–10 kg P ha⁻¹ for the succeeding rice crops. While inorganic soil P pools extractable with ion exchange resins and 0.1 M NaOH appeared to be the main source of P absorbed by rice, microbial and organically-bound P pools responded dynamically to variation in soil water regimes of the main wet, dry and early wet seasons. The schema needs to be developed further to incorporate site-specific conditions and management factors that directly or indirectly affect P cycling, especially loss of soil-water saturation during the rice cropping cycle. The paper discusses the application of results for acid sandy soils to other significant rice soils in the rainfed lowlands of southeast Asia.     

The influence of host plant,nitrogen fertilization and fungicide application on the abundance and seasonal dynamics of vesicular-arbuscular mycorrhizal fungi in arable soils of northern Germany

The present investigation examines whether the crop plant, increased nitrogen (N) fertilization and fungicide application influence the pattern of vesicular-arbuscular mycorrhizal fungi (VAM) populations. For this purpose, two arable field locations in Lower Saxony (Hotteln and Langreder) were chosen and the formation of mycorrhiza, spore density, number of infectious propagules (MPN) and frequency of spore types within VAM populations were investigated. The influence of crop plants was examined over two cultivation periods (1986/1987 and 1987/1988) in Hotteln, comparing winter wheat, winter barley and sugar beet. The effects of increased N fertilization and fungicide application were investigated on winter wheat in Langreder in the cultivation period 1988 only. Both the frequency of mycorrhizal infection and the spore dynamics in soil differed with the crops grown. Spore density and MPN increased until harvesting when host plants (winter wheat, winter barley) were cultivated, whereas both diminished with a non-host plant (sugar beet). Different spore types increased or decreased, according to the plant species grown, but the predominating types of the location remained constant. Increased N fertilization caused marginal inhibition of mycorrhizal infection and sporulation on winter wheat, whereas both leaf and base application of fungicides resulted in minor increases in mycorrhizal colonization of roots and sporulation in soil. Both increased N fertilization and fungicide application distinctly decreased the sporulation of one type in May, but the characteristic compositions of the VAM populations remained unchanged.     

Influence of inoculum density of two vesicular–arbuscular mycorrhizal fungi and temperature/light intensity on Trifolium repens

The influence of three inoculum densities of Glomus caledonius and G. epigaeus and two temperature/light intensity conditions was investigated on Trifolium repens. The significance of inoculation was compared to the significance of naturally occurring vesicular–arbuscular mycorrhizal (VAM) fungi and to application of soluble phosphate fertilizer. Increasing density of inoculum and the highest temperature/light intensity condition tested increased VAM infection, whereas only small differences were found between efficiency of the two introduced VAM fungi. The presence of naturally occurring VAM fungi proved as efficient in establishing infection as the most successful inoculations. Some interactions among the investigated parameters were found for several recordings. The increase in VAM infection was followed by an increase in number of nodules; in uptake of phosphorus, nitrogen, zinc, and copper; and in growth of roots and shoots. The calculated inflow of phosphate, zinc, and copper into roots was not associated with inoculum density, VAM species or temperature/light conditions. Compared to an uninoculated control without application of phosphate, inoculation with the highest spore density increased (after 18 weeks growth) the dry weight of shoot 52 fold and 7 fold for G. caledonius , and 121 fold and 9 fold for G. epigaeus at low and high temperature/light conditions, respectively. It was also found that VAM increased weight per nodule 52% when roots with no or sparse VAM infection were compared to roots with low to maximal VAM infection and 98% when roots with low VAM infection were excluded. Application of phosphate fertilizer enhanced nodulation and growth of non–mycorrhizal plants to a level similar to that of the most heavily VAM infected plants.     

Mycorrhizal Interactions in Sustainable Agriculture

Abstract

Vesicular-arbuscular mycorrhizal (VAM) fungi are an intimate link between the roots of most crop plants and soils, thereby affecting the development of host plants and host soils. The role of VAM fungi in improving plant nutrition and their interactions with other soil biota have been investigated with reference to host plant growth, but little is known about how these interactions affect soil structure. The impact of cultural practices and the particular role that VAM fungi play in improving soil structure are discussed in the context of sustainable farming.     

Interactions between vesicular-arbuscular mycorrhizal fungi and asparagus

Summary The vesicular-arbuscular mycorrhizal (VAM) fungus,Glomus versiforme increased significantly the growth ofAsparagus officinalis under controlled conditions using Turface as the growth medium. The growth responses, including increases in root fresh weight, numbers of shoots, shoot dry weight, and shoot height follow a pattern similar to other mycorrhizal systems. Indigenous VAM fungi appeared to have negative effects on average shoot fresh and dry weight, number of shoots per pot and average shoot height on one year oldA. officinalis seedlings obtained from the field and grown under controlled conditions. These results may be due either to the high levels of soluble phosphate present in the soil or the ineffectiveness of the particular indigenous fungi as mycorrhizal fungi in asparagus. Indigenous mycorrhizal fungi overwinter in asparagus root crown as vesicles and as external and internal hyphae. Soil obtained from the same fields as the one year old crowns was a good source of mycorrhizal inoculum for sterile seedlings.     

Field nursery inoculation of Hevea brasiliensis Muell. Arg. seedling rootstock with vesicular-arbuscular mycorrhizal (VAM) fungi

The growth response of Hevea brasiliensis to vesicular-arbuscular mycorrhizal (VAM) fungi inoculation was assessed in two field nursery sites containing indigenous mycorrhizal fungi (IMF). Seedling rootstocks were inoculated with mixed VAM-fungal species in a factorial combination with phosphorus (P) fertilizer application, and planted in randomised blocks on sandy (site 1) and clayey (site 2) soils. Plants were harvested after 26 weeks for measurements of shoot dry weight (DW), stem diameter, height, mycorrhizal root colonization and leaf nutrient contents. At site 1, VAM increased shoot DW, stem diameter and plant height only in treatments without P applied. Increases in shoot DW due to VAM were 70% greater than the uninoculated controls although this was reduced to 5% when P was applied. At site 2, VAM inoculation also increased shoot DW and stem diameter but the magnitude of the increases was smaller. Shoot DW response due to VAM was only 29%. At this second site, applying phosphate to uninoculated plants did not increase shoot yields further. Leaf concentrations of all nutrients were unaffected by VAM at both sites, except for copper (Cu) which was increased by VAM in treatments where P was not applied. However, leaf contents of P, potassium (K), magnesium (Mg) and Cu were increased by VAM at site 1, and of leaf nitrogen (N) and K at site 2. These experiments demonstrate that VAM-fungi could be introduced into field nursery sites to improve growth and P uptake by H. brasiliensis. The relevance of VAM-fungi to H. brasiliensis seedling rootstock development and the influence of IMF in determining field responses is discussed.     

Preinoculation of lettuce and onion with VA mycorrhizal fungi reduces deleterious effects of soil salinity

The hypothesis that inoculation of transplants with vesicular-arbuscular mycorrhizal (VAM) fungi before planting into saline soils alleviates salt effects on growth and yield was tested on lettuce (Lactuca sativa L.) and onion (Allium cepa L.). A second hypothesis was that fungi isolated from saline soil are more effective in counteracting salt effects than those from nonsaline soil. VAM fungi from high- and low-salt soils were trap-cultured, their propagules quantified and adjusted to a like number, and added to a pasteurized soil mix in which seedlings were grown for 3–4 weeks. Once the seedlings were colonized by VAM fungi, they were transplanted into salinized (NaCl) soil. Preinoculated lettuce transplants grown for 11 weeks in the saline soils had greater shoot mass compared with nonVAM plants at all salt levels [2 (control), 4, 8 and 12 dS m^–1] tested. Leaves of VAM lettuce at the highest salt level were significantly greener (more chlorophyll) than those of the nonVAM lettuce. NonVAM onions were stunted due to P deficiency in the soil, but inoculation with VAM fungi alleviated P deficiency and salinity effects; VAM onions were significantly larger at all salt levels than nonVAM onions. In a separate experiment, addition of P to salinized soil reduced the salt stress effect on nonVAM onions but to a lesser extent than by VAM inoculation. VAM fungi from the saline soil were not more effective in reducing growth inhibition by salt than those from the nonsaline site. Colonization of roots and length of soil hyphae produced by the VAM fungi decreased with increasing soil salt concentration. Results indicate that preinoculation of transplants with VAM fungi can help alleviate deleterious effects of saline soils on crop yield.     

Response ofLeucaena leucocephala to vesicular-arbuscular mycorrhizal colonization and rock phosphate fertilization in an Oxisol

Response ofLeucaena leucocephala (Lam) de Wit to rock phosphate application and inoculation with the vesicular-arbuscular mycorrhizal (VAM) fungusGlomus aggregatum (Schenck and Smith emend Koske) was evaluated in a pot experiment. VAM colonization increased as rock phosphate application increased. Using phosphorus concentration in pinnules as an indicator of VAM activity, significant VAM activity occurred at 25 days after planting at the lower levels of rock phosphate application (0, 0.34 and 0.68 g P kg^–1). The time required for significant VAM activity was shortened by 5 days at the higher P levels (1.36, 2.72 and 5.44 g P kg^–1). The highest VAM activity was associated with the highest rate of rock phosphate application.Inoculation withG. aggregatum significantly increased the uptake of Cu, P and Zn and dry-matter yield at all levels of rock phosphate applied. Copper concentrations in roots of mycorrhizal Leucaena were significantly higher than that of shoots. The results indicated that Leucaena in symbiotic association with VAM fungi effectively utilized P from rock phosphate. However, high rates of rock phosphate are required to attain growth comparable to that obtained with the application of water-soluble phosphate.Contribution from Hawaii Institute of Tropical Agriculture and Human Resources, Journal Series No. 3243.     

VA-mycorrhizae and mycorrhiza stimulating isoflavonoid compounds reduce plant herbicide injury

Imazaquin, imazethapyr and pendimethalin showed high toxicity to sorghum plants grown in a greenhouse soil mix. However, mycorrhizal sorghum plants were less affected by herbicide toxicity than non-mycorrhizal ones, at low to moderate herbicide concentrations. VAM herbicide safening effects were more evident on imazaquin-treated plants, than for those treated with the other two herbicides. Applications of imazethapyr and pendimethalin at the two highest concentrations, but not imazaquin, reduced VAM colonization rates in sorghum. Applications of the VAM stimulating isoflavonoids, biochanin A and formononetin, at 5 ppm solutions to a field soil sample containing toxic levels of imazaquin (13 ppb) and indigenous VAM fungi, reduced herbicide-induced injury in corn and sorghum under growth chamber conditions. The benefits of isoflavonoids were reduced when additional propagules of Glomus intraradix were added into field-soil samples, and were eliminated when VAM fungi were inactivated by autoclaving. This indicates that herbicide safening effects of biochanin A, and formononetin are VAM-mediated and also suggests the potential use of these isoflavonoids as herbicide safeners.     

Interaction of Endomycorrhizal Fungi,Superphosphate, and Meloidogyne incognita on Cotton in Microplot and Field Studies

Microplot and field experiments were conducted to determine the effects of two vesicular-arbuscular mycorrhizal (VAM) fungi, Glomus intraradices (Gi) and Gigaspora margarita (Gm), and dicalcium phosphate (P) on Meloidogyne incognita (Mi) reproduction and seed cotton yield of the Mi-susceptible cotton cultivar, Stoneville 213. In 1983 population densities of Mi juveniles were significantly lower 60 and 90 days after planting in microplots receiving Gi. Mycorrhizal fungi reduced the severity of yield losses to Mi, whereas P fertilization increased yield losses to Mi. In 1984 microplot yields were reduced linearly as nematode inoculum densities increased in treatments of Mi alone, Gm, or P, but the response was curvilinear with Gi. Yield suppressions in the 1984 field experiment occurred only in plots infested with Mi alone. In the 1984 microplots, numbers of Mi juveniles penetrating seedling roots increased Iinearly with increasing nematode inoculum densities and was favored when mycorrhizal fungi or superphosphate were added. Juvenile penetration of roots was negatively correlated with yields in all treatments (r = -0.54 to -0.81) except Gm and with number of bolls in Mi alone (r = -0.85) and P (r = -0.81) treatments. Mycorrhizal fungi can increase host tolerance to M. incognita in field conditions and may function as important biological control agents in soils infested with high population densities of efficient VAM species.     

Effects of rhizosphere soil,vesicular-arbuscular mycorrhizal fungi and phosphate onPlantago major L. ssp.pleiosperma Pilger

Biotic factors in the rhizosphere and their effect on the growth ofPlantago major L. ssp.pleiosperma Pilger (Great plantain) were studied. In a pot experiment the effect on shoot growth of the addition of 2.5% rhizosphere soil at four levels of phosphate was highly dependent on the availability of phosphate: a promoting effect at low phosphate levels was observed while a reducing effect occurred at higher phosphate levels. As the roots were infected with vesicular-arbuscular mycorrhizal (VAM) fungi in the treatment with rhizosphere soil, two other experiments were set up to separate effects of the indigenous VAM fungi from effects of the total rhizosphere population. The uptake of phosphate and shoot growth was not decreased at higher phosphate availability when VAM inoculum was added alone or in combination with rhizosphere soil. The growth reducing effect of the rhizosphere soil could therefore not be ascribed only to mycorrhizal infection. The results suggest that biotic factors in the rhizosphere soil affect the phosphate uptake ofPlantago major ssp.pleiosperma. This may, under conditions of phosphate limitation, lead to an increase of phosphate stress and, subsequently, a growth reduction. Futhermore, it is concluded that VAM fungi, as part of the rhizosphere population, may compensate this phosphate stress by enhancing the phosphate uptake.Grassland Species Research Group Publication No. 148.     

Nutrient availability effects on vesicular-arbuscular mycorrhizal bell pepper (Capsicum annuum) seedlings and transplants

Pepper (Capsicum annuum) seeds were sown in nutrient-poor sand or nutrient-rich peat/vermiculite amended or not amended with Glomus macrocarpum. The vesicular-arbuscular mycorrhizal (VAM) seedlings were irrigated with three levels of nutrient solution, and transplanted into four levels of P-amended soil, each of which was irrigated with two levels of nutrient solution minus P. Mycorrhizal seedlings in sand were responsive to increasing nutrient levels; in nutrient-rich peat the seedlings did not respond to additional fertilisation. The greatest seedling development accompanied by good fungus colonisation was in nutrient-poor medium irrigated with the highest nutrient solution tested (18 mM N, 1.2 mM P, and 7 mM K). Non-VAM plants almost ceased growing between the weeks 4 and 5, whereas VAM plants increased in weight by 41–188%. After transplanting, sand-grown seedlings benefited from VAM when 300 mg P/kg or more was added to the soil but peat-grown plants did not. Fruit development was delayed in all non-VAM plants compared with VAM ones.