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.     

Practical aspects of mycorrhizal technology in some tropical crops and pastures

Summary Greenhouse and field experiments were conducted on the effect of VA mycorrhiza (VAM) on the growth of cassava, various tropical grass and legume species, as well as beans, coffee and tea. A large number of VAM fungal species were evaluated for effectivity in increasing cassava growth and P uptake in acid low-P soils. The effectivity of VAM species and isolates was highly variable and dependent on soil pH and fertilizer applications, as well as on soil temperature and humidity. Two species,Glomus manihotis andEntrophospora colombiana were found to be most effective for a range of crops and pastures, at low pH and at a wide range of N, P, and K levels. At very low P levels nearly all crops and pasture species were highly mycorrhizal dependent, but at higher soil P levels cassava and several pasture legumes were more dependent than grass species. Mycorrhizal inoculation significantly increased cassava and bean yields in those soils with low or ineffective indigenous mycorrhizal populations. In these soils cassava root yields increased on the average 20–25% by VAM inoculation, both at the experiment station and in farmers’ fields. VAM inoculation of various pasture legumes and grasses, in combination with rock phosphate applications, increased their early growth and establishment. Agronomic practices such as fertilization, crop rotations, intercropping and pesticide applications were found to affect both the total VAM population as well as its species composition. While there is no doubt about the importance of VA mycorrhiza in enhancing P uptake and growth of many tropical crops and pastures grown on low-P soils, much more research is required to elucidate the complicated soil-plant-VAM interactions and to increase yields through improved mycorrhizal efficiency.     

Dual inoculation with Aspergillus fumigatus and Glomus mosseae enhances biomass production and nutrient uptake in wheat (Triticum aestivum L.) supplied with organic phosphorus as Na-phytate

In a pot experiment, wheat was grown for 50 days in two heat-sterilized low-phosphorus (P) soils supplied with organic P as Na-phytate. Seed inoculation with the phosphatase-producing fungus (PPF) Aspergillus fumigatus or soil inoculation with the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus mosseae increased shoot and root dry weight and root length, phosphatase activity in the rhizosphere and shoot concentrations of P and to a lesser extent of K and Mg. As a rule, the greatest effects on those parameters were most in the combined inoculation treatment (PPF + VAM). Shoot concentrations of Cu and Zn were only enhanced by VAM, not by PPF. At harvest, depletion of organic P in the rhizosphere soil increased in the order of: sterilized soil < PPF < VAM < PPF + VAM which corresponded with the enhanced P concentrations in the plants. The results demonstrate that organic P in form of Na-Phytate is efficiently used by VAM and that use of organic P can be increased by simultaneous inoculation with phosphatase-producing fungi.     

Intersporal Genetic Variation of Gigaspora margarita, a Vesicular Arbuscular Mycorrhizal Fungus, Revealed by M13 Minisatellite-Primed PCR

Spores of vesicular arbuscular mycorrhizal (VAM) fungi contain thousands of nuclei. In order to understand the karyotic structure of a VAM fungus spore, the genetic variation of the first generation of spores from a VAM fungus (Gigaspora margarita) was examined. Spores originating from both single- and multispore inoculations of the species G. margarita were analyzed by M13 minisatellite-primed PCR. In both cases, different fingerprints were obtained from individual spores with few spores exhibiting similar fingerprints. These results can be explained only by a heterokaryotic status of the nuclear population within a spore.     

Dependency of cassava (Manihot esculanta Crantz) on vesicular-arbuscular mycorrhizal fungi

Cassava plants were started in the greenhouse either from small cuttings (2.0 mg P/cutting) or large cuttings (20.2 mg P/cutting) in a subsurface Oxisol not inoculated or inoculated with Glomus aggregatum at target soil solution P concentrations of 0.003–0.2 mg l^-1. Vesicular-arbuscular mycorrhizal (VAM) fungal colonization levels in excess of 60% were attained on cassava roots irrespective of the size of cutting material used or target soil solution P status. However, plants started from large cuttings grew faster and better than those started from smaller ones. Cassava was found to be very highly dependent on VAM fungi if grown from small cutting but only marginally dependent if grown from large cuttings. The lower dependence of cassava on VAM fungi when started from larger cuttings appears to be related to the high P reserve in these cuttings and hence the low requirement of the plants for soil P until the P reserve in the cuttings is significantly depleted.A contribution from the Hawaii Institute of Tropical Agriculture and Human Resources Journal Series No. 3896     

Growth and nutrient status of citrus plants as influenced by mycorrhiza and phosphorus application

To test the hypothesis that high levels of soluble phosphate applied in combination with VAM fungi, to citrus plants, can cause growth depression even in the absence of other limiting factors, and also to test if rock phosphate, under these conditions, may be a satisfactory P source, a greenhouse experiment was conducted using sterilized soil with four levels of phosphate (0, 50, 100 and 200 ppm P) supplied either as soluble P or as rock phosphate. Citrus seedlings were either inoculated with the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus etunicatum or left uninoculated. Six months after the start of the experiment, the plants were harvested and shoot dry weight, P and K uptake, root colonization and the number of spores in 50 cm³ of soil were determined. Significant increases were found in dry matter yields and in P and K contents, due to VAM fungus inoculation, at the zero and 50 ppm soluble P levels and at all rock phosphate levels. At 100 ppm soluble P, the development of VAM plants was equilvalent to that of non-VAM plants, and at 200 ppm, growth was significantly less than that of non-VAM plants. Root colonization and sporulation were reduced at higher P levels. The absolute growth depression of VAM plants at the higher P level was likely due to P toxicity. In addition, high leaf P and K concentrations may have interfered with carbohydrate distribution and utilization in these symbioses. Rock phosphate may be used with VAM citrus to substitute for medium amounts of soluble phosphate.     

The management of populations of vesicular-arbuscular mycorrhizal fungi in acid-infertile soils of a savanna ecosystem

A field trial was conducted at two sites in the savanna ecosystem of eastern Colombia to compare the effects of inoculation with vesicular-arbuscular mycorrhizal fungi (VAMF) ofBrachiaria dictyoneura (a tropical grass), cassava (Manihot esculenta), the tropical forage legume kudzu (Pueraria phaseoloides) andSorghum sp., and two phosphate sources. The second stage of the trial studied the effect of these pre-crop treatments on the subsequent growth, nutrition and VAM status of cowpea (Vigna unguiculata) andStylosanthes capitata in the following season, compared with both crops sown in native savanna. Inoculation significantly increased the levels of VAM and plant yields in the early growth stages of all crops during the first season, particularly with the rock phosphate (RP) source. The most significant increases were observed in the mycorrhiza-dependent cassava and kudzu crops up to 15 weeks after sowing, and were associated with increased foliar uptake of P and Mg. The effectiveness of the introduced inoculum was greater at the field site with a sandier soil. In the second season the levels of VAM in roots of cowpea andS. capitata were all increased significantly in pre-cropped plots compared with a savanna control. The increased presence of VAM was associated with significantly increased yields on plots previously sown to cassava, kudzu andSorghum sp. The data support the idea that increasing the VAMF inoculum potential of these acid-infertile soils by inoculation or pre-crops can greatly increase the rate of establishment of mycorrhiza-dependent host plants.     

A fungal root symbiont modifies plant resistance to an insect herbivore

Vesicular-arbuscular mycorrhizal (VAM) fungi are common root-colonizing symbionts that affect nutrient uptake by plants and can alter plant susceptibility to herbivores. I conducted a factorial experiment to test the hypotheses that colonization by VAM fungi (1) improves soybean (Glycine max) tolerance to grazing by folivorous Mexican bean beetle (Epilachna varivestis), and (2) indirectly affects herbivores by increasing host resistance. Soybean seedlings were inoculated with the VAM fungus Glomus etunicatum or VAM-free filtrate and fertilized with high-[P] or low-[P] fertilizer. After plants had grown for 7 weeks first-instar beetle larvae were placed on bagged leaves. Growth of soybean was little affected by grazing larvae, and no effects of treatments on tolerance of soybeans to herbivores were evident. Colonization by VAM fungus doubled the size of phosphorus-stressed plants but these plants were still half the size of plants given adequate phosphorus. High-[P] fertilizer increased levels of phosphorus and soluble carbohydrates, and decreased levels of soluble proteins in leaves of grazed plants. Colonization of grazed plants by VAM fungus had no significant effect on plant soluble carbohydrates, but increased concentration of phosphorus and decreased levels of proteins in phosphorus-stressed plants to concentrations similar to those of plants given adequate phosphorus. Mexican bean beetle mass at pupation, pupation rate, and survival to eclosion were greatest for beetles reared on phosphorus-stressed, VAM-colonized plants, refuting the hypothesis that VAM colonization improves host plant resistance. VAM colonization indirectly affected performance of Mexician bean beetle larvae by improving growth and nutrition of the host plant. Received: 28 February 1997 / Accepted: 23 June 1997     

Defoliation effects on mycorrhizal colonization, nitrogen fixation and photosynthesis in the Glycine-Glomus-Rhizobium symbiosis

Soybean [ Glycine max (L.) Merr. cv. Wells] plants grown in a greenhouse were inoculated with Rhizobium japonicum strain 61A118 and the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus fasciculatum (Thaxt. sensu Gerd.) Gerd. & Trappe. Plants were defoliated (26, 48 and 66%) throughout the growth period and evaluated for VAM colonization, N₂, fixation and photosynthesis at harvest (six weeks). Photosynthate stress as a result of defoliation affected nodulation and nodule activity most severely. Colonization of the roots by the VAM fungus was little affected in comparison, and the intensity of colonization increased with increasing stress. The CO₂-exchange rate decreased less with defoliation than did leaf mass, and photosynthetic efficiency increased with the severity of defoliation. The increase in photosynthetic efficiency was significantly correlated with increases in leaf P (r = 0.91) and N (r = 0.97) concentrations. The results suggest that the VAM fungus should not be regarded as a simple P source and C sink in the tripartite legume association. Threeway source/sink relationships (VAM-P, Rhizobium-N, and host leaf-C) are discussed.     

Growth,micronutrient content and vesicular-arbuscular fungi infection of herbaceous plants on lignite mine spoils: A greenhouse pot experiment

A greenhouse experiment was carried out to determine the growth, micronutrient content and vesicular-arbuscular mycorrhizal (VAM) colonization of fescue and clover plants growing on a composite lignite mine soil. Five VAM fungus inocula were used. Results show that mine spoils had a high potential acidity and poor physical and chemical conditions for plant growth. Toxic levels of Al, Mn and Zn were found in plant tissues, with fescue showing apical necrosis. Glomus deserticola was the only VAM fungus that succeeded in colonizing plant roots, although it failed to improve the establishment or development of either species.     

Modelling Fungal (Neozygites cf. Floridana) Epizootics in Local Populations of Cassava Green Mites (Mononychellus Tanajoa)

The fungus, Neozygitis cf. floridana is parasitic on the cassava green mite, Mononychellus tanajoa (Bondar) (Acari: Tetranychidae) in South America and may be considered for classical biological control of cassava green mites in Africa, where cassava is an important subsistence crop, cassava green mites are an imported pest and specific natural enemies are lacking. Spider mites generally have a viscous structure of local populations, a trait that would normally hamper the spread of a fungus that is transmitted by the contact of susceptible hosts with the halo of capilliconidia surrounding an infectious host. However, if infected mites search and settle to produce capilliconidia on sites where they are surrounded by susceptible mites before becoming infectious, then the conditions for maximal transmission in a viscous host population are met. Because the ratio between spider mites and the leaf area they occupy is constant, parasite-induced host searching behaviour leads to a constant per capita transmission rate. Hence, the transmission rate only depends on the number of infectious hosts. These assumptions on parasite-induced host search and constant host density lead to a simple, analytically tractable model that can be used to estimate the maximal capacity of the fungus to decimate local populations of the cassava green mite. By estimating the parameters of this model (host density, per capita transmission rate and duration of infected and infectious state) it was shown that the fungal pathogen can reduce the population growth of M. tanajoa, but cannot drive local mite populations to extinction. Only when the initial ratio of infectious to susceptible mites exceeds unity or the effective growth rate of the mite population is sufficiently reduced by other factors than the fungus (e.g. lower food quality of the host plant, dislodgement and death by rain and wind and predation), will the fungal pathogen be capable of decimating the cassava green mite population. Under realistic field conditions, where all of these growth-reducing factors are likely to operate, there may well be room for effective control by the parasitic fungus. This revised version was published online in November 2006 with corrections to the Cover Date.     

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.     

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.     

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.     

Influence of Vesicular-Arbuscular Mycorrhizal Fungi on the Response of Potato to Phosphorus Deficiency

Morphological and biochemical interactions between a vesicular-arbuscular mycorrhizal (VAM) fungus (Glomus fasciculatum [Thaxt. sensu Gerdemann] Gerdemann and Trappe) and potato (Solanum tuberosum L.) plants during the development of P deficiency were characterized. Nonmycorrhizal (NM) plants grown for 63 d with low abiotic P supply (0.5 mM) produced 34, 52, and 73% less root, shoot, and tuber dry matter, respectively, than plants grown with high P (2.5 mM). The total leaf area and the leaf area:plant dry weight ratio of low-P plants were substantially lower than those of high-P plants. Moreover, a lower shoot:root dry weight ratio and tuber:plant dry weight ratio in low-P plants than in high-P plants characterized a major effect of P deficiency stress on dry matter partitioning. In addition to a slower rate of growth, low-P plants accumulated nonreducing sugars and nitrate. Furthermore, root respiration and leaf nitrate reductase activity were lower in low-P plants than in high-P plants. Low abiotic P supply also induced physiological changes that contributed to the greater efficiency of P acquisition by low-P plants than by high-P plants. For example, allocation of dry matter and P to root growth was less restricted by P deficiency stress than to shoot and tuber growth. Also, the specific activities of root acid phosphatases and vanadate-sensitive microsomal ATPases were enhanced in P-deficient plants. The establishment of a VAM symbiosis by low-P plants was essential for efficient P acquisition, and a greater root infection level for P-stressed plants indicated increased compatibility to the VAM fungus. By 63 d after planting, low-P VAM plants had recovered 42% more of the available soil P than low-P NM plants. However, the VAM fungus only partially alleviated P deficiency stress and did not completely compensate for inadequate abiotic P supply. Although the specific activities of acid phosphatases and microsomal ATPases were only marginally influenced by VAM infection, VAM roots characteristically had a higher protein concentration and, consequently, enhanced microsomal ATPase and acid phosphatase activities on a fresh weight basis compared with NM roots. Morphological and ultrastructural details of VAM plants are discussed in relation to the influence of the VAM symbiosis on P nutrition of potato.     

Parasitic and mutualistic associations between a mycorrhizal fungus and soybean: The effect of phosphorus on host plant-endophyte interactions

Soybean [ Glycine max (L.) Merr. cv. Kent] plants were colonized by the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus fasciculatum (Thaxt. sensu Gerd.) Gerd. and Trappe in pot cultures using an inert medium and a nutrient solution. Phosphorus was provided initially as 0, 25,50, 100 or 200 mg hydroxyapatite [HAP, Ca₁₀(PO₄)₆(OH)₂] per pot. Under the low (0 mg HAP) and high (100 and 200 mg HAP) P regimes, VAM plants showed 20, 25 and 38% growth retardation, respectively, relative to non-colonized controls. At 50 mg HAP, VAM plant growth was significantly enhanced (14%). Dry weight and P content of both VAM and control plants increased with increased P availability throughout the HAP gradient. Intraradical VAM fungal biomass increased linearly with increasing P availability. Extraradical VAM fungal biomass was smaller than the intraradical component of the fungus at the lowest and highest levels of P addition in the growth medium. The ratio of extra- to intraradical mycelium, a suggested index of VAM fungal effectiveness, was greatest for the 50 mg HAP treatment, coinciding with growth enhancement of the host plant. This enhanced growth of the host at an intermediate P level was apparently a result of increased P uptake by the endophyte.     

Effect of vesicular-arbuscular mycorrhizal and phosphatase-producing fungal inoculation on growth and nutrition of white clover supplied with organic phosphorus

In a pot experiment white clover was grown for 50 d in two sterilized low phosphorus soils supplied with organic phosphorus as sodium phytate. Seed inoculation withAspergillus fumigatus, a phosphatase-producing fungus (PPF), and soil inoculation withGlomus mossece, a vesicular-arbuscular mycorrhizal (VAM) fungus resulted in increased shoot and root dry mass and root length, phosphatase activity in soil and shoot concentrations of P and to a lesser extent of Mg. The effects on these parameters were most distinct upon combined inoculation treatment (PPF+VAM). A significant increase in hyphal length density and decrease in Mn concentration was observed only upon PPF+VAM treatment. Shoot concentrations of Cu and Zn were highly enhanced by VAM but not by PPF. Of the total P uptake the PPF contribution accounted for 39–41 %, VAM contributed 41–50 % and their combination resulted in 55–58 %, in comparison to only 38–40 % contribution due to soil microorganisms under unsterilized conditions. The depletion of organic phosphorus in the rhizosphere soil increased in the order (PPF+VAM)>VAM>PPF> unsterilized soil>sterilized soil. The results demonstrate the efficient use of phytate phosphorus byA. fumigatus andG. mosseœ and suggest that dual inoculation is superior to a single one.     

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.     

Resistance Responses of Potato to Vesicular-Arbuscular Mycorrhizal Fungi under Varying Abiotic Phosphorus Levels

In mycorrhizal symbioses, susceptibility of a host plant to infection by fungi is influenced by environmental factors, especially the availability of soil phosphorus. This study describes morphological and biochemical details of interactions between a vesicular-arbuscular mycorrhizal (VAM) fungus and potato (Solanum tuberosum L. cv Russet Burbank) plants, with a particular focus on the physiological basis for P-induced resistance of roots to infection. Root infection by the VAM fungus Glomus fasciculatum ([Thaxt. sensu Gerdemann] Gerdemann and Trappe) was extensive for plants grown with low abiotic P supply, and plant biomass accumulation was enhanced by the symbiosis. The capacity of excised roots from P-deficient plants to produce ethylene in the presence or absence of exogenous 1-amino cyclopropane-1-carboxylic acid (ACC) was markedly reduced by VAM infection. This apparent inhibition of ACC oxidase (ACC_ox) activity was localized to areas containing infected roots, as demonstrated in split-root studies. Furthermore, leachate from VAM roots contained a potent water-soluble inhibitor of ethylene generation from exogenous ACC by nonmycorrhizal (NM) roots. The leachate from VAM-infected roots had a higher concentration of phenolics, relative to that from NM roots. Moreover, the rates of ethylene formation and phenolic concentration in leachates from VAM roots were inversely correlated, suggesting that this inhibitor may be of a phenolic nature. The specific activity of extracellular peroxidase recovered in root leachates was not stimulated by VAM infection, although activity on a fresh weight basis was significantly enhanced, reflecting the fact that VAM roots had higher protein content than NM roots. Polyphenol oxidase activity of roots did not differ between NM and VAM roots. These results characterize the low resistance response of P-deficient plants to VAM infection. When plants were grown with higher abiotic P supply, the relative benefit of the VAM symbiosis to plant growth decreased and root infection was lower. The in vivo ACC_ox activity was also greater in roots of plants grown on high levels of P compared with those grown on low levels, although the influence of VAM infection was partially to counteract the nutritional effect of P on ACC_ox activity. Similar to ACC_ox activity, extracellular peroxidase activity of roots increased linearly with increasing abiotic P supply, thus indicating a greater potential for resistance to VAM infection. These findings suggest that VAM fungi may alter phenolic metabolism of roots so as to hinder ethylene production and the root's ability to invoke a defense response. Raising the abiotic P supply to plants at least partially restores the capacity of roots to produce ethylene and may, in this way, increase the root's resistance to VAM infection.     

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.