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.     

Live-cell imaging elaborating epidermal invasion and vascular proliferation/colonization strategy of Verticillium dahliae in host plants

The soilborne ascomycete fungus Verticillium dahliae causes destructive vascular wilt disease in hundreds of dicotyledonous plant species. However, our understanding of the early invasion from the epidermis to the vasculature and the prompt proliferation and colonization in the xylem tissues remains poor. To elaborate the detailed infection strategy of V. dahliae in host plants, we traced the whole infection process of V. dahliae by live-cell imaging combined with high-resolution scanning electron microscopy. The 4D image series demonstrated that the apex of invading hyphae becomes tapered and directly invades the intercellular space of root epidermal cells at the initial infection. Following successful epidermal invasion, the invading hyphae extend in the intercellular space of the root cortex toward the vascular tissues. Importantly, the high-resolution microscopic and live-cell images demonstrated (a) that conidia are formed via budding at the apex of the hyphae in the xylem vessels to promote systemic propagation vertically, and (b) that the hyphae freely cross adjacent xylem vessels through the intertracheary pits to achieve horizontal colonization. Our findings provide a solid cellular basis for future studies on both intracellular invasion and vascular colonization/proliferation during V. dahliae infection and pathogenesis in host plants.     

An Exploratory Study on the Influence of Vesicular-arbuscular Mycorrhizal Fungi on the Success of Weed Biological Control with Insects

This paper presents the results of an exploratory study to determine whether vesicular-arbuscular mycorrhizal (VAM) fungi affect the success of insects used as weed biological agents. No direct correlation between the colonization by VAM fungi alone and the success of root- and shoot-feeding insects was found. However, in Centaurea diffusa , plants colonized by VAM fungi had a lower shoot root ratio and increased attack and survival of the root beetle, Sphenoptera jugoslavica , in the field. Beetle success was also best on C. diffusa when the plant was growing among determinate grasses that largely cease growing after flowering in the spring or early summer. This is consistent with other reports that VAM hyphae can transfer nutrients between plant species from low to high sinks. From these results and related reports in the literature, it is suggested that plants with many stenophagous root-insects are those in which VAM fungi colonization increases nutrient partitioning to the roots with little or no decrease in plant growth. Root-feeding, rather than foliage- or seed-feeding, insects seem to be particularly effective as biocontrol agents for plants of this type.     

Seasonality of vesicular-arbuscular mycorrhizae in sedges in a semi-arid tropical grassland

Vesicular-arbuscular mycorrhizal (VAM) colonization and spore numbers in the rhizosphere of Cyperus iria L. and C. rotundus L., growing in a semi-arid tropical grassland, was studied during the 1993 and 1994 monsoons. In addition, climatic and chemical properties of the soils were determined in order to investigate their influence on mycorrhizal variables. VAM fungal association in the sedges was confirmed by plant- and root-trap culture techniques. The soil nutrients exhibited seasonal variations, but were highly variable between years. Intercellular hyphae and vesicles with occasional intraradical spores characterized mycorrhizal association in sedges. Dark septate fungi also colonized roots of sedges. Temporal variations in mycorrhizal colonization and spore numbers occurred, indicating seasonality. However, the patterns of mycorrhizal colonization and spore numbers were different during both the years. The VAM fungal structures observed were intercellular hyphae and vesicles. Changes in the proportion of root length with VAM structures, total colonization levels and spore numbers were related to climatic and edaphic factors. However, the intensity of influence of climatic and soil factors on VAM tended to vary with sedge species.     

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.     

Effect of soil liming and vesicular-arbuscular-mycorrhizal inoculation on the growth and micronutrient content of the teff plant

In a greenhouse experiment involving an acid soil teff [Eragrostis tef (Zucc.) Trotter] plants failed to grow unless the soil was limed or inoculated with either of two vesicular-arbuscular-mycorrhizal (VAM) fungi,Glomus mosseae orGlomus macrocarpum. Plant growth increased by liming and to a lesser extent by VAM fungal inoculation. Liming also enhanced root colonization by VAM fungi. Shoot micronutrient content generally increased as a result of inoculation, and decreased by increased lime applications.     

Arbuscular mycorrhizal structure and fungi associated with mosses

We investigated the colonization and diversity of arbuscular mycorrhizal (AM) fungi associated with 24 moss species belonging to 16 families in China. AM fungal structures, i.e. spores, vesicles, hyphal coils (including intracellular hyphae), or intercellular nonseptate hyphae, were found in 21 moss species. AM fungal structures (vesicles, hyphal coils, and intercellular nonseptate hyphae) were present in tissues of 14 moss species, and spores and nonseptate hyphae on the surface of gametophytes occurred in 15 species. AM fungal structures were present in 11 of the 12 saxicolous moss species and in six of the ten terricolous moss species, but absent in two epixylous moss species. AM fungal structures were only observed in moss stem and leaf tissues, but not in rhizoids. A total of 15 AM fungal taxa were isolated based on trap culture with clover, using 13 moss species as inocula. Of these AM fungi, 11 belonged to Glomus, two to Acaulospora, one to Gigaspora, and one to Paraglomus. Our results suggest that AM fungal structures commonly occur in most mosses and that diverse AM fungi, particularly Glomus species, are associated with mosses.     

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 dependency and growth enhancing ability of VA-mycorrhizal fungi for Eleusine coracana (finger millet)

Inoculation of finger millet (Eleusine coracana Gaertn.) plants with one of six different vesicular, arbuscular, mycorrhizal (VAM) fungi increased plant biomass, height, leaf area and absolute growth rate; however, effectiveness of the various VAM fungi varied significantly. Maximum root colonization and mycorrhizal efficacy was observed with plants inoculated with Glomus caledonicum. Among five host genotypes tested for mycorrhizal dependency against G. caledonicum, genotype HR-374 gave the highest plant biomass, mycorrhizal efficacy and root colonization, the inoculation resulting in increased mineral (phosphate, nitrogen, Zn²⁺ and Cu²⁺) content and uptake in shoots.     

Vesicular-arbuscular mycorrhizal fungi,particularly Glomus tenue,in Venezuelan bromeliad epiphytes

The mycorrhizal status of water-impounding tank bromeliad epiphytes from three locales differing in altitude and moisture regime within Venezuelan cloud forest was examined. Species of vesicular-arbuscular mycorrhizal (VAM) fungi found in arboreal soils were compared to VAM fungi found in terrestrial soils. Sixteen of the 19 epiphytes examined for the presence of VAM fungi had roots with infection stages; 14 of these specimens showed growth of the fine endophyte Glomus tenue. Fine endophyte was the only VAM fungus found associated with epiphytes in the driest locale studied, while coarse VAM fungi (Gigaspora and Scutellospora spp.) were found at sampling locales receiving more moisture. Root infection was usually composed of intercellular hyphae and peletons; few arbuscules were observed. However, abundant extracellular hyphae were often observed tangled about roots in arboreal soil. It is concluded that epiphytic bromeliads probably benefit, at least periodically, from VAM fungi scavenging for sporadically available nutrients in arboreal soils. Glomus tenue may be particularly important as a colonizing VAM fungus in drier sites of Venezuelan cloud forest. The species composition of VAM fungi in arboreal soils was different to that of terrestrial soils sampled directly under epiphytic bromeliad perches, suggesting that VAM fungi species associated with bromeliads are dispersed to their hosts by vagile animal vectors.     

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.     

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.     

Comparative study of mycorrhizal susceptibility and anatomy of four palm species

A morphological and anatomical study of the root systems of the palm species Brahea armata S. Watson, Chamaerops humilis L., Phoenix canariensis Chabaud and Phoenix dactylifera L. has been carried out to determine possible mycorrhizal colonization sites. Furthermore, the arbuscular mycorrhizal (AM) anatomical types formed by the four palm species in association with Glomus mosseae (Nicol. & Gerd.) Gerdemann & Trappe have been examined. The presence of a continuous sclerenchymatic ring in the outer cortex and aerenchyma in the inner cortex that are anatomical indicators of mycorrhizal nonsusceptibility in all four palm species is observed. The root systems of B. armata and C. humilis present only one group of third-order roots, while the third-order roots of P. canariensis and P. dactylifera may be divided into five different groups: short thick roots, mycorrhizal thickened roots, fine short roots, fine long roots, and pneumatorhizas. Third-order and some second-order roots of B. armata and C. humilis are susceptible to colonization by AM fungi, while only the mycorrhizal thickened roots form mycorrhizas with arbuscules in the Phoenix species. The root system of the Phoenix species also presents AM colonization in fine roots with only intraradical hyphae and spores, but without arbuscules, and pseudomantles of spores anchored in the pneumatorings of the second-order roots, which are described for the first time. The mycorrhizas formed by the four palm species are of an intermediate type, between the Arum and the Paris types, and are characterized by intercalary arbusculate coils and not only by intracellular but also by intercellular fungal growth. Our study suggests that a different degree of adaptation may exist among palm mycorrhizas toward the slow growth of palms and low spore numbers in the soil where they grow.     

Influence of edaphic and climatic factors on dynamics of root colonization and spore density of vesicular-arbuscular mycorrhizal fungi in Acacia farnesiana Willd. and A. planifrons W.et.A.

 A study was conducted to assess the dynamics of vesicular-arbuscular mycorrhizal (VAM) fungi associated with Acacia farnesiana and A. planifrons in moderately fertile alkaline soils. The intensity of root colonization by VAM fungi and the distribution of VAM fungal structures varied with host species over a period of time. The occurrence of vesicles with varied morphology in the mycorrhizal roots indicates infection by different VAM fungal species. This was further confirmed from the presence of spores belonging to different VAM fungal species in the rhizosphere soils. Root colonization and spore number ranged from 56% – 72% and 5 – 14 g^{–  1}soil in A. farnesiana and from 60% – 73% and 5 – 15 g^{–  1} soil in A. planifrons. Per cent root colonization and VAM spore number in the rhizosphere soil were inversely related to each other in both the Acacia species. However, patterns of the occurrence of VAM fungal structures were erratic. Spores of Acaulospora foveata, Gigaspora albida, Glomus fasciculatum, G. geosporum and Sclerocystis sinuosa were isolated from the rhizosphere of A. farnesiana whereas A. scrobiculata, G. pustulatum, G. fasciculatum, G. geosporum and G. microcarpum were isolated from that of A. planifrons. The response of VAM status to fluctuating edaphic factors varied with host species. In A. farnesiana though soil nitrogen (N) was positively correlated with root colonization, soil moisture, potassium and air temperature were negatively correlated to both root colonization and spore number. Per cent root colonization and spore number in A. planifrons were negatively related to each other. Further, in A. planifrons as the soil phosphorus and N were negatively correlated with the density of VAM fungal spores, the same edaphic factors along with soil moisture negatively influenced root colonization. Received: 16 May 1995 / Accepted: 7 February 1996     

The propagules of vesicular-arbuscular mycorrhizal (VAM) fungi capable of initiating VAM infection after topsoil disturbance

Summary The removal and storage of topsoil decreases the infectivity of vesicular-arbuscular mycorrhizal (VAM) fungi. The propagules of VAM fungi include spores, root fragments containing hyphae and vesicles, and soil hyphae. The viability of each type of propagule after disturbance will determine the initiation of VAM associations with plants recolonizing the disturbed site. This study aimed to examine which of the propagules of VAM fungi are capable of initiating VAM infection after soil disturbance. Soil from an open woodland site of low soil fertility, in southeastern Australia was wetsieved through a tier of three sieves (1 mm, 250 m and 106 m), and the following fractions were extracted: (i) root fragments, (ii) fungal hyphae, and (iii) VAM spores. Each fraction was tested to determine its potential to initiate VAM. Hyphae of VAM fungi grew from root fragments within 14 days. The VAM spore fraction initiated VAM infection after 28 days. VAM hyphal fragments did not produce any VAM infection even after 42 days.     

Characteristic responses of three tropical legumes to the inoculation of two species of VAM fungi in Andosol soils with different fertilities

The growth and mineral nutrition responses were evaluated of three tropical legumes, cowpea (Vigna unguiculata L. cv Kuromame), pigeonpea [Cajanus cajan L. (Millsp.) cv ICPL 86009] and groundnut (Arachis hypogaea cv Nakateyutaka) inoculated with two different species of VAM fungi, Glomus sp. (Glomus etunicatum-like species) and Gigaspora margarita, and grown in Andosols with different fertilities [Bray II-P: topsoil (72 ppm), subsoil (<0.1 ppm)]. Percent fungal root colonization was high in cowpea and groundnut but relatively low in pigeonpea in both soil types. Despite the low rate of root infection, significant growth responses were produced, especially in the inoculated pigeonpea plant. In all legumes, shoot dry matter production was favoured by the inoculations. Increases in shoot biomass due to mycorrhizae were greater in the subsoil than in the topsoil. Mycorrhization raised shoot concentrations of P and Ca (in cowpea and groundnut) and P and K (in pigeonpea) in the topsoil. Whereas the P concentration in shoots in the subsoil was not positively affected by VAM fungi, particularly in cowpea and pigeonpea, the concentration of K in such plants was significantly increased by VAM treatment. The results also showed that mycorrhizal enhancement of shoot micronutrient concentrations was very rare in all plants, with negative effects observed in certain cases. Cu concentration, in particular, was not affected by VAM formation in any of the plants, and Mn and Fe in pigeonpea and groundnut, respectively, remained the same whether plants were mycorrhizal or not. In both soils the three legumes responded to Glomus sp. better than to Gigaspora margarita, and the effects of the VAM fungi on each of the crops relative to the controls were greater in the subsoil than in the topsoil. However, shoot growth of groundnut was not affected as much as cowpea and pigeonpea by the type of soil used. In spite of the relatively low infection of its root, pigeonpea was generally the most responsive of the three legume species in terms of mycorrhizal/nonmycorrhizal ratios.     

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.     

Mycorrhizae in Adenostoma fasciculatum Hook. & Arn.: a combination of unusual ecto- and endo-forms

 The mycorrhizal status of Adenostoma fasciculatum, the dominant shrub in California chaparral, has been unclear. In two typical, nearly monospecificstands, A. fasciculatum was found to have arbuscules and intercellular hyphae. Antisera detected hyphae of the arbuscular mycorrhizal (AM) fungal genera Acaulospora, Glomus, and Scutellospora, although we found only spores of Glomus. Some roots had partial sheaths and inter- and intracellular septate fungi without indications of root necrosis. Ectomycorrhizal root tips were also found, including Cenococcum and other unknown taxa. Sporocarps of EM fungi including species of Rhizopogon, Pisolithus, Balsamia, Laccaria, Hygrophorus, and Cortinarius were found in the stand, with no other EM or arbutoid mycorrhizal plants nearby. These observations indicate that A. fasciculatum forms mycorrhizae with AM, septate, and EM fungi, but often fails to form easily recognizable mycorrhizal structures. Accepted: 5 September 1998     

Asexual reproduction of <Emphasis Type="Italic">Hygrophorus olivaceoalbus</Emphasis> by intracellular microsclerotia in root cells of <Emphasis Type="Italic">Picea abies</Emphasis> – A winner of ozone stress?

Hygrophorus olivaceoalbus has long been known as an ectomycorrhizal fungus, formerly designated with the artificial binomen Piceirhiza gelatinosa. Recently, it has been found to be abundant and very frequent under double ambient ozone free air fumigation of mature Norway spruce trees. As it has already been reported that this fungus can form intercellular hyphae within the root meristem, a more detailed study was performed to clarify its type of root colonization. The present study not only revealed intercellular hyphae within the meristematic zone but also intracellular hyphae within root cortex cells which grow towards and into large tannin droplets—phenolic compounds usually deposited as defensive aids—to finally fill them completely. The hyphal assemblages become globular which at first sit as separated hyphal balls within cells still containing cytoplasm. Later on, they are apparently released from the root cells, presumably as microsclerotia for dispersal of the species. Old ectomycorrhizae (ECM) show an apical pore, and later a large orifice of a tube-like cylinder formed by the thick, persistent hyphal mantle. The root tissue is progressively degrading towards proximal parts. Disintegrating root cells apparently liberate the microsclerotia through the orifice. Further studies have to find out the mechanism by which the microsclerotia are liberated and whether they operate as asexual propagules and lastly how and by whom they are propagated. As these ECM are found under ozone stress, and with identical features at higher altitudes, stress impact on trees might be the causative agent for the high frequency and abundance of Hygrophorus olivaceoalbus ECM.     

Medicago truncatula Vapyrin is a novel protein required for arbuscular mycorrhizal symbiosis

Arbuscular mycorrhizal (AM) symbiosis is a widespread mutualism formed between vascular plants and fungi of the Glomeromycota. In this endosymbiosis, fungal hyphae enter the roots, growing through epidermal cells to the cortex where they establish differentiated hyphae called arbuscules in the cortical cells. Reprogramming of the plant epidermal and cortical cells occurs to enable intracellular growth of the fungal symbiont; however, the plant genes underlying this process are largely unknown. Here, through the use of RNAi, we demonstrate that the expression of a Medicago truncatula gene named Vapyrin is essential for arbuscule formation, and also for efficient epidermal penetration by AM fungi. Vapyrin is induced transiently in the epidermis coincident with hyphal penetration, and then in the cortex during arbuscule formation. The Vapyrin protein is cytoplasmic, and in cells containing AM fungal hyphae, the protein accumulates in small puncta that move through the cytoplasm. Vapyrin is a novel protein composed of two domains that mediate protein–protein interactions: an N‐terminal VAMP‐associated protein (VAP)/major sperm protein (MSP) domain and a C‐terminal ankyrin‐repeat domain. Putative Vapyrin orthologs exist widely in the plant kingdom, but not in Arabidopsis, or in non‐plant species. The data suggest a role for Vapyrin in cellular remodeling to support the intracellular development of fungal hyphae during AM symbiosis.