Receptivity of various wheat cultivars to infection by VA-mycorrhizal fungi as influenced by inoculum potential and the relation of VAM-effectiveness to succinic dehydrogenase activity of the mycelium in the roots

Wheat cultivars grown in pots in a greenhouse were inoculated either once or twice with the vesicular-arbuscular mycorrhizal fungus (VAMF) Glomus mosseae. If inoculum was only added to the soil once (before planting) the cultivars showed differences in subsequent VAM development. If additional inoculum was added ten days after planting, VAM development was much increased and cultivars which remained without VAM after only one inoculum application now became mycorrhizal.Succinate dehydrogenase (SDH) activity decreased throughout the experiment, but this decrease was less rapid in mycelium in twice-inoculated plants.No close relationship between SDH-activity and plant growth (VAM effectiveness) was found.     

Effect of aromatic plant species on vesicular-arbuscular mycorrhizal establishment in Pistacia terebinthus

The inoculation of Pistacia terebinthus with vesicular-arbuscular mycorrhizal (VAM) fungi and the spread of the infection were studied using a mixed cropping system, under glasshouse conditions, with Salvia officinalis, Lavandula officinalis and Thymus vulgaris colonized by Glomus mosseae as an inoculation method. This method was compared with soil inoculum placed under the seed or distributed evenly in the soil. Indirect inoculation with all the aromatic plants tested significantly increased VAM root colonization of P. terebinthus compared with the use of soil inoculum, although the effect on plant growth was different for each one of the aromatic species used as inoculum source. Inoculation with L. officinalis and T. vulgaris were the best treatments resulting in high VAM colonization and growth enhancement of P. terebinthus.     

Categories of vesicular-arbuscular mycorrhizal dependency of host species

Summary Brassica nigra and selected species of Leucaena and Sesbania were used as indicator hosts in a greenhouse experiment designed to establish distinct categories of mycorrhizal dependence. The plants were grown in an oxisol with different concentrations of established soil solution P in the presence or absence of the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus aggregatum. The extent to which the plant species depended on the fungus for dry matter production diminished with increased concentrations of soil solution P, but the magnitude of this decrease varied from species to species. Five distinct mycorrhizal categories are proposed based on the differences observed, ranging from non-dependent to very highly dependent. The critical soil solution P concentrations that were useful for separating host species into distinct VAM-dependency groups were 0.02 and 0.2 mg/l. Species differing in their mycorrhizal dependency differed with respect to the soil solution P concentration required for the expression of maximum VAM effectiveness, the degree to which increasing concentrations of P depressed VAM infection and the pattern of immobile nutrient accumulation.Contribution from the Hawaii Institute of Tropical Agriculture and Human Resources Journal Series No. 3547     

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.     

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.     

Alteration of growth ofSolanum opacum andPlantago drummondii and inhibition of regrowth of hyphae of vesicular-arbuscular mycorrhizal fungi from dried root pieces by manganese

Addition of MnSO₄ or MnCl₂ to a fine sandy soil from South Australia had a negative effect on shoot growth and root elongation ofSolanum opacum in the absence of significant presence of vesicular-arbuscular mycorrhiza (VAM). VAM ameliorated the reduction of plant growth by Mn, even though mycorrhizal development was decreased. Mn inhibited infection of roots by a fine endophyte less than that by some coarse endophytes. High concentrations of available Mn inhibited growth of hyphae of VAM fungi from dried root pieces, a significant source of infection by mycorrhizal fungi in the soil used.     

Occurrence and importance of mycorrhizae in aquatic trees of New South Wales,Australia

Vesicular arbuscular mycorrhizal (VAM) infection was found in KOH-cleared and lactophenolblue-stained roots of Salix babylonica, Melaleuca quinquenervia and Casuarina cunninghamiana. These are all trees growing on creeks and river banks, in stationary or slowly flowing fresh or brackish waters in swamps, creeks, drains and channels, and in seepage areas of New South Wales, Australia. Larger and older roots lacked VAM infection in the inner cortex, probably due to suberisation of cells, and the endophyte was restricted to the epidermal layers. Spores and sporocarps of the VAM fungi Glomus fasciculatus, G. mosseae, Sclerocystis rubiformis, Gigaspora margarita and an unidentified Scutellospora sp. were wet sieved and decanted from aquatic sediments and soils. The presence of similar VAM fungal spores in the aquatic sediments and terrestrial soil suggests that they probably enter the aquatic sediments through run off from the land ecosystem. All three plants formed vesicular arbuscular (VA) mycorrhizae almost exclusively in the marshy, periodically inundated soils, but the same plant species formed endo-/ ectomycorrhizae when growing in soil with higher redox potentials (E _h). Salix and Melaleuca tree roots possessed both VAmycorrhizae and ectomycorrhizae. VAM roots of Casuarina were equipped with both N-fixing Frankia nodules and proteoid roots. VAM endophytes did not invade nodular cortical tissues, suggesting the presence of an exclusion mechanism which needs further study. The highest VAM infection was found in nodulated specimens. Free-floating roots growing in water close to the banks were non-mycorrhizal but were mycorrhizal in the bottom-rooting state. VAM spore number and mycorrhizal infection seem to be associated with redox-potential, i.e. lower at sites such as swamps, water or sediments with lower E _h values than in terrestrial soils with higher E _h values. A relationship between soil moisture gradient and VAM infection pattern became apparent from the study of a C. cunninghamiana transect on a creek embankment, i.e. typical vesicles and arbuscules were found in roots from drier soils, there was a lack of arbuscules in relatively wet soils but large lipid-filled intracellular vesicles were present, and typical vesicles and arbuscules were absent in flooded creek beds where roots were associated with coenocytic intercellular hyphae with abundant lipid droplets. The importance of VA mycorrhiza, ectomycorrhizae, N-fixing root nodules and proteoid roots at the land-water interface is discussed with reference to the use of these trees as pioneering species for stabilising river and stream banks, reducing erosion, windbreaking, and as a long-term and inexpensive means of achieving biological control of aquatic weeds by shading waterways.     

ROLE OF MYCORRHIZAL FUNGUS ORIGIN IN GROWTH AND NUTRIENT UPTAKE BY GERANIUM ROBERTIANUM

Prior field studies have shown that populations of forest herbs on relatively nutrient poor soils have higher vesicular-arbuscular mycorrhizal (VAM) infection intensity than plants on rich soils. However, the growth responses and ability to take up P against the soil nutrient gradient are often not linearly related to infection intensity. To determine if intraspecific differences among populations of the common VAM fungus Glomus occultum could differentially affect growth and nutrient uptake, Geranium robertianum seedlings were inoculated with Glomus occultum isolated from four forest types along a gradient of soil fertility, and grown in a greenhouse at P levels typical of the extremes of that gradient. Plants given inoculum from relatively infertile forest sites generally produced greater root, shoot, and total mass than plants given inoculum from fertile sites or uninoculated plants, especially at the low P supply rate. Total P uptake and both P and N uptake efficiency were also highest in plants given inocula from low fertility sites. These results indicate that local adaptation and intraspecific variations in the ability of VAM fungi to induce growth and nutrient uptake effects on host plants may be as important as interspecific differences among VAM fungus species.     

Effects of P-fertilization and inoculation by two vesicular-arbuscular mycorrhizal fungi on growth and nodulation ofCalopogonium caeruleum

The growth response ofCalopogonium caeruleum, a leguminous covercrop in plantation agriculture, to inoculation with two vesicular-arbuscular mycorrhizal (VAM) fungi was investigated in five phosphorus (P)-deficient soils supplied with various levels of rock phosphate. Significant shoot yield increases over the uninoculated controls were obtained in most sterilised or unsterilised soils at all applied P levels, although the inoculant VAM fungi differed in their effectiveness in the soils used. Responses in mycorrhizal root infections, P and nitrogen (N) concentrations in tops and plant nodulation varied. The results are discussed in relation to the edaphic environment of the mycorrhizal association.     

Vesicular-arbuscular mycorrhizal inoculum potential affects the growth of Stryphnodendron microstachyum seedlings in a Costa Rican human tropical lowland

This study used a plant bioassay to investigate the vesicular-arbuscular mycorrhizal (VAM) inoculum potential of soil from three vegetation types (fern, secondary forest, and grass) in an abandoned pasture in the tropical humid lowlands at La Selva, in northeastern Costa Rica. Growth, measured as seedling height, number of leaves, and total (above- and belowground) biomass, of Stryphnodendron microstachyum Poepp. et Endl. (Synon. S. excelsum Harms) seedlings was significantly lower when grown in soil inoculum from the fern areas than in soil inoculum from the forest and grass areas. However, S. microstachyum seedlings grown in the fern inoculum had significantly greater VAM colonization than seedlings grown in the forest and grass inoculum. In addition, roots collected from a dominant plant species from each of the three vegetation types showed that the fern (Nephrolepsis biserrata) had significantly greater mycorrhizal colonization than the tree (Pentaclethra macroloba (Willd.) Kuntze or the grass (Brachiaria spp.). The results of this study suggest that differences in mycorrhizal inoculum potential among vegetation types and its effects on seedling growth may have important implications for the restoration and management of degraded lands.     

Yellow leaf spot of cashew: A case of molybdenum deficiency

Summary Rooted cuttings ofSeverinia buxifolia were inoculated with the vesicular-arbuscular mycorrhizal (VAM) fungusGlomus intraradices or provided an inoculum filtrate (non-VAM plants) and grown in one of seven media combinations of fired montmorillonite clay (FMC) and Canadian peatmoss (CP) at ratios of 100%, 80%, 67%, 50%, 33%, 20%, or 0% FMC. Mycorrhizal infection increased with higher proportions of FMC, but the growth of both VAM and non-VAM plants was reduced with increased FMC amendment. The growth benefit (top and root fresh-dry weights) conferred by mycorrhizal infection was greater at higher levels of FMC in the media. Improved phosphorus uptake by inoculated severinia plants appeared at least partially responsible for increased growth compared to non-VAM plants under conditions of high soluble salts and pH associated with high FMC composition. Florida Agr. Expt. Sta. Journal Series No. 6319.     

Effectiveness of VAM fungi in nonsterile soils before and after optimization of P in soil solution

Five tropical soils were either not inoculated or inoculated with the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus aggregatum. The degree to which VAM effectiveness was expressed in the soils was evaluated prior and after solution P status was adjusted for optimal VAM activity. VAM effectiveness determined by monitoring P concentrations of pinnules of Leucaena leucocephala leaves as a function of time and as dry matter yield determined at the time of harvest, indicated that in three of the soils VAM effectiveness was either very restricted or altogether unexpressed irrespective of vesicular-arbuscular mycorrhizal fungal (VAMF) inoculation if soil solution P was not optimized for VAM effectiveness. After P optimization, effectiveness was significantly increased by VAMF inoculation although in four of the soils, densities of indigenous VAMF propagules greatly exceeded that attained by the inoculum after it was mixed with soil. Mycorrhizal fungal inoculation effects varied from soil to soil, depending on the extent to which the effectiveness of indigenous and introduced endophytes was enhanced by P optimization and the similarity of inherent soil solution P concentrations to the range known to be optimum for VAM effectiveness. Of the indicator variables monitored, VAMF colonization was least sensitive to treatment effects followed by shoot P concentration measured at the time of harvest.Contribution from Hawaii Institute of Tropical Agriculture and Human Resources Journal series No. 3781.Contribution from Hawaii Institute of Tropical Agriculture and Human Resources Journal series No. 3781.     

Vesicular-arbuscular mycorrhizal effectiveness in an acid soil amended with fresh organic matter

A greenhouse investigation was undertaken to determine the influence of fresh organic matter on the formation and functioning of vesicular-arbuscular mycorrhizal symbiosis in Leucaena leucocephala grown in an acid aluminum-rich ultisol. In soil not amended with fresh organic matter or lime, plants failed to grow. Mycorrhizal infection level, mycorrhizal effectiveness measured in terms of pinnule P content of L. leucocephala leaves and dry matter yield of the legume increased with increase in fresh organic matter. Although VAM colonization level and dry matter yield of L. leucocephala were significantly higher if the test soil was limed (7.2 cmole OH^–) than if amended with fresh organic matter, the latter was as effective as lime in off-setting the detrimental effect of aluminum on mycorrhizal effectiveness. The lower mycorrhizal colonization level and the lower dry matter yield noted in the soil treated with fresh organic matter appears to be related to the inadequacy of Ca in the soil amended with fresh organic matter. These observations are supported by the low calcium status of soil and plant tissues in the absence of lime. It is concluded that while fresh organic matter, in appropriate amounts, could protect sensitive plants and VAM symbiosis against Al toxicity in acid soils, maximum mycorrhizal inoculation effects are not likely to be attained unless the soils are also amended with Ca.Contribution from Hawaii Institute of Tropical Agriculture and Human Resources Journal Series No 3740.     

Use of soil transfer for reforestation on abandoned mined lands in Alaska

Soil transfers from an intermediate successional site and a mature forest site were applied to Populus balsamifera L. cuttings and Alnus crispa (Ait.) Pursh seedlings placed on an abandoned mined site in south central Alaska to improve plant establishment. Mycorrhizal fungi in the soil transfers from the two successional stages were hypothesized to have different effects on plant species that colonize disturbed sites at different times or on different substrates. The site consisted of coarse, dry, low-nutrient spoils and was naturally colonized by scattered P. balsamifera but not A. crispa, although seed sources for both were adjacent to the site. Physical dimensions of the transplanted seedlings and cuttings were measured at the beginning and end of each growing season. Selected plants were harvested at the end of the 2-year study and examined for mycorrhizal formation, current growth, and leaf tissue nutrient concentrations. Both plant species were taller when treated with the soil transfers from the mature forest than with soils from the intermediate site although the increase for A. crispa was greater. Physical dimensions, current growth, and nutrient concentrations were greater when A. crispa was treated with the mature soil transfer compared with the intermediate soil transfer. Mycorrhizae which infected Alnus were predominantly a brown woody type, while other types accounted for greater relative mycorrhizal infection percentage on Populus. Insufficient quantities of mycorrhizal inoculum of suitable species, as well as low moisture and low nutrient conditions, may be factors limiting A. crispa colonization on primary disturbed sites in south central Alaska.     

Effects of belowground grazing by collembola on growth,mycorrhizal infection,and P uptake of Geranium robertianum

We hypothesized that the grazing of vesicular-arbuscular mycorrhizal (VAM) hyphae by soil animals could be responsible for the lack of a direct relationship between mycorrhizal infection intensity and nutrient uptake under field conditions. To test this hypothesis, we determined the effect of a range of densities of the collembola, Folsomia candida, on growth, VAM infection, and P uptake in Geranium robertianum, a common forest herb, under greenhouse conditions. Total and aboveground growth were greater at low collembola density than either at higher collembola density or without collembola. These differences were greater when the plants were grown in a high organic content soil mix than when grown in sand. Root mass was not affected by collembola density. In the soil mix, root length decreased with increasing collembola density, but not in the sand. The percent of root length infected with VAM was lower at any collembola density than when collembola were absent. Total infected root length decreased linearly with increasing collembola density. Few significant differences in P uptake or tissue concentration were found. Thus, plant growth (but not P uptake) may be stimulated at low collembola density and inhibited at high. We discuss mechanisms which may be responsible for this non-linear response, and the implications of the pattern of response to studies of plant competition, nutrient turnover, and revegetation.     

Effect of tillage and farming system upon VAM fungus populations and mycorrhizas and nutrient uptake of maize

Low-input agricultural systems that do not rely on fertilizers may be more dependent on vesicular-arbuscular mycorrhizal [VAM] fungi than conventionally managed systems. We studied populations of spores of VAM fungi, mycorrhiza formation and nutrient utilization of maize (Zea mays L.) grown in moldboard plowed, chisel-disked or no-tilled soil under conventional and low-input agricultural systems. Maize shoots and roots were collected at four growth stages. Soils under low-input management had higher VAM fungus spore populations than soils under conventional management. Spore populations and colonization of maize roots by VAM fungi were higher in no-tilled than in moldboard plowed or chisel-disked soil. The inoculum potential of soil collected in the autumn was greater for no-till and chisel-disked soils than for moldboard plowed soils and greater for low-input than conventionally farmed soil. The effects of tillage and farming system on N uptake and utilization varied with growth stage of the maize plants. The effect of farming system on P use efficiency was significant at the vegetative stages only, with higher efficiencies in plants under low-input management. The effect of tillage was consistent through all growth stages, with higher P use efficiencies in plants under moldboard plow and chisel-disk than under no-till. Plants grown in no-tilled soils had the highest shoot P concentrations throughout the experiment. This benefit of enhanced VAM fungus colonization, particularly in the low-input system in the absence of effective weed control and with likely lower soil temperatures, did not translate into enhanced growth and yield.     

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.     

A field study using the fungicide benomyl to investigate the effect of mycorrhizal fungi on plant fitness

Summary The effect of vesicular-arbuscular mycorrhiza (VAM) on the fecundity ofVulpia ciliata ssp.ambigua was investigated at two field sites in eastern England by applying the fungicide benomyl to reduce VAM infection. The application of benomyl at the two sites produced very different results. At one site the application of the fungicide reduced the fecundity of plants whereas at the other fecundity was increased. At the first site the reduction in fecundity was linked to a significant reduction in VAM infection on the sprayed plants. The mechanism of the benefit associated with the VAM infection is however unclear: there was no treatment effect on morphology or on phosphorus inflow. At the second site, where fecundity was increased, there was only a negligible amount of VAM infection amongst the unsprayed plants and it is suggested that the increase in fecundity with the application of benomyl may have resulted from a reduction in infection by other, presumably pathogenic, fungi. The value of VAM fungi to the host plant may therefore not be restricted to physiological benefits. They may also provide protection to the plant by competing for space with other species of pathogenic fungi.     

Potentials and limitations of mycorrhizal inoculation illustrated by experiments with field-grown cassava

Summary Field inoculation trials with cassava (Manihot esculenta Crantz) were conducted in Quilichao (typic Dystropept soil) and Carimagua (Haplustox soil). In Quilichao, with a large and effective native VA-mycorrhizal (VAM) population, inoculation withGlomus manihotis did not increase cassava yields significantly, neither when different sources and levels of inoculum material were used, nor with different cassava cultivars, or after stabilizing soil temperature through mulching. Field inoculation did result in a decrease of the coefficient of variation with respect to yield. The high dependency of cassava on an effective VAM association was indicated by a marked decrease in yield after eradication of native VAM by soil sterilization. In Carimagua, with a lower native VAM population, mycorrhizal inoculation withG. manihotis increased yields significantly at intermediate levels of 100 kg/ha of applied P, using either inoculum of cassava orPanicum maximum roots or inoculum of a soil-root mixture of maize or tropical kudzu. Higher or lower levels of P decreased the effect of inoculation on yield. There were no significant differences among P sources, ranging from highly soluble triple superphosphate to low solubility rock phosphates. Inoculation with different VAM isolates had a variable effect on cassava yields, and showed that there may be an interaction between P fertilizer level and isolate efficiency. It is concluded that there may be a potential to increase yields or decrease the fertilizer P requirements of cassava through field inoculation with effective VAM isolates, in the vast areas of acid infertile Oxisols and Ultisols with low native VAM fungal populations, represented by Carimagua.     

Controls for rhizosphere microorganisms to study effects of vesicular-arbuscular mycorrhizae on Artemisia tridentata

Seven treatments were set up to test the effects of vesicular-arbuscular (VA) mycorrhizal fungi and other rhizosphere microorganisms on the growth of Artemisia tridentata ssp. tridentata. Soil sievings had no significant effect on root or shoot mass. Spores and surface-sterile spores were a poor inoculum source, but roots and fresh soil caused 45–75% mycorrhizal infection. Whereas root-inoculated plants still had low growth responses by the end of the experiment, fresh soil inoculum caused the greatest response, and partial fresh inoculum caused a lesser response. These results suggest that fresh soil is an appropriate inoculum for this plant-fungal-soil system, and that the major effect on plant growth of the fresh soil inoculum is from the mycorrhizal fungi and not from the other microorganisms, because the sievings had no effect on plant growth. In addition, soil dilution plating of saprophytic fungi showed 85% species similarity between sterile and fresh soil inoculum by the end of the experiment. Since the effects of non-VA microorganisms are complex and varied, we suggest that researchers work out the type of mycorrhizal controls that best suit their system.