Mycorrhizal responses of barley cultivars differing in P efficiency

The purpose of this study was to investigate how barley cultivars which are different in dry matter yield at low phosphorus (P) supply (i.e. they differ in agronomic P efficiency) respond to mycorrhizal infection. In a preliminary experiment, six mycorrhizal fungi were tested for their ability to colonize barley (Hordeum vulgare L.) roots at a soil temperature of 15°C.Glomus etunicatum was the most effective species and was used in the main experiment. The main experiment was conducted under glasshouse conditions in which soil temperature was maintained at 15°C. Treatments consisted of a factorial arrangement of 8 barley cultivars, 2 mycorrhiza (inoculated and non-inoculated), and 3 rates of P (0, 10 and 20 mg kg^-1). P utilization efficiency (dry matter yield per unit of P taken up) and agronomic P efficiency among the barley cultivars was significantly negatively correlated with mycorrhizal responses. However, the response to mycorrhizal infection was positively correlated with response to P application. Poor correlation was observed between P concentration when neither mycorrhiza nor P were supplied and the percentage of root length infected. The extent of mycorrhizal infection among the barley cultivars in soil without P amendment varied from 8.6 to 28.6%. Significant interactions between cultivar and P addition, and between mycorrhiza and P addition were observed for shoot dry weight but not root dry weight.     

Effects of vesicular-arbuscular mycorrhizal infection and phosphate on Plantago major ssp. pleiosperma in relation to the internal phosphate concentration

Two experiments were carried out to study physiological effects of vesicular-arbuseular mycorrhizal infection on Plantago major L., ssp. pleiosperma (Pilger). In the first experiment, infection by the Glomus fasciculatum (Thaxt. sensu Gerdemann) Gerdemann and Trappe increased growth, shoot to root ratio, P concentrations in both shoot and roots and total uptake of P per plant. The percentages of dry matter in both shoot and roots were lower in mycorrhizal plants.
In the second experiment different P treatments were applied to both mycorrhizal and non-mycorrhizal P. major plants to separate any effects of mycorrhizal infection from increased uptake of P. In addition to the effects found in the first experiment, mycorrhizal, P, and mycorrhizal x P interaction effects were found on root respiration rate and the concentration of soluble sugars in the roots. No clear effects on total dry weight, N and starch concentrations in shoot and roots and sugar concentraion in the shoot were found. Irrespective of the mycorrhizal treatment, increased P concentration in the shoot correlated with an increased shoot to root ratio and root respiration rate, and a decreased percentage dry matter and sugar concentration in the roots. However, the root respiration rate and the P concentration in the roots of mycorrhizal plants were enhanced more than expected from the increased P concentrations in the shoots of these plants.     

Mycorrhizal dependency of banana (Musa acuminata,AAA group) cultivar

Seven banana cultivars (Musa acuminata, AAA group) were inoculated with two species of vesicular arbuscular mycorrhizal (VAM) fungi (Glomus mosseae and Glomus macrocarpum) in a greenhouse experiment. Inoculated plants had generally greater shoot dry weight and shoot phosphorus concentrations compared to the noninoculated plants. A great variation in dependency on mycorrhizal colonization was observed among the banana cultivars. Cv. Williams showed the highest relative mycorrhizal dependency (RMD) and cv. Poyo the lowest. For all the cultivars studied, inoculation with G. macrocarpum resulted in the highest RMD values. Both root dry weight and root hair length or density of the noninoculated plants were inverserly correlated with the RMD values of cultivars.     

Effects of vesicular-arbuscular mycorrhizal infection and phosphate on Plantago major ssp. pleiosperma in relation to internal cytokinin concentrations

Relations between cytokinin concentrations and effects of P and vesicular-arbuscular mycorrhizal (VAM) infection were investigated in Plantago major L. ssp. pleiosperma Pilger. Both mycorrhizal infection by Glomus fasciculatum (Thaxt. sensu Gerdemann) Gerdemann and Trappe and P addition increased the shoot to root ratio, specific leaf area (SLA), and P concentrations of shoot and roots, and decreased the percentage of dry matter in the shoot during the experiment. In general, P concentration in the shoot and roots of each treatment correlated positively with the shoot to root ratio and specific leaf area, and negatively with the percentage of dry matter in the shoot. Cytokinin concentrations in the tissue of shoots and roots were determined using an enzyme-linked immunosorbent assay. Concentrations of zeatin and zeatin-ribosides in the free base and nucleotide fractions had increased more after P addition than in the case of mycorrhizal infection in both shoot and roots, whereas the P concentrations had increased less. It is suggested that zeatin and zeatin-ribosides are not the primary growth-substances involved in mediating VAM effects.     

Root colonization of different hosts by the vesicular-arbuscular mycorrhizal fungus Glomus dimorphicum

Colonization of the roots of beans, alfalfa, onions, red clover, corn, and four barley cultivars (Bonanza, Klondike, Gateway 63, and Olli) by Glomus dimorphicum Boyetchko and Tewari, a vesicular-arbuscular mycorrhizal fungus isolated from a barley field in Alberta, Canada, was studied under greenhouse conditions. Infection levels were low in all four barley cultivars but were higher in beans, alfalfa, and onions and were highest in red clover and corn roots. The infection patterns of G. dimorphicum varied among all the hosts. Coiling of intracellular hyphae occurred in corn, alfalfa, and red clover roots. Appreciable numbers of intraradical vesicles were found only in red clover and bean roots, while arbuscules formed in all hosts except barley. It was concluded that the pattern of root colonization by G. dimorphicum is influenced by the host genome and that the fungal morphology in the roots is variable and, thus, not diagnostic for the mycorrhizal species.     

Effects of liming and mycorrhizal colonization on soil phosphate depletion and phosphate uptake by maize (Zea mays L.) and soybean (Glycine max L.) grown in two tropical acid soils

The effects of liming and inoculation with the arbuscular mycorrhizal fungus, Glomus intraradices Schenck and Smith on the uptake of phosphate (P) by maize (Zea mays L.) and soybean (Glycine max [L.] Merr.) and on depletion of inorganic phosphate fractions in rhizosphere soil (Al-P, Fe-P, and Ca-P) were studied in flat plastic containers using two acid soils, an Oxisol and an Ultisol, from Indonesia. The bulk soil pH was adjusted in both soils to 4.7, 5.6, and 6.4 by liming with different amounts of CaCO₃.In both soils, liming increased shoot dry weight, total root length, and mycorrhizal colonization of roots in the two plant species. Mycorrhizal inoculation significantly increased root dry weight in some cases, but much more markedly increased shoot dry weight and P concentration in shoot and roots, and also the calculated P uptake per unit root length. In the rhizosphere soil of mycorrhizal and non-mycorrhizal plants, the depletion of Al-P, Fe-P, and Ca-P depended in some cases on the soil pH. At all pH levels, the extent of P depletion in the rhizosphere soil was greater in mycorrhizal than in non-mycorrhizal plants. Despite these quantitative differences in exploitation of soil P, mycorrhizal roots used the same inorganic P sources as non-mycorrhizal roots. These results do not suggest that mycorrhizal roots have specific properties for P solubilization. Rather, the efficient P uptake from soil solution by the roots determines the effectiveness of the use of the different soil P sources. The results indicate also that both liming and mycorrhizal colonization are important for enhancing P uptake and plant growth in tropical acid soils.     

磷水平对接种丛枝菌根真菌甜玉米苗期生长的影响

研究了不同外源磷水平条件下,接种丛枝菌根真菌根内球囊霉(Glomus intraradices)对寄主植物甜玉米菌根侵染率、地上部和地下部鲜重、氮磷含量、精氨酸含量影响。结果表明:丛枝菌根真菌能够很好的侵染于玉米植株根系。且不同磷水平条件下,菌根侵染率差异较显著。在低磷水平下,菌根侵染率较高。孢子数量随着磷水平提高而增加。菌丝室根外菌丝鲜重在P40时最高。菌根化的甜玉米生物量及氮磷含量显著高于对照组。此外,低磷水平促使甜玉米地上部和地下部鲜重显著提高。甜玉米地上部总氮和地下部总氮含量分别在P40、P80和P20、P40时最高。地上部总磷和地下部总磷含量分别在P80和P160时最高。菌根精氨酸含量在低磷(P20)时最高。研究表明接种丛枝菌根真菌可促进甜玉米幼苗生长并与外源磷水平有关。     

Growth response and phosphorus uptake of rye with long and short root hairs: Interactions with mycorrhizal infection

Plant growth and phosphorus (P) uptake of two selections of rye (Secale cereale L.) differing in length of root hairs, in response to mycorrhizal infection were investigated. Rye plants with short root hairs (SRH) had a greater length of root infected by Glomus intraradices (up to 32 m pot^–1) than those with long root hairs (LRH) (up to 10 m pot^–1). Application of P decreased the percentage of root length infected in both selections. In low-P soil, mycorrhizal infection increased shoot and root P concentration, especially in LRH plants. Generally, LRH had higher shoot dry weight than SRH plants. P uptake was increased both by LRH and by mycorrhizal infection. Differences in specific P uptake and P utilization efficiency between SRH and LRH plants were observed in non-mycorrhizal plants. With low P supply, P utilization efficiency (dry matter yield per unit of P taken up) of LRH plants increased with time. However, mycorrhizal infection reduced P utilization efficiency, particularly of SRH plants. SRH plants, which were agronomically less efficient (i.e. low dry matter yield at low P supply) were more responsive to either mycorrhizal infection or P addition than the LRH plants. No interaction was observed between mycorrhizal infection and root hair length.     

Variability among lettuce cultivars grown at two levels of available phosphorus

Fifteen lettuce cultivars representing three different morphological types were grown in a sand alumina system under conditions of low (deficient) and high (sufficient) P supply. An efficient plant was defined as one that produced a large shoot fresh weight under low P conditions. Cultivars within their respective groups varied significantly for some traits that appeared to be important in determining adaptation to P. This led to the conclusion that accumulation of P in shoot tissue or the total plant was the main difference between efficient and inefficient cultivars. Accumlation of P seemed to be due to a greater absorption capability of roots or greater root mass (weight), depending on the different lettuce groups. Differences in internal use of P did not contribute to differences in shoot fresh weight.The butterhead cultivars were the least efficient plants when grown under low P. Compared to the other groups, plants has lower translocation efficiency and a greater root: shoot ratio. Never-the-less, butterhead cultivars as efficient as the best cultivars of other groups were found. There were no differences between Brazilian and American cultivars for any of the traits analysed, probably due to the fact that in both countries vegetables are bred under high fertility levels and grown with heavy applications of fertilizers. The results of this study demonstrate that there were genotypic variability and/or genotype×environment interaction effects for shoot weight (yield) among the lettuce cultivars grown under low P conditions imposed in the sand-alumina system.     

Influence of soil pH on the soybean-endomycorrhiza symbiosis

Summary Soybean (Glycine max {L.} Merr.) cultivars were inoculated withGigaspora gigantea andGlomus mosseae to determine mycorrhizal: cultivar relationships as affected by soil pH. The specific cultivarfungal response was dependent on soil pH. Overall cultivar responses in unlimed soil (pH 5.1) were greater forG. gigantea thanG. mosseae. The Bossier —G. gigantea combination was particularly responsive in unlimed soil and showed an increase of 10% in shoot length, 35% in shoot dry weight. 75% in root dry weight, and 397% in nodule dry weight over uninoculated controls. Little cultivar response was observed withG. mosseae inoculation in unlimed soil. In limed soil (pH 6.2), the larger responses were obtained withG. mosseae inoculated plants, although inoculation with eitherG. mosseae orG. gigantea appeared effective. In general, nodulation was greater on mycorrhizal roots than on control roots.     

Mass production of Glomus mosseae spores

Five crops inoculated with Glomus mosseae were grown for 10 weeks and the development of mycorrhizal infection and sporulation were assessed. Infected roots from pot cultures of different ages were used to examine the host effect on the development of mycorrhizae. The effectiveness of each host was assessed by measuring spore numbers. For all hosts, the percentage of root length infected increased rapidly up to 10 weeks after sowing. Infectivity of root inocula increased with increasing percentage of root length infected with the inoculum for all crops, except where large numbers of mature spores (1755) had been produced on barley. The highest spore numbers were achieved in the rhizosphere of barley plants, followed by chickpea and beans. The lowest spore numbers were found in the rhizosphere of corn and okra plants. The type of the crop as well as the harvest date greatly influenced the size of the spore population and the extent of root colonization of G. mosseae.     

The effect of phosphorus availability on the carbon economy of contrasting common bean (Phaseolus vulgaris L.) genotypes

A common response to low phosphorus availability is increased relative biomass allocation to roots. The resulting increase in root:shoot ratio presumably enhances phosphorus acquisition, but may also reduce growth rates by diverting carbon to the production of heterotrophic rather than photosynthetic tissues. To assess the importance of increased carbon allocation to roots for the adaptation of plants to low P availability, carbon budgets were constructed for four common bean genotypes with contrasting adaptation to low phosphorus availability in the field ("phosphorus efficiency"). Solid-phase-buffered silica sand provided low (1 microM), medium (10 microM), and high (30 microM) phosphorus availability. Compared to the high phosphorus treatment, plant growth was reduced by 20% by medium phosphorus availability and by more than 90% by low phosphorus availability. Low phosphorus plants utilized a significantly larger fraction of their daytime net carbon assimilation on root respiration (c. 40%) compared to medium and high phosphorus plants (c. 20%). No significant difference was found among genotypes in this respect. Genotypes also had similar rates of P absorption per unit root weight and plant growth per unit of P absorbed. However, P-efficient genotypes allocated a larger fraction of their biomass to root growth, especially under low P conditions. Efficient genotypes had lower rates of root respiration than inefficient genotypes, which enabled them to maintain greater root biomass allocation than inefficient genotypes without increasing overall root carbon costs.     

Role of mycorrhizal infection in the growth and reproduction of wild vs. cultivated plants

Summary We tested the hypothesis that mycorrhizal infection benefits wild plants to a lesser extent than cultivated plants. This hypothesis stems from two observations: (1) mycorrhizal infection improves plant growth primarily by increasing nutrient uptake, and (2) wild plants often possess special adaptations to soil infertility which are less pronounced in modern cultivated plants. In the first experiment, wild (Avena fatua L.) and cultivated (A. sativa L.) oats were grown hydroponically at four different phosphorus levels. Wild oat was less responsive (in shoot dry weight) to increasing phosphorus availability than cultivated oat. In addition, the root: shoot ratio was much more plastic in wild oat (varying from 0.90 in the low phosphorus solution to 0.25 in the high phosphorus solution) than in cultivated oat (varying from 0.44 to 0.17). In the second experiment, mycorrhizal and non-mycorrhizal wild and cultivated oats were grown in a phosphorus-deficient soil. Mycorrhizal infection generally improved the vegetative growth of both wild and cultivated oats. However, infection significantly increased plant lifespan, number of panicles per plant, shoot phosphorus concentration, shoot phosphorus content, duration of flowering, and the mean weight of individual seeds in cultivated oat, while it had a significantly reduced effect, no effect, or a negative effect on these characters for wild oat. Poor positive responsiveness of wild oat in these characters was thus associated with what might be considered to be inherent adaptations to nutrient deficiency: high root: shoot ratio and inherently low growth rate. Infection also increased seed phosphorus content and reproductive allocation.     

Arbuscular mycorrhizal fungal-induced alteration to root architecture in strawberry and induced resistance to the root pathogenPhytophthora fragariae

Three strawberry cultivars Elsanta, Cambridge Favourite and Rhapsody were inoculated with eitherGlomus fasciculatum orGlomus etunicatum and their growth compared with non-inoculated plants. The roots of all inoculated plants were 55 to 70% colonised after 98 days. Increases in both root and shoot dry weights were measured. Root architecture was also determined and increases in branching were evident in AMF colonised root systems. The remaining plants were then inoculated with the root pathogenPhytophthora fragariae and allowed to grow for a further 58 days before harvest. In two of the cultivars, Cambridge Favourite and Elsanta, AMF reduced root necrosis by approximately 60 and 30% respectively. Only in the least susceptible cultivar, Rhapsody, was no reduction measured in AMF colonised plants. There were differences in the control conferred by the two arbuscular mycorrhizal fungi and this suggests there may be practical benefits of inoculation. Relationships between the presence of roots of different orders, on inoculation with the pathogen, and subsequent necrosis provided a mechanism for identifying root-architecture driven alteration to susceptibility. Root system necrosis was positively correlated with the proportion of the root system made up of higher order roots (3° to 4°) in non-colonised plants and negatively correlated in AMF colonised plants. These data suggest that root-architecture changes are not important per se but factors expressed concurrently may be.     

Rice cultivar evaluation for phosphorus use efficiency

Phosphorus deficiency is one of the most growth-limiting factors in acid soils in various parts of the world. The objective of this study was to screen 25 rice cultivars (Oryza sativa L.) at low, medium, and high levels of soil P. Number of tillers, root length, plant height, root dry weight and shoot dry weight were related to tissue P concentrations, P uptake and P-use efficiency. Shoot weight was found to be the plant parameter most sensitive to P deficiency. Significant cultivar differences in P use efficiency were found. Phosphorus use efficiency was higher in roots than shoots and decreased with increasing levels of soil P. Positive correlations were found among growth parameters such as plant height, tillers, root and shoot weight, and P content of roots and shoots. These results indicate selection of rice cultivars for satisfactory performance under low P availability can be carried out using shoot and root dry weight as criteria.     

不同磷效率小麦品种对缺磷胁迫反应的比较

在营养液培养条件下,以根据相对产量为指标筛选出的6个不同磷效率的小麦(Triticum aestivum L.)品种为材料,对其苗期在缺磷条件下生长、根冠磷含量及其分配,以及叶片韧皮部汁液中磷浓度等进行了比较研究。结果表明,缺磷抑制植株地上部生长,但刺激根系生长,导致植株根／冠比增加。无论在供磷或缺磷条件下,磷高效品种的根冠生长速率都低于磷低效品种。缺磷导致植株体内的磷含量下降与根系相比,地上部磷含量的下降速率更快。但在缺磷条件下,不同磷效率的小麦品种根冠间的磷分配变化没有差异。研究发现,在正常供磷条件下,磷高效小麦品种的叶片韧皮部汁液中磷浓度较低,而磷低效品种的叶片韧皮部汁液中磷浓度较高。但开始缺磷后,磷高效品种的叶片韧皮部汁液中的磷浓度下降较慢,使其相对磷浓度较高。缺磷后10天,磷低效品种叶片韧皮部汁液中的磷浓度为供磷对照的35．9％,而磷高效品种叶片韧皮部汁液中的磷浓度为供磷对照的59％。     

Effects of ultraviolet-B radiation on growth, mycorrhizas and mineral nutrition of silver birch (Betula pendula Roth) seedlings grown in low-nutrient conditions

The effects of increased ultraviolet‐B (UV‐B) radiation on the growth, mycorrhizas and mineral nutrition of silver birch (Betula pendula Roth) seedlings were studied in greenhouse conditions. Seedlings—planted in a birch‐forest top soil and sand substrate—were grown without additional nutrient supply. Ultraviolet treatment started immediately after the seedlings emerged and the daily integrated biologically effective UV‐B irradiance on the UV‐B‐treated plants was equivalent to a 25% depletion of stratospheric ozone under clear sky conditions. Visible symptoms of UV‐B damage or nutrient deficiency were not observed throughout the experiment. Seedling height and dry weight (DW) (measured after 58 days and 76 days of treatment) were not affected by increased UV‐B. However, a significant shift in DW allocation toward roots resulted in a lower shoot/root ratio and leaf area ratio in UV‐B‐treated plants compared to control plants. At the first harvest (after 58 days of treatment), the percentage of various mycorrhizal morphotypes and the number of short roots per unit of root length or weight were not affected by increased UV‐B despite significantly increased DW allocation toward roots. Subtle reduction in the allocation of nitrogen (N) to leaves and increased allocation of phosphorus (P) to roots may suggest cumulative effects that could affect the plant performance over the long‐term.     

Effect of Arbuscular Mycorrhizal Fungi On Yield and Phytoremediation Performance of Pot Marigold (Calendula officinalis L.) Under Heavy Metals Stress

In order to study the effect of mycorrhizal fungi (inoculated and non-inoculated) and heavy metals stress [0, Pb (150 and 300 mg/kg) and Cd (40 and 80 mg/kg)] on pot marigold (Calendula officinalis L.), a factorial experiment was conducted based on a randomized complete block design with 4 replications in Research Greenhouse of Department of Horticultural Sciences, University of Tehran, Iran, during 2012–2013. Plant height, herbal and flower fresh and dry weight, root fresh and dry weight and root volume, colonization percentage, total petal extract, total petal flavonoids, root and shoot P and K uptakes, and Pb and Cd accumulations in root and shoot were measured. Results indicated that with increasing soil Pb and Cd concentration, growth and yield of pot marigold was reduced significantly; Cd had greater negative impacts than Pb. However, mycorrhizal fungi alleviated these impacts by improving plant growth and yield. Pot marigold concentrated high amounts of Pb and especially Cd in its roots and shoots; mycorrhizal plants had a greater accumulation of these metals, so that those under 80 mg/kg Cd soil^?1 accumulated 833.3 and 1585.8 mg Cd in their shoots and roots, respectively. In conclusion, mycorrhizal fungi can improve not only growth and yield of pot marigold in heavy metal stressed condition, but also phytoremediation performance by increasing heavy metals accumulation in the plant organs.     

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

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

Manganese reduction in the rhizosphere of mycorrhizal and nonmycorrhizal maize

The influence of rhizosphere microorganisms and vesicular-arbuscular (VA) mycorrhiza on manganese (Mn) uptake in maize (Zea mays L. cv. Tau) plants was studied in pot experiments under controlled environmental conditions. The plants were grown for 7 weeks in sterilized calcareous soil in pots having separate compartments for growth of roots and of VA mycorrhizal fungal hyphae. The soil was left either uninoculated (control) or prior to planting was inoculated with rhizosphere microorganisms only (MO-VA) or with rhizosphere microorganisms together with a VA mycorrhizal fungus [Glomus mosseae (Nicol and Gerd.) Gerdemann and Trappe] (MO+VA). Mycorrhiza treatment did not affect shoot dry weight, but root dry weight was slightly inhibited in the MO+VA and MO-VA treatments compared with the uninoculated control. Concentrations of Mn in shoots decreased in the order MO-VA > MO+VA > control. In the rhizosphere soil, the total microbial population was higher in mycorrhizal (MO+VA) than nonmycorrhizal (MO-VA) treatments, but the proportion of Mn-reducing microbial populations was fivefold higher in the nonmycorrhizal treatment, suggesting substantial qualitative changes in rhizosphere microbial populations upon root infection with the mycorrhizal fungi. The most important microbial group taking part in the reduction of Mn was fluorescent Pseudomonas. Mycorrhizal treatment decreased not only the number of Mn reducers but also the release of Mn-solubilizing root exudates, which were collected by percolation from maize plants cultivated in plastic tubes filled with gravel quartz sand. Compared with mycorrhizal plants, the root exudates of nonmycorrhizal plants had two fold higher capacity for reduction of Mn. Therefore, changes in both rhizosphere microbial population and root exudation are probably responsible for the lower acquisition of Mn in mycorrhizal plants.