Nodulated N2-fixing Casuarina cunninghamiana is the sink for net N transfer from non-N2-fixing Eucalyptus maculata via an ectomycorrhizal fungus Pisolithus sp. using 15NH4+ or 15NO3- supplied as ammonium nitrate

To determine the effects of nitrogen source on rates of net N transfer between plants connected by a common mycorrhizal network, we measured transfer of N supplied as 15NH4 14NO3 or 14NH4 15NO3 in three Casuarina/Eucalyptus treatments interconnected by a Pisolithus sp. The treatments were nonnodulated nonmycorrhizal/nonmycorrhizal; nonnodulated mycorrhizal/mycorrhizal; and nodulated mycorrhizal/mycorrhizal. Mycorrhization was 67% in Eucalyptus and 36% in Casuarina. N2 fixation supplied 38% of the N in Casuarina. Biomass, N and 15N contents were lowest in nonmycorrhizal plants and greatest in plants in the nodulated/mycorrhizal treatment. Nitrogen transfer was enhanced by mycorrhization and by nodulation, and was greater when N was supplied as 15NH4+ than 15NO3-. Nitrogen transfer rates were lowest in the nonmycorrhizal treatment for either 15N source, and greatest in the nodulated, mycorrhizal treatment. Transfer was greater to Casuarina than to Eucalyptus and where ammonium rather than nitrate was the N source. Irrespective of 15N source and of whether Casuarina or Eucalyptus was the N sink, net N transfer was low and was similar in both nonnodulated treatments. However, when Casuarina was the N sink in the nodulated, mycorrhizal treatment, net N transfer was much greater with 15NH4+ than with 15NO3-. High N demand by Casuarina resulted in greater net N transfer from the less N-demanding Eucalyptus. Net transfer of N from a non-N2-fixing to an N2-fixing plant may reflect the very high N demand of N2-fixing species.     

Distribution of arbuscular mycorrhizal fungi in marshy and shoreline vegetation of Deepar Beel Ramsar Site of Assam,India

Arbuscular mycorrhizal fungal (AMF) symbiosis was thought to be rare in wetland plant roots, although several recent studies suggested that this association might be important in wetland ecosystems. In this research work we have studied the distribution of AMF in the marshy and shoreline vegetation of Deepar Beel Ramsar site of Assam, India. The study reveals the percentage of mycorrhizal colonization in the roots of different plant species which were observed from 20.89 to 86.47% and particularly found larger among the members of the family Poaceae. The Vetiveria zizanioides L. from the family Cyperaceae showed the highest (86.47%) percentage of root colonization, however, only one plant species viz. Scirpus lateriflorus Gmel. from the same family was found to be nonmycorrhizal. The rhizospheric soil samples of most of the plant species were found to be dominated by Glomus morphotypes. All total 18 AMF morphotypes were recorded which comprises four genera viz. Glomus (66.67%), Acaulospora (16.66%), Gigaspora (11.11%) and Scutellospora (5.56%). The observation of diversity of AMF in 25 different plant species among the wetland plants gives a glimpse of AMF diversity and their host selectivity in the said ecosystem.     

Benefit and cost analysis and phosphorus efficiency of VA mycorrhizal fungi colonizations with sorghum (Sorghum bicolor) genotypes grown at varied phosphorus levels

Sorghum [Sorghum bicolor (L.) Moench] was grown in a greenhouse in a low P (3.6 mg kg^-1) soil (Typic Argiudolls) inoculated with the vesicular-arbuscular mycorrhizal fungi (VMAF) Glomus fasciculatum and P added at 0, 12.5, 25.0, and 37.5 mg kg^-1 soil to determine the effects of VAMF-root associations on plant growth, benefit and cost analysis, and P efficiency (dry matter produced/unit P absorbed). Root colonization with VAMF and shoot growth enhancements decreased with increased soil P applications. Mycorrhizal plants were less P efficient than nonmycorrhizal plants. Shoot dry matter differences between mycorrhizal and nonmycorrhizal plants were considered the benefit derived by plants from VAMF-root associations. Shoot dry matter differences between mycorrhizal and nonmycorrhizal plants with similar P concentrations were considered the costs paid by plants for VAMF-root associations. Values of benefit and cost analysis for VAMF-root associations were highest when soil P was lowest and decreased with increasing P applications. Genotypic differences for calculated costs were pronounced, but not benefits. Benefit and cost analysis.may be helpful to evaluate host plant genotypes and VAMF species to optimize efficiencies of VAMF symbiosis in different soil environments.     

Occurrence of arbuscular mycorrhizal fungi in a phosphorus-poor wetland and mycorrhizal response to phosphorus fertilization

The presence of arbuscular mycorrhizas in fens has received little attention, but because fen plants are often phosphorus limited, the plant-fungus interaction could be an important factor in plant competition for phosphorus. In this field study, we determined mycorrhizal colonization rates for 18 fen plant species. Also in the field, we examined the effect of four different forms of phosphorus on the percentage colonization for one fen plant species, Solidago patula. We found that in a species-rich, phosphorus-poor wetland both mycorrhizal and nonmycorrhizal species were common. Nine of ten dicotyledonous species examined formed arbuscular mycorrhizas, while all monocotyledonous species were at most very weakly mycorrhizal. A morphological explanation for this pattern is that the monocots in our study have more extensive aerenchyma, especially in coarse roots. Therefore, monocots are able to transport oxygen to their roots more effectively than dicots. In the organic wetland soil, additional oxygen in the rhizosphere promotes phosphorus mineralization and availability. Two of the monocot species (Typha latifolia and Carex lasiocarpa), which have been described previously as mycorrhizal in other wetland types, are surprisingly nonmycorrhizal in our phosphorus-poor study site, suggesting that a mycorrhizal association would not offer improved phosphorus nutrition to these species. In contrast, our field phosphorus addition decreased mycorrhizal colonization in S. patula, suggesting that one benefit to S. patula of the mycorrhizas is phosphorus uptake.     

The influence of arbuscular mycorrhizae and light on Wisconsin (USA) sand savanna understories 2. Plant competition

Wisconsin (USA) oak savannas are endangered plant communities that have remarkably high plant species diversity. To investigate factors underlying this richness, we experimentally investigated the potentially interacting effects of light gradients and arbuscular mycorrhizal fungi (AMF) on plant competition in the greenhouse, using a fully randomized block design. We used four plant species, soil, and AMF from a remnant sand savanna, under two light and five AMF treatments. Plants were grown four per pot under two competition treatments (either one or four species per pot) for 20 weeks. Using ANOVA, we found that all species showed significant treatment effects on total and shoot biomass, primarily due to differences in competition and light, less to AMF. However, effects were the opposite of predictions. Putatively mycorrhizal plants showed neutral to negative responses to AMF, and a nonmycorrhizal species outcompeted AMF species in infected pots. We concluded that our experimental setup of small pots, sandy soil, and long growing period had induced parasitism by the AMF on susceptible hosts. This unexpected result is consistent with field data from the sand savanna, and may help explain how nonmycorrhizal plants can compete successfully with AMF species in established, species-rich communities.     

Influence of Mycorrhizal Fungus, Phosphorus,and Burrowing Nematode Interactions on Growth of Rough Lemon Citrus Seedlings

Rough lemon seedlings were grown in mycorrhizal-infested or phosphorus-amended soil (25 and 300 mg P/kg) in greenhouse experiments. Plants Were inoculated with the citrus burrowing nematode, Radopholus citrophilus (0, 50, 100, or 200 nematodes per pot). Six months later, mycorrhizal plants and nonmycorrhizal, high-P plants had larger shoot and root weights than did non-mycorrhizal, low-P plants. Burrowing nematode population densities were lower in roots of mycorrhizal or nonmycorrhizal, high-P plants than in roots of nonmycorrhizal, low-P plants; however, differences in plant growth between mycorrhizal and nonmycorrhizal plants were not significant with respect to initial nematode inoculum densities. Phosphorus content in leaf tissue was significantly greater in mycorrhizal and nonmycorrhizal, high-P plants compared with nonmycorrhizal, low-P plants. Nutrient concentrations of K, Mg, and Zn were unaffected by nematode parasitism, whereas P, Ca, Fe, and Mn were less in nematode-infected plants. Enhanced growth associated with root colonization by the mycorrhizal fungus appeared to result from improved P nutrition and not antagonism between the fungus and the nematode.     

Mycorrhizal status of herb-layer plants in a fertilized oak-pine forest

In the years 1986–1988 investigations were carried out on the influence of fertilizers on the mycorrhizal status of herb-layer plants of a mixed oak-pine forest in the Niepolomice Forest (Southern Poland). The site enrichment with N, P, K altered the frequency of mycorrhizal infection in most species investigated and decreased the percentage of colonized root cells. Some species disappear after fertilization (e.g. Vaccinium myrtillus L.). Among the species vigorously expanding after the treatment, mainly nonmycorrhizal plants (e.g. Festuca gigantea (L.) Vill.) were found. Some species (e.g. Milium effusum L.) favourably reacting to mineral fertilization, lost their mycorrhizae.     

Mycorrhizal associations in the Patagonian steppe,Argentina

The mycorrhizal status of plant species in north-west Patagonia was examined. Communities representative of Patagonian steppe and marshes were compared with respect to the mycorrhizal status of their species.Most of both native and exotic plant species sampled were arbuscular mycorrhizal (AM). The percentage of species with mycorrhizal association was higher for perennial herbs and shrubs than for annual herbs. The higher ratio of mycorrhizal/nonmycorrhizal (NM) species found for dicotyledons than for monocotyledons, could reflect the presence of a considerable number of NM monocotyledons in the marsh. The mycorrhizal status of plants differed slightly between the steppe and the marsh. In the steppe, native AM species were more frequent than in the marsh. In contrast, in the marsh, the NM species were proportionally more represented than in the steppe. The Juncaceae and Cyperaceae, which include hydrohytic NM plants, accounted for many of these differences. Moreover, the dominant species in the marshes, Juncus arcticus, is a NM species.In the present study, most of species belonging to the same taxonomic family tended to have the same mycorrhizal associations, in agreement with studies on plants from other regions. Exceptions to this general behaviour were observed in the families Cyperaceae, Scrophulariaceae, Berberidaceae and Amaryllidaceae. The most represented families in which mycorrhizal behaviour differed between species of the same family were Asteraceae, Fabaceae and Poaceae. Senecio neaei (Asteraceae) and Boopis australis (Calyceraceae) showed facultative mycorrhizal behaviour.     

Response to ammonium and nitrate by a mycorrhizal annual invasive grass and native shrub in southern California

The goal of this study was to determine the interaction of mycorrhizae and two N sources, ammonium (NH(4)(+)) and nitrate (NO(3)(-)), on the growth of a coastal sage scrub (CSS) species, Artemisia californica, and an exotic annual grass, Bromus madritensis ssp. rubens. Anthropogenic nitrogen deposition may be influencing the decline of CSS and replacement by exotic grasses, but the extent to which mycorrhizae are involved in shrubland decline is unknown. NO(3)(-) is the dominant form of deposition in southern California, although the native, uneutrophied soils have a greater concentration of NH(4)(+). Seeds of each species were germinated in pots of sterile soil, inoculated with native soil containing mycorrhizal spores and infective root fragments, and fertilized with 50 μg/g of either NO(3)(-) or NH(4)(+). NH(4)(+) enhanced the growth of both mycorrhizal species, while NO(3)(-) did not. Control plants of B. madritensis under low N had a significant response to mycorrhizae, but A. californica did not. Nitrate increased the growth of nonmycorrhizal A. californica as much as the mycorrhizal NH(4)(+)-treated plants. There is no evidence in this study to suggest that the decline of A. californica or increase in B. madritensis is due to a mycorrhizal response to NO(3)(-). Other life history traits of the two species must be used to explain the invasive behavior of the annual grass. Mycorrhizae may be more important in controlling plant growth in native uneutrophied soils dominated by NH(4)(+) rather than NO(3)(-).     

Mycorrhizal status of the genus Carex (Cyperaceae)

The Cyperaceae have generally been considered nonmycorrhizal, although recent evidence suggests that mycotrophy may be considerably more widespread among sedges than was previously realized. This study surveyed 23 species of Carex occurring in upland and wetland habitats in northeastern Illinois. Mycorrhizal infection by arbuscular fungi was found in the roots of 16 species of Carex and appears to occur in response to many factors, both environmental and phylogenetic. While some species appear to be obligately nonmycorrhizal, edaphic influences may be responsible for infection in others. In five of the seven Carex species that were nonmycorrhizal, a novel root character, the presence of bulbous-based root hairs, was identified. The taxonomically patchy distribution of the distinctive root hair trait suggests that these structures may have evolved several times within the genus. Evidence of multiple independent origins of the root hair trait lends support to the hypothesis that root hairs represent an adaptation to nonmycotrophy. Although taxonomic position does seem to be of importance in determining the mycorrhizal dependence of sedges, the pattern may be a patchwork of both mycorrhizal clades and clades that have adapted to the nonmycorrhizal state.     

Field responses to added organic matter, arbuscular mycorrhizal fungi, and fertilizer in reclamation of taconite iron ore tailing

A three season study was conducted to determine the effect of added composted yard waste, arbuscular mycorrhizal (AM) fungi, and fertilizer on plant cover, standing crop biomass, species composition, AM fungal infectivity and spore density in coarse taconite iron ore tailing plots seeded with a mixture of native prairie grasses. Plant cover and biomass, percent seeded species, mycorrhizal infectivity and spore density were greatly increased by additions of composted yard waste. After three seasons, total plant cover was also greater in plots with added fertilizer. Third season plant cover was also greater in plots amended with the higher rate (44.8 Mg ha^{–1) of compost than the moderate rate (22.4 Mg ha-1}). Field inoculation with AM fungi also increased plant cover during the second season and infectivity during the first two seasons. Seeded native species, consisting mostly of the cover species Elymus canadensis, dominated plot vegetation during the second and third seasons. Dispersal of AM fungal propagules into nonmycorrhizal plots occurred rapidly and increased infectivity in compost-amended plots during the third season. In plots with less than 10% plant cover, AM fungal infectivity of inoculated plots was greatly reduced after the second season. The high level of plant cover and the trend of increasing proportion of mycorrhizal-dependent warm-season grasses, along with increases in infectivity, forecast the establishment of a sustainable native grass community that will meet reclamation goals.     

Benefit, cost and water-use efficiency of arbuscular mycorrhizal durum wheat grown under drought stress

 Arbuscular mycorrhizal fungi (AMF) living symbiotically with host plants enhance plant growth by improving the acquisition of mineral nutrients and water relations. This study determined the effects of AMF inoculation on growth, benefit/cost and water-use efficiency (grams dry matter produced per kilogram water evapotranspired) in two durum wheat genotypes (drought sensitive and drought tolerant) under water-stressed and well-watered conditions. Plants were grown in a low-P silty clay (Typic Xerochrept) soil mix in a greenhouse. Shoot and root dry matter (DM) and root AMF colonization were higher for well-watered than for water-stressed plants. The mycorrhizal plants were more water-use efficient than nonmycorrhizal plants. Shoot DM differences between mycorrhizal and nonmycorrhizal plants represent the benefit derived by plants from AMF-root associations. Shoot DM differences between mycorrhizal and nonmycorrhizal plants under similar conditions of water treatment represent the cost to the plant of AMF-root associations. Values of benefit/cost for AMF-root associations were highest when plants were water-stressed and decreased under well-watered conditions. Genotypic differences in calculated costs and benefits were pronounced. Benefit/cost analysis may be helpful in evaluating host plant genotypes in order to optimize efficiencies of AMF symbiosis under different environmental conditions. Accepted: 4 April 1998     

Phosphorus nutrition-mediated effects of arbuscular mycorrhiza on leaf morphology and carbon allocation in perennial ryegrass

The aim of this work was to disentangle phosphorus status-dependent and -independent effects of arbuscular mycorrhizal fungus (AMF) on leaf morphology and carbon allocation in perennial ryegrass (Lolium perenne). To this end, we assessed the P-response function of morphological components in mycorrhizal and nonmycorrhizal plants of similar size. AMF (Glomus hoi) stimulated relative P-uptake rate, decreased leaf mass per area (LMA), and increased shoot mass ratio at low P supply. Lower LMA was caused by both decreased tissue density and thickness. Variation in tissue density was almost entirely caused by variations in soluble C, while that in thickness involved structural changes. All effects of AMF were indistinguishable from those mediated by increases in relative P-uptake rate through higher P-supply rates. Thus the relationships between relative P-uptake rate, leaf morphology and C allocation were identical in mycorrhizal and nonmycorrhizal plants. No evidence was found for AMF effects not mediated by changes in plant P status.     

Growth responses of tropical forage plant species to vesicular-arbuscular mycorrhizae

Summary Growth and mineral uptake of twenty-four tropical forage legumes and grasses were compared under glasshouse conditions in a sterile low P oxisol, one part inoculated and the other not inoculated with mycorrhizal fungi. Shoot and root dry weights and total uptake of P, N, K, Ca, and Mg of all the test plants were significantly increased by mycorrhizal inoculation. Mycorrhizal inoculation, with few exceptions, decreased the root/shoot ratio. Non-mycorrhizal plants contained always lower quantities of mineral elements than mycorrhizal plants. Plant species showed differences in percentage mycorrhizal root length and there was no correlation between percentage mycorrhizal infection and plant growth parameters. A great variation in dependence on mycorrhiza was observed among forage species. Total uptake of all elements by non-mycorrhizal legumes and uptake of P, N and K by non-mycorrhizal grasses correlated inversely with mycorrhizal dependency. Mycorrhizal plants of all species used significantly greater quantities of soil P than the nonmycorrhizal plants. Utilization of soil P by non-mycorrhizal plants was correlated inversely with mycorrhizal dependency.     

Mycorrhizal Status Influences the Rate but not the Temperature Sensitivity of Soil Respiration

Mycorrhizal fungi, which can produce a large portion of total soil respiration, respond strongly to global changes such as elevated CO₂, N-deposition, and land-use change. Predictions of future ecosystem C sequestration hinge on respiration budgets, but the mycorrhizal influence on total soil respiration remains unknown. In this study, sunflowers (Helianthus annuus) were subjected to various mycorrhizal treatments, and their root and soil systems were enclosed in chambers that continuously monitored belowground (root + mycorrhizal + heterotrophic) CO₂ production during plant growth, death, and decomposition. Rhizocosms with high mycorrhizal colonization exhibited higher soil respiration rates as plants matured, an increase that was in proportion to the mycorrhizal stimulation of plant growth. Living mycorrhizal plants behaved like nonmycorrhizal ones in that total rhizocosm respiration had the same relationship to plant mass and the same temperature sensitivity as nonmycorrhizal plants. Upon removal of the shoots though, mycorrhizal plants exhibited the largest relative reduction in respiration resulting in a unique relationship of soil respiration with plant mass. The mycorrhizal influence on heterotrophic respiration merits as much attention from experimenters and modelers as the mycorrhizal contribution to autotrophic respiration.     

Early events in the life of apple roots: variation in root growth rate is linked to mycorrhizal and nonmycorrhizal fungal colonization

Early events of mycorrhizal and nonmycorrhizal fungal colonization in newly-emerging roots of mature apple (Malus domestica Borkh) trees were characterized to determine the relationship of these events to fine root growth rate and development. New roots were traced on root windows to measure growth and then collected and stained to quantify microscopically the presence of mycorrhizal and nonmycorrhizal fungal structures. Most new roots were colonized by either mycorrhizal or nonmycorrhizal fungi but none less 25 days old were ever internally colonized by both. Compared to nonmycorrhizal colonization, mycorrhizal colonization was associated with faster growing roots and roots that grew for a longer duration, leading to longer roots. While either type of fungi was observed in roots as soon as 3 days after root emergence, intraradical colonization by mycorrhizal fungi was generally faster (peaking at 7 to 15 days) than that by nonmycorrhizal fungi and often occurred more frequently in younger roots. Only 15 to 35% of the roots had no fungal colonization by 30 days after emergence. This study provides the first detailed examination of the early daily events of mycorrhizal and nonmycorrhizal fungal colonization in newly emerging roots under field conditions. We observed marked discrimination of roots between mycorrhizal and nonmycorrhizal fungi and provide evidence that mycorrhizal fungi may select for faster growing roots and possibly influence the duration of root growth by non-nutritional means.     

THE ROLE OF ENDOMYCORRHIZAE IN REVEGETATION PRACTICES IN THE SEMI-ARID WEST. I. A COMPARISON OF INCIDENCE OF MYCORRHIZAE IN SEVERELY DISTURBED VS. NATURAL ENVIRONMENTS

A comparison of a natural, undisturbed ecosystem, a mid-elevation sage community, with a severely disturbed old roadbed through this community revealed that more than 99% of the plant cover in the natural community was mycorrhizal (vesicular-arbuscular), whereas less than 1% of the plant cover in the disturbed area (roadbed) was mycorrhizal. Examples of nonmycorrhizal plants as primary successional species in severely disturbed habitats are discussed. The importance of maintaining or re-establishing the mycorrhizal fungal component in reclamation programs designed to produce stable ecosystems is emphasized.     

Behavior of mercury in a soil–plant system as affected by inoculation with the arbuscular mycorrhizal fungus Glomus mosseae

Effects of inoculation with the arbuscular mycorrhizal (AM) fungus Glomus mosseae on the behavior of Hg in soil–plant system were investigated using an artificially contaminated soil at the concentrations of 0, 1.0, 2.0, and 4.0 mg Hg kg⁻¹. Mercury accumulation was lower in mycorrhizal roots than in nonmycorrhizal roots when Hg was added at the rates of 2.0 and 4.0 mg kg⁻¹, while no obvious difference in shoot Hg concentration was found between mycorrhizal and nonmycorrhizal treatments. Mycorrhizal inoculation significantly decreased the total and extractable Hg concentrations in soil as well as the ratio of extractable to total Hg in soil. Equilibration sorption of Hg by soil was investigated, and the results indicated that mycorrhizal treatment enhanced Hg sorption on soil. The uptake of Hg was lower by mycorrhizal roots than by nonmycorrhizal roots. These experiments provide further evidence for the role of mycorrhizal inoculation in increasing immobilization of Hg in soil and reducing the uptake of Hg by roots. Calculation on mass balance of Hg in soil suggests the presence of Hg loss from soil presumably through evaporation, and AM inoculation enhanced Hg evaporation. This was evidenced by a chamber study to detect the Hg evaporated from soil.     

Effects of mycorrhizal infection on drought tolerance and recovery in safflower and wheat

The influence of arbuscular mycorrhizal fungi on drought tolerance and recovery was studied in safflower (Carthamus tinctorius L.) and wheat (Triticum aestivum L.). Plants were grown with and without the mycorrhizal fungus, Glomus etunicatum Becker & Gerd., in nutrient-amended soil under environmentally-controlled conditions to yield mycorrhizal and nonmycorrhizal with similar leaf areas, root length densities, dry weights, and adequate tissue phosphorus. When drought stress was induced, mycorrhizal infection did not affect changes in leaf water, osmotic or pressure potentials, or osmotic potentials of leaf tissue rehydrated to full turgor in either safflower or wheat. Furthermore, in safflower, infection had little effect on drought tolerance as indicated by the level of leaf necrosis. Mycorrhizal wheat plants, however, had less necrotic leaf tissue than uninfected plants at moderate levels of drought stress (but not at severe levels) probably due to enhanced phosphorus nutrition. To determine the effects of infection on drought recovery, plants were rewatered at a range of soil water potentials from –1 to –4 MPa. We found that although safflower tended to recover more slowly from drought after rewatering than wheat, mycorrhizal infection did not directly affect drought recovery in either plant species. Daily water use after rewatering was reduced and was correlated to the extent that leaves were damaged by drought stress in both plant species, but was not directly influenced by the mycorrhizal status of the plants.     

Growth and Water Relations in Mycorrhizal and Nonmycorrhizal Pinus Halepensis Plants in Response to Drought

Mycorrhizal and nonmycorrhizal Pinus halepensis plants were subjected to water stress by withholding irrigation for four months and then rehydrated for 30 d. Water stress affected plants growth and mycorrhizal association was unable to avoid the effects of drought on plant growth. However, when irrigation was re-established the increase in height, number of shoots, total dry mass, and chlorophyll content in the mycorrhizal plants were greater than in non-mycorrhizal plants. The decrease in soil water content decreased the leaf water potential, leaf pressure potential and stomatal conductance. These decreases were higher for nonmycorrhizal than for mycorrhizal plants, indicating that the mycorrhizal fungi permit a higher water uptake from the dry soils. The total content of inorganic solutes was not changed by presence of mycorrhizae.