Effects of microtopography on mycorrhizal infection in Atlantic white cedar (Chamaecyparis thyoides (L.) Mills.)

 The presence and intensity of mycorrhizal infection in wetland trees has received relatively little attention. We report here a study of mycorrhizal infection in Atlantic white cedar (Chamaecyparis thyoides), a member of the Cupressaceae, which forms monospecific stands in swamps throughout the Atlantic Coastal Plain of North America. The trees grow on the tops of elevated hummocks, but the fine roots extend along the sides of the hummocks to the flooded hollows. Roots from all microtopographic positions on the hummocks are colonized by vesicular-arbuscular mycorrhizae (VAM). In addition to arbuscules and vesicles, occasional hyphal coils are observed within the cortex cells. However, frequencies of occurrence of vesicles and arbuscles are significantly higher on the tops and sides than at the bottoms of the hummocks. These differences correspond to higher concentrations of acetylglucosamine in the roots at these positions. Frequencies of all mycorrhizal structures (arbuscles, vesicles and hyphae) in roots at the base of the hummocks are very low. These results suggest that mycorrhizal colonization in wetland trees is greater in aerobic microsites, a finding in accord with results from studies of both herbaceous wetland plants and other wetland trees. Accepted: 5 September 1998     

VESICULAR-ARBUSCULAR MYCORRHIZAE IN HAWAIIAN ERICALES

Vesicular-arbuscular mycorrhizae (VAM) are reported for the first time in four species of Hawaiian Ericales, Vaccinium calycinum, V. dentatum, and V. reticulatum of the Ericaceae and Styphelia tameiameiae of the Epacridaceae. The coarse roots (> 1.5 mm diam) of many specimens were densely colonized by VAM fungi, with up to 90% of the length of roots containing arbuscules, vesicles, coils, and internal hyphae. Spores of an undescribed Glomus sp. were associated with two species of Vaccinium. The hair roots of all species bore the ericoid mycorrhizae typical of certain families of this order. The high frequency of VAM in Hawaiian populations of Ericales suggests that ancestral Ericales possessed the capacity to form both VA and ericoid mycorrhizae. An evolutionary sequence of mycorrhizal dependency in the Ericales is presented.     

A tripartite culture system for endomycorrhizal inoculation of micropropagated strawberry plantlets in vitro

The objective of the current investigation was to develop a reliable method to obtain vesicular arbuscular mycorrhizae (VAM) in micropropagated plantlets and to determine their influence on growth. An in vitro system for culturing the VA mycorrhizal fungus Glomus intraradices with Ri T-DNA-transformed carrot roots or nontransformed tomato roots was used in this study as a potential active source of inoculum for the colonization of micropropagated plantlets. After root induction, micropropagated plantlets grown on cellulose plugs (sorbarod) were placed in contact with the primary mycorrhizae in growth chambers enriched with 5000 ppm CO₂ and fed with a minimal medium. After 20 days of tripartite culture, all plantlets placed in contact with the primary symbiosis were colonized by the VAM fungus. As inoculum source, 30-day-old VA mycorrhizal transformed carrot roots had a substantially higher infection potential than 5-, 10-or 20-day-old VAM. Colonized plantlets had more extensive root systems and better shoot growth than control plants. The VAM symbiosis reduced the plantlet osmotic potential. This response may be a useful pre-adaptation for plantlets during transfer to the acclimatization stage.     

MYCORRHIZAE IN HAWAIIAN ANGIOSPERMS: A SURVEY WITH IMPLICATIONS FOR THE ORIGIN OF THE NATIVE FLORA

Endemic, indigenous, naturalized and cultivated angiosperms growing in a variety of habitats in Hawaii were examined for mycorrhizae. Of 147 species (representing 61 families) examined, 122 were mycorrhizal, and 98% of the latter formed vesicular-arbuscular mycorrhizae (VAM). Ericoid, orchid, and ectomycorrhizae also were found. Mycorrhizae were most frequent in endemic species (90.3%) and least in indigenous species (71.9%). The incidence of mycotrophy in Hawaiian endemic species was significantly greater than in mainland species. In addition, VAM often were found in species belonging to families that typically lack mycorrhizae (Aizoaceae, Brassicaceae, Caryophyllaceae, Cyperaceae, Hydrophyllaceae, Juncaceae, Pandanaceae, and Urticaceae). Naturalized weedy species were highly mycorrhizal. The high incidence of mycorrhizae in endemic species suggests that VAM fungi and obligately mycotrophic plant species may have been present in the Hawaiian Islands from the earliest stages of the development of the angiosperm flora.     

Vesicular-arbuscular mycorrhizae of epiphytes

This article introduces reports concerning the occurrence of mycorrhizae on epiphytes in Costa Rica, Ethiopia, Venezuela, Malaysia, and Mexico. Association of vesicular-arbuscular mycorrhizal fungi with the roots of epiphytes is not well known. Vesicular-arbuscular mycorrhizal fungi (VAM) do occur in the canopy, but are uncommon except in certain sites and host taxa. Occurrence of VAM on epiphytes may be constrained by mineral nutrient availability and spatial heterogeneity in the canopy. Nevertheless, epiphytes present unique opportunities to study influences of mycorrhizae on vascular plant community composition and on the evolution of mycorrhizal associations.     

Axenic culture and encapsulation of the intraradical forms of Glomus spp.

In recent years there have been many attempts to cultivate in vitro vesicular-arbuscular mycorrhizal (VAM) fungi which are obligate symbionts. Resting spores extracted from soils are often used as inoculum. Mycorrhizal root pieces are also used for inoculation but the role of intra-radical structures has not been clearly established. On agar medium vegetative mycelium was regenerated from individual intra-radical vesicles and from hyphae extracted by enzymatic maceration. After cell penetration, the mycelium probably accumulates substances which allow growth of VAM fungi in pure culture. When associated with tomato roots, this mycelium forms typical mycorrhizae. Encapsulation stabilized the biological properties of mycorrhizal roots and isolated vesicles. The immobilization also preserved the infectivity of the intra-radical hyphae and vesicles. After 25 years of exclusive utilization of resting spores as starting material for axenic and dual cultures of VAM fungi, it appears that intra-radical vesicles may be preferable propagules.D.-G. Strullu and C. Romand are with the Université d'Angers, Laboratoire de Phytonique, 2 Boulevard Lavoisier, 49045 Angers Cedex, France. C. Plenchette is with INRA, Station d'Agronomie, 17 rue Sully, 21034 Dijon Cedex, France.     

The topsoil as the major store of the propagules of vesicular-arbuscular mycorrhizal fungi in southeast Australian sandstone soils

The formation of vesicular-arbuscular mycorrhizae (VAM) in intact soil profiles from two sites in southeastern Australia were measured at two depths using a bioassay grown in intact soil cores. Intact soil cores were taken from (1) topsoil (0–15 cm) and (2) subsoil (15–30 cm) four times during 1990. Seeds of Acacialinifolia (Vent.) Willd. (Mimosaceae) were sown into the cores and plants harvested 8 and 12 weeks after sowing. For 1990, at both sites and in all seasons, VAM most readily developed in the roots of seedlings of A. linifolia grown in topsoil. Limited VAM occurred in roots grown in subsoil cores. Most colonisation of roots by VAM occurred from cores collected during spring and summer. Spore numbers were quantified for each site and depth by wet-sieving 100-g samples of air-dried soil and counting turgid spores containing oil droplets. Three types of spores were found in the soils. Few spores were extracted from all soils sampled, and for the most abundant of the spore types at least twice as many spores occurred in the topsoil than in the subsoil for all seasons examined. As most of the propagules that initiate VAM infection were observed in the topsoil, disturbances which involve the removal and storage of the top 15 cm will adversely affected these fungi.     

Mycorrhizae and their potential use in the agricultural and forestry industries

Mycorrhizal fungi associated with plant roots increase the absorption of nutrients, particularly phosphorus, and thus enhance the growth of crop plants and trees. Vesicular-arbuscular mycorrhizae (VAM) occur in approximately 90% of all vascular plants including most of the important agricultural species, whereas ectomycorrhizae are found in most of the economically important tree species of the temperate regions of the world, and in some tropical trees. These symbiotic associations are, therefore, important in crop and biomass production. For this reason they are receiving considerable attention in agriculture and forestry. Currently, VAM are utilized in fumigated soils, greenhouse crops, and in the reclamation of disturbed sites. Ectomycorrhizae are employed in the establishment of trees in nurseries, in reforestation programs, and in the production of containerized seedlings. Production of VAM and ectomycorrhiza inoculum for large scale projects is now feasible but many basic questions related to persistence of these fungi in field situations, competition with other microorganisms, and particularly the most efficient fungi to use for particular hosts remain largely unanswered.     

Occurrence of vesicular-arbuscular mycorrhizae in Pseudotsuga menziesii and Tsuga heterophylla seedlings grown in Oregon Coast Range soils

 Vesicular-arbuscular mycorrhizae (VAM) were common in seedlings of Pseudotsuga menziesii and Tsuga heterophylla grown in a greenhouse soil bioassay in soils collected from the Oregon Coast Range. Although root samples were heavily colonized by ectomycorrhizal fungi (EM), VAM colonization was observed in the cortical cells of both secondary and feeder roots. Vesicles, arbuscules, and hyphae typical of VAM occurred in 48% of 61 P. menziesii and 25% of 57 T. heterophylla seedlings. The ecological significance of VAM presence in the Pinaceae, as well as interactions among VAM, EM, and the plant host, deserve future investigation. Accepted: 16 August 1995     

Host genotype and the formation and function of VA mycorrhizae

VA mycorrhizae, the most ancient type of mycorrhizal symbiosis, are present in the most phytogenetically advanced groups. Few plants have evolved mechanisms to completely prevent infection by VAM fungi. Yet, plant species that are less dependent on VA mycorrhizae for nutrient acquisition (e.g., grasses) generally have less root colonization in the field than more dependent species (e.g., Citrus). Among closely related Citrus genotypes, there is a greater tendency for less dependent species to limit the rate but not the extent of colonization, even in high-P soils. We hypothesize that colonization represents a significant carbon cost that may be regulated by the host genotype. Carbon expenditure on the fungus at high P may result in mycorrhizal-induced growth depression. The potential value of breeding plants for greater susceptibility to colonization will depend on the cost/benefit of VA mycorrhizae for the specific crop, soil and environmental conditions. Although the genetics and physiology of host control over VAM colonization are barely known, recently discovered mycorrhizal colonization mutants (myc^-) of pea offer great promise for the study of host-fungus compatibility. Florida Agricultural Experiment Station Journal Series No. R-02765. Florida Agricultural Experiment Station Journal Series No. R-02765.     

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.     

Vesicular-arbuscular mycorrhizae in relation to plant disease

Vesicular-arbuscular mycorrhizae (VAM) enhance plant growth through increased nutrient uptake, stress tolerance and disease resistance. As an integral part of the root system, they interact with other microorganisms in soil and result in increased root exudation approaching about 25% of the plant dry matter production. Roots support a multitude of microorganisms that, in concert, can have profound influence on growth and survival of the plant. VAM fungi can alter the root exudation pattern, enhance chitinolytic activity and alter photosynthetic/respiratory deficiencies. VAM-positive plants are known to exhibit varied resistance towards soil-borne and foliar pathogens. The known interactions include a number of mechanisms, such as exclusion of the pathogen, lignification of plant cell walls, changed phosphate nutrition resulting in altered exudation by roots, and formation of inhibitory low molecular weight compounds. The purpose of this review is to discuss VAM-plant-pathogen interactions and the possible mechanisms involved in altered resistance. Based on these observations, a working model is proposed to explain the VAM-disease interaction under varied environmental conditions.     

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.     

Development of two endomycorrhizal symbioses on soybean and comparison with phosphorus fertilization

Summary Soybean (Glycine max L. Merr. cv. Amsoy 71) plants were grown in a greenhouse in a soil very low in plant-available P, and plants were harvested 5 times over a 21-week growth period. Soybeans were inoculated with one of two species of VAM fungi or received daily one of three nutrient solutions of different P concentrations (0.0, 0.2, or 1.0mMP). Until week 9, the dry weights, leaf areas and developmental stage of soybeans inoculated withG. fasciculatum orG. mosseae were similar to the 1.0 or 0.2mMP-treated plants, respectively. Phosphorus concentrations were significantly lower in VAM plants at weeks 6 and 9 as compared to non-VAM soybeans given 1.0mMP, suggesting P input in VAM plants was immediately used for new growth. Total P input for VAM plants was linear over 21 weeks, and the average rate of P uptake for these plants was 0.19mg P d⁻¹. Estimated specific P uptake rates (SPUR) for the mycorrhizae (VAM roots) were twice that of the control (0.0mMP) roots. The calculated SPURs forG. fasciculatum andG. mosseae hyphae were 95 and 120μg P g⁻¹ VAM d⁻¹ respectively, a 4 to 5 fold increase over non-inoculated roots, indicating more attention must be paid to P assimilation by VAM fungi in P-fixing substrates. Contribution from the Western Regional Research Center, USDA-ARS (CRIS No. 5325-20580-003).     

THE MYCORRHIZAE OF PIONEER SPECIES IN DISTURBED ECOSYSTEMS IN WESTERN PENNSYLVANIA

Study plots with ten types of disturbances were established in a Solidago-Aster community in western Pennsylvania. One year after disturbance, 93% of all species had vesicular-arbuscular mycorrhizae (VAM) and accounted for 92% of all plant cover. After 3 yr, 96% of the species had VAM and 96% canopy coverage. An adjacent 10-yr Solidago-Aster community had 100% VAM species. The VAM annuals, Ambrosia artemisiifolia and Setaria glauca, were major first year dominants while VAM perennials, Agropyron repens, Solidago canadensis, and Aster ericoides, dominated at 3 yr. The mean percent mycorrhizae of all dominants at 1 yr was 39% and 34% at 3 yr. Four disturbed sites had 100% VAM colonization after 1 yr and six sites after 3 yr. Severe physical disturbances (topsoil removed and topsoil-subsoil removed) resulted in complete colonization by VAM species and mean mycorrhizal infections of more than 75%. The plowed-disked-prometone and plowed-disked-vapam disturbances resulted in low levels of mycorrhizae after 3 yr. The plowed-disked-atrazine plot showed an initial severe decrease in percentage of mycorrhizae, but doubled its infection rate by 3 yr. Three non-mycorrhizal species (Chenopodium album, Rumex acetosella, and Polygonuspersicaria) were observed during the 3-yr period. C. album was the major dominant after 1 yr in the plowed-disked-fertilized site and R. acetosella a dominant in the plowed-disked-prometone plot.     

Influence of phosphorus and formononetin on isozyme expression in the Zea mays‐Glomus intraradices symbiosis

Maize (Zea mays L. cv. Great Lakes 586) plants were either inoculated with the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus intraradices Schenck and Smith, or grown in the presence of the isoflavone formononetin or were provided with both G. intraradices and formononetin. All plants were grown in soil containing one of five levels of inorganic P (between 8 and 110 µg g^?1 soil). By 3 weeks there were significant differences in a number of enzyme activities and in the pattern of isoenzymes in roots colonized by the VAM fungus or treated with formononetin. One NAD-malate dehydrogenase (MDH) isozyme was expressed only in mycorrhizal roots, whether treated or not with formononetin. Despite differences in the soil P level, the expression of this isozyme was not observed in non-mycorrhizal roots, indicating specific expression in the mycorrhizae. We suggest that MDH isozyme could serve as a specific, early indicator of the Zea-Glomus symbiosis. Differences in the esterase (EST) isozyme pattern were not detectable between VAM and non-VAM roots, suggesting that this enzyme system is not a good parameter for the evaluation of mycorrhizal colonization. As available P in the soil increased, total EST activity appeared to increase as well. Interestingly, total peroxidase (POX) activity increased along with P suggesting that as plant P nutrition improved, both cell wall ramification and the quantity of defense peroxidases increased as well. Total POX activity from mycorrhizal roots was inversely correlated with root colonization, indicating that there was suppression of POX activity by the host under low soil P. Most interestingly, formononetin further decreased POX activity regardless of the level of P or mycorrhizal status. This may suggest one mechanism by which formononetin enhances root VAM colonization. The presence of this isoflavone suppressed POX activity in mycorrhizal roots allowing a rapid penetration and spread of the fungus in the root cortex. The interplay between host root, soil P levels, secondary metabolites and endogenous host enzyme activities and a particular VAM fungus has a profound effect on the efficiency, duration and functioning of an endomycorrhizal symbiosis.     

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.     

Fungi in neotropical epiphyte roots

Roots of thirty-eight Ecuadoran vascular epiphytes, representing eleven angiosperm families, were examined for the presence of symbiotic microorganisms. Most orchid roots contained fungal endophytes like those that regularly infect terrestrial counterparts. Hyphae were also common in and on nonorchid roots, but assignments of these relationships to known mycorrhizal morphologies was not possible in all cases. Evidence of vesicular-arbuscular mycorrhizae (VAM) existed in a number of subjects while in Ericaceae and Campanulaceae a fungal association similar to the demateaceous surface fungi (DSF) described for alpine and prarie plants was usually present. Some associations were characterized by multicellular propagules on root surfaces. The significance of these findings and the factors likely to influence occurrence and consequences of root-fungus mutualisms in tropical forest canopies are discussed. Facts and considerations that could aid future inquiry on these systems are provided.     

Hyphal contribution to water uptake in mycorrhizal plants as affected by the fungal species and water status

Vesicular-arbuscular mycorrhizae may increase resistance of plants to drought by a number of mechanisms, such as increased root hydraulic conductivity, stomatal regulation, hyphal water uptake and osmotic adjustment. However, a substantial contribution of vesicular-arbuscular mycorrhizal (VAM) hyphae to water uptake has not been demonstrated unequivocally. The objective of this investigation was to examine the contribution of hyphae from two VAM fungi to water uptake and transport by the host plant. Lettuce (Lactuca sativa L.) plants were grown in a container divided by a screen into two compartments. One was occupied by roots, the other only by VAM hyphae, which the screen permitted to pass. Roots were colonized by the VAM fungi Glomus deserticola or Glomus fasciculatum, or were left uninoculated but P-supplemented. Water was supplied to the hyphal compartment at a distance of 10 cm from the screen (root). CO₂ exchange rate, water-use efficiency, transpiration, stomatal conductance and photosynthetic phosphorus-use efficiency of VAM or P-amended control plants were evaluated at three levels of water application in the hyphal compartment. Results indicate that much of the water was taken up by the hyphae in VAM plants. VAM plants, which had access to the hyphal compartment, had higher water and nutrient contents. G. deserticola functioned efficiently under water limitation and mycelium from G. fasciculatum-colonized plants was very sensitive to water in the medium. This discrepancy in VAM behaviour reflects the various abilities of each fungus according to soil water levels. Different abilities of specific mycelia were also expressed in terms of nutritional and leaf gas-exchange parameters. G. fasciculatum caused a significant increase in net photosynthesis and rate of water use efficiency compared to G. deserticola and P-fertilized plants. In contrast, the G. deserticola treatment was the most efficient affecting N, P and K nutrition, leaf conductance and transpiration. Since no differences in the intra- and extra-radical hyphal extension of the two endophytes were found, the results demonstrate that mycorrhizal hyphae can take up water and that there are considerable variations in both the behaviour of these two VAM fungi and in the mechanisms involved in their effects on plant water relations.     

香港次生林下植物菌根的调查

本文报道了香港６４种野生植物根系天然感染菌根菌的状况：９８．４％的植物都不同程度地感染上了菌根菌,９８．１％的根样发现受内生菌根（ＶＡＭ）的感染,其中小叶青冈（Ｑｕｅｒｃｕｓｍｙｒｓｉｎａｅｆｏｌｉａ）同时受外生菌根（ＥＣＭ）和内生菌根的感染。该研究揭示了香港次生林植物在自然条件下受菌根菌感染的普遍性,为进一步探讨华南地区菌根与退化地区植物群落演替的关系提供了重要的本底资料。