The enhanced growth of plants infected by vesicular–arbuscular (VA) mycorrhizal fungi results primarily from improved uptake of soil phosphate. Extra phosphate reaches the root through the fungal hyphae, which tap the soluble P in soil beyond the phosphate-depletion zone near the root surface. This mechanism can explain the many corrrelations between root geometry and mycotrophy and other nutritional effects of VA mycorrhizae such as increased uptake of zinc and copper ions. Recently VA mycorrhizae have been shown to increase the levels of chlorophyll and some hormones in plants and to alleviate water stress. Legumes are now receiving considerable attention because VA mycorrhiza affects nitrogen fixation in them indirectly by its action on P uptake. In this review the physiology of the VA mycorrhizal symbiosis is discussed in categories reflecting successive stages in its formation and function: (i) activation of the VA mycorrhizal propagules; (ii) penetration and initial infection of the host plant; (iii) spread of infection in roots; (iv) response of the plant; the components and mechanisms of VA mycorrhizal systems; (v) benefits to the fungus; carbon sinks; and (vi) imbalances in the symbiosis. It is suggested that studies on the physiological complexities of VA mycorrhizal associations should take more account of the biological diversity of VA mycorrhizal fungi and the wide range of host–endophyte–soil specificities.
Light and temperature greatly influenced the development of vesicular-arbuscular mycorrhiza and growth of onions in a phosphate-deficient soil. There were more large arbuscules and host growth was stimulated more with 25,000 lux than with 13,000 lux at 23° C and in a 14-23° C diurnal cycle. At 14° C and 13,000 lux mycorrhiza caused no growth stimulation even in three low-phosphate soils. At 18° C infection was much sparser in a 6 h daylength than in 12 and 18 h. Mycorrhizal plants kept in daylengths of 6 h at 13,000 lux were 3.5 times heavier than their non-inoculated controls. The effect of infection increased in longer daylengths and higher light intensities to 14.2 times the weight of controls with 18 h at 25,000 lux. The addition of soluble phosphate stimulated growth to the same extent as mycorrhizal inoculation did in the highest light conditions but phosphate stimulated growth more than did mycorrhiza under intermediate light conditions. The amounts of soluble carbohydrate in the roots of plants given phosphate and in those that were mycorrhizal did not differ significantly, but there was more soluble carbohydrate in plants growing in most light. Plants both with and without mycorrhiza contained much glucose, fructose, sucrose and an unidentified sugar with a low RF value, but there was no indication of fungal carbohydrates such as trehalose and mannitol.
Summary Mycorrhizal infections formed by different endophytes were examined in 10 crop species grown separately and in pairs in sterilized and unsterile soils. No infection was observed in cabbage, kale, rape or swede (in the supposedly non‐mycorrhizal family Cruciferae) and only traces were seen in sugar beet (supposedly non‐mycorrhizal Chenopodiaceae) when these plants were grown alone. However, slight (< 5 %) infection (cortical mycelium and vesicles, but no arbuscules) developed in some when a mycorrhizal host plant was present and there were many clumps of endophyte mycelium on their root surfaces, usually attached to entry points which had often aborted. Glomus fasciculatus‘E3’ was a more infective endophyte than Gigaspora margarita. Infection was usually well developed in the host plants barley, lettuce, maize, potato and onion. It was depressed only in a few pairs but no more by the presence of a ‘non‐host’ plant than by a host plant. The results suggest that the barriers to mycorrhizal infection in ‘non‐hosts’ are intrinsic and more probably related to characteristics of the root cortex or epidermis than to any infection‐inhibiting factors that might be released in root exudates.
SUMMARYLight and temperature greatly influenced the development of vesicular-arbuscular mycorrhiza and growth of onions in a phosphate-deficient soil. There were more large arbuscules and host growth was stimulated more with 25,000 lux than with 13,000 lux at 23° C and in a 14-23° C diurnal cycle. At 14° C and 13,000 lux mycorrhiza caused no growth stimulation even in three low-phosphate soils. At 18° C infection was much sparser in a 6 h daylength than in 12 and 18 h. Mycorrhizal plants kept in daylengths of 6 h at 13,000 lux were 3.5 times heavier than their non-inoculated controls. The effect of infection increased in longer daylengths and higher light intensities to 14.2 times the weight of controls with 18 h at 25,000 lux. The addition of soluble phosphate stimulated growth to the same extent as mycorrhizal inoculation did in the highest light conditions but phosphate stimulated growth more than did mycorrhiza under intermediate light conditions. The amounts of soluble carbohydrate in the roots of plants given phosphate and in those that were mycorrhizal did not differ significantly, but there was more soluble carbohydrate in plants growing in most light. Plants both with and without mycorrhiza contained much glucose, fructose, sucrose and an unidentified sugar with a low RF value, but there was no indication of fungal carbohydrates such as trehalose and mannitol.
Summary Onion and Coprosma plants were grown in a range of soils mostly containing very little available phosphate. Very large increases of shoot dry weight (up to eighteen‐fold with Coprosma and nineteen‐fold with onion) were obtained by adding phosphate. Similar increases (up to fifteen‐fold with Coprosma and twelve‐fold with onion) were obtained by mycorrhizal inoculation. The response to phosphate equalled or slightly exceeded the response to mycorrhiza, except in two soils rich in phosphate in which there was no response to either, and in one soil in which the mycorrhizal onion and Coprosma plants were twice as heavy as those given phosphate. The greatly increased growth from either treatment was associated with a large increase in the uptake of phosphorus. The probability of such growth responses, but not their size, was usually predictable in soils containing very little phosphorus soluble in CaCl2. In all soils with inoculum mycorrhizal infection was extensive. The sources of phosphorus available to mycorrhizal roots but not to non‐mycorrhizal roots differed in the various soils. In some inhibitory factors might make vesicular‐arbuscular mycorrhiza less effective in promoting the growth of the host plants.
SUMMARY Interactions between vesicular‐arbuscular (VA) mycorrhiza, utilization of rock phosphate and nodulation were examined in three legumes (clover, Stylosanthes and Centrosema) and in onions, grown in eight P‐deficient soils ranging from pH 8.1 to 5.3. Irrespective of pH, inoculation with VA endophytes increased P uptake in all host plants in all the soils when the indigenous endophytes had been removed by irradiation, but appreciable increases in plant dry weight only occurred when P concentrations of the uninoculated plants were low, generally below 0.15%. In the acid soils adding rock phosphate generally improved growth of the non‐mycorrhizal plants and inoculation with VA endophytes greatly improved its utilization. The effects persisted when the soil was used a second time. In neutral and alkaline soils rock phosphate was unavailable to non‐mycorrhizal plants and remained so after inoculation with VA endophytes. Legumes inoculated with the appropriate Rhizobium strain only nodulated in the most P‐deficient soils when they were also mycorrhizal, and added rock phosphate greatly improved nodulation and nitrogen fixation of the mycorrhizal plants. Some pilot experiments in unsterile soils are also described and the bearing of these results on field inoculation with VA endophytes is discussed.
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