Regulation of mycorrhizal infection by hormonal factors produced by hosts and fungi

Summary An overview of current research on hormonal factors produced by plants and fungi in mycorrhizal associations is presented. On the one hand, growth hormones in roots and their exudates influence the metabolism and growth of fungi. On the other, fungal hormones influence root morphology, metabolic changes and the growth of the entire plant.     

Acquisition of nutrients from organic resources by mycorrhizal autotrophic plants

Summary Evidence exists to suggest that mycorrhizal fungi are capable of producing enzymes allowing them to access carbon, nitrogen and phosphorus from complex organic resources in soil. This facility is mainly demonstrated in ectomycorrhizal and ericaceous endomycorrhizal fungi associated with highly organic soils and climatically stressed environments. These data support a direct nutrient cycling hypothesis proposed for tropical ectomycorrhizal forests. In terms of forest succession, the evidence agrees with a major contribution of the mycorrhizal symbiosis in late stages of the succession, where elemental cycling becomes increasingly more conservative and process rates limited by biotic factors. Here, tree growth benefits from direct nutrient cycling mediated by their mycorrhizal symbionts.     

Assimilation of mineral nitrogen and ion balance in the two partners of ectomycorrhizal symbiosis: Data and hypothesis

Summary Assimilation pathways of mineral nitrogen and ion balances of the two partners of ectomycorrhizal symbiosis (fungi and woody plants) are reviewed. Data are presented about the partners both in pure culture and in mycorrhizal association. The two forms of mineral nitrogen, ammonium and nitrate, differ in their mobility in the soil, their transport into the cells, their uptake rates by plants and their assimilation pathways. These metabolic differences are related to differences in adjustment of ion balances and carbon metabolism under conditions of nitrate or ammonium nutrition. The data obtained on the partners of ectomycorrhizal symbiosis are discussed from this point of view and the observations composed with those on herbaceous angiosperms.     

Influence of phosphorus supply and light intensity on mycorrhizal response inPisum-Rhizobium-Glomus symbiosis

The influence of mycorrhizal colonization withGlomus mosseae on parameters of N₂ fixation and plant growth was studied in pot experiments with pea plants (Pisum sativum L.) infected withRhizobium leguminosarum and supplied with varied levels of phosphorus (P) and nitrogen (N). Reduced light intensities were used to evaluate the dependence of the microsymbionts on assimilate supply. In plants grown with low P supply, mycorrhization increased the concentration of P in shoots, and thus N₂ fixation. Reduced light intensity significantly depressed mycorrhizal colonization and nodule growth in low-P plants. When P supply did not limit plant growth and N₂ fixation, however, the percentage of mycorrhizal colonization was reduced due to the higher P status, and the microsymbionts were not impaired by low light intensities. To maximize carbohydrate supply, another experiment was carried out at high light intensity of 900 mol m^–2s^–1 and with non-limiting P supply. Nitrogen fertilization, given as starter N, enhanced plant growth, but delayed nodule formation. Towards flowering, nodulation rapidly increased, but less so inGlomus inoculated plants. After 28 days mycorrhizal plants were lower in shoot dry weight, nodule dry weight and nitrogenase activity. The results suggest that under many, but not all, environmental conditions the host plant is able to restrict mycorrhizal colonization and, thus, to prevent impairment ofRhizobium symbiosis.deceased in May 1994     

Biological interactions in the mycorrhizosphere

Summary Microbial communities in the mycorrhizosphere (i.e. in the small volume of soil immediately surrounding a mycorrhizal root) actively interact with the establishment and functioning of the mycorrhizal symbiosis. Examples of competitive and mutualistic interactions are given, and are discussed in the light of recent observations and experimental results. The significance of biological interactions in the mycorrhizosphere is considered from the standpoints of plant ecology and of practical application.     

植物修复重金属与多环芳烃污染土壤强化措施研究进展

重金属和多环芳烃是土壤环境中的重要污染物,其复合污染土壤的修复已成为环境科学研究的热点问题。植物修复技术是目前修复土壤复合污染重要方法之一,但植物本身修复能力有限,需借助化学、微生物、基因工程等手段对其修复效果进行强化。本文对国内外近几年来植物修复重金属-多环芳烃复合污染强化措施研究成果进行综述,并重点讨论了植物根际生长促进菌及菌根在强化修复中的应用,在此基础上对未来该领域需要深入研究的科学问题进行了阐述。     

Costs and benefits of mycorrhizas: Implications for functioning under natural conditions

Summary It is paradoxical that most plants under natural conditions are infected with vesicular-arbuscular mycorrhizal fungi, yet that it is often difficult to demonstrate that infected plants receive any benefit from the association. The costs and benefits of infection are analysed and a hypothesis formulated that infection only yields benefits at times during the life cycle when P demand by the plant exceeds the capacity of the root system. A simulation model is described that suggests that infection density should be more or less constant below a threshold value of root P uptake rate, but that above this value roots should be non-mycorrhizal. More extensive study of mycorrhizas under field conditions is needed to test such predictions.     

Biochemical approaches to the study of plant-fungal interactions in arbuscular mycorrhiza

This communication compares some biochemical methods for quantifying colonization by arbuscular mycorrhizal (AM) fungi. The degree of mycorrhizal colonization can conveniently be measured by determining fungal specific sterols. AM-colonized plants show a specific synthesis of 24-methylene cholesterol and an enhanced level of campesterol (=24-methyl cholesterol). A gene probe for nitrate reductase, the key enzyme for nitrogen assimilation, has been developed, which allows the monitoring of the distribution of this enzyme in fungi. Among the phytohormones tested, only abscisic acid (ABA) is found at a considerably higher level in AM-colonized plants than in controls. The concentration of ABA is about twenty times higher in spores and hyphae of the AM fungusGlomus than in maize roots. Other phytohormones (auxins, cytokinins) do not show such alterations after mycorrhizal colonization. The roots of gramineous plants become yellow as a result of mycorrhizal colonization. The yellow pigment(s) formed is (are) deposited in larger quantities in the vacuole(s) of the root parenchyma and endodermis cells during the development of the gramineous plants. A substance isolated from such roots has now been identified as a C-14 carotenoid with two carboxylic groups, and named mycorradicin.     

Mycorrhizas in ecosystems

Summary The results of analyses of the distribution, structure and function of ericoid, ecto and vesicular-arbuscular mycorrhizas are used to challenge the conventional view, which was based largely upon studies of isolated plants and excised plant roots under controlled conditions, that the symbiosis is primarily involved in the capture of phosphate ions. In nature, each mycorrhizal type is associated with an ecosystem and soil environment with distinctive characteristics in which selection has favoured the development of a particular range of attributes. These attributes are evaluated and their importance for the individual plant and for the ecosystems in which they occur is assessed. It is concluded that knowledge of the full range of functions of each mycorrhizal type is essential for an understanding of the distribution and dynamics of the ecosystem in which it predominates.     

Phytoalexin response is elicited by a pathogen (Rhizoctonia solani) but not by a mycorrhizal fungus (Glomus mosseae) in soybean roots

Summary A container system was constructed to study the response of soybean roots to infection by mycorrhizal or pathogenic fungi. The system allows a rapid and synchronous inoculation byGlomus mosseae orRhizoctonia solani. The phytoalexin glyceollin was measured in roots of inoculated and uninoculated plants for a period of 30 days. A significantly increased content of phytoalexin was found inR. solani-infected roots as compared to uninfected control roots. However, there was no difference in the glyceollin contents of the mycorrhizal and the control roots for up to 23 days after inoculation. The accumulation of glyceollin inR. solani-infected roots was not influenced by a subsequent inoculation withG. mosseae. Moreover glyceollin accumulated in mycorrhizal plants to the same extent as in control plants when they were inoculated withR. solani. The two fungi did not mutually influence the course of infection when they were inoculated together.     

Phenols in mycorrhizal roots ofArachis hypogaea

Summary Roots of mycorrhizal groudnut plants had higher amounts of phenols compared to non-mycorrhizal roots. Histochemical study of the mycorrhizal root revealed the accumulation of phenols in hyphae and arbuscules of the fungus within the host.     

Response of maize to different inoculum densities of vesicular arbuscular mycorrhizal endophytes

Summary The effect of the inoculum density of vesicular arbuscular mycorrhizal endophytes on growth and development in maize was investigated in sterilized soil under glass-house conditions. Mycorrhizal plants grew robust and produced three times more dry weight than non-mycorrhizal plants. 40 or more endophytes per plant produced the highest mycorrhizal association and the maximum growth in maize. The uninoculated plants exhibited the symptoms of chlorosis.     

Comparative anatomy of the host-fungus interface in mycorrhizas

Summary There are several types of mycorrhizal symbiosis (ectomycorrhiza, endomycorrhiza, ectendomycorrhiza), and the interfaces between the host-plant and the fungal symbiont have different organizations. The interfaces between the partners are always limited on the one side by the fungal plasmalemma and on the other side by the plasmalemma of the host plant or the perisymbiont membrane derived from it The cytoplasms of the partners are therefore separated by a mixed apoplast consisting of a fungal wall and a host wall or an apposition layer.     

Morphological,biochemical and molecular changes during ectomycorrhiza development

Summary An ectomycorrhiza, a specialized root organ, is the result of a complex interaction leading to a finely-tuned symbiosis between a plant and a compatible ectomycorrhizal fungus. Ultrastructural observations combined with cytochemical and biochemical studies reveal that structural and metabolic changes in the symbiont cells lead to the final phenotype of the active ectomycorrhiza. In the present review these changes are interpreted as changes in gene expression and discussed within the context of ectomycorrhiza development. Recent genetic data indicate that the continued vegetative growth of the ectomycorrhizal hyphae and the root tissues, and their ability to switch to symbiotic organ formation, is basically controlled by developmentally critical genes. The activity of these symbiotic genes during the differentiation of ectomycorrhizas is associated with extensive changes in the concentration of particular polypeptides and protein biosynthesis. The present state of knowledge about the developmental biology of ectomycorrhizas allows only speculation about the events during their development.Puisant mes forces aux sources des galaxies En buvant la sève des arbres M. Jonasz     

The use of metabolomics to dissect plant responses to abiotic stresses

Plant metabolism is perturbed by various abiotic stresses. As such the metabolic network of plants must be reconfigured under stress conditions in order to allow both the maintenance of metabolic homeostasis and the production of compounds that ameliorate the stress. The recent development and adoption of metabolomics and systems biology approaches enable us not only to gain a comprehensive overview, but also a detailed analysis of crucial components of the plant metabolic response to abiotic stresses. In this review we introduce the analytical methods used for plant metabolomics and describe their use in studies related to the metabolic response to water, temperature, light, nutrient limitation, ion and oxidative stresses. Both similarity and specificity of the metabolic responses against diverse abiotic stress are evaluated using data available in the literature. Classically discussed stress compounds such as proline, γ-amino butyrate and polyamines are reviewed, and the widespread importance of branched chain amino acid metabolism under stress condition is discussed. Finally, where possible, mechanistic insights into metabolic regulatory processes are discussed.     

Bacterial phytotoxins: Mechanisms of action

Many species of phytopathogenic procaryotes produce toxins that appear to function in disease development. The affect the plant in different ways, the end result of which is the elicitation of chlorosis, necrosis, watersoaking, growth abnormalities or wilting. The most extensively studied toxins cause chlorosis. They specifically inhibit diverse enzymes, all critical to the plant cell. This inhibition results in a complex series of metabolic dysfunctions ultimately resulting in symptom expression. Substances causing growth abnormalities consist of known phytohormones and other compounds with plant hormone-like activities, but which have no structural relationship to the known hormones. The former act in the usual manner but, because of their elevated levels and imbalances, the host's regulatory mechanisms are overwhelmed and abnormal growth results (hyperplasia, shoot or root formation); the mechanisms of action of the latter group are unknown. High molecular weight, carbohydrate-containing substances, also acting in unknown ways, cause tissue watersoaking or wilting. Likewise, we know little about toxins causing necrosis except for syringomycin which affects ion transport across the plasmalemma.     

The significance of mycorrhizas for protective ecosystems

Summary On the basis of the reviews presented in this issue, the ecological significance of mycorrhizal symbioses is discussed. Mycorrhizas may have some importance in the acquisition of mineral nutrients during the productive phase of ecosystems in early stages of succession, but their main role is played during the protective phase of ecosystems in the final stages of succession when most resources are incorporated into biomass. In these successional stages, mycorrhizas short-circuit nutrient cycles by directly reacquiring nutrients in organic form from plant (and fungal) litter, and they may reallocate resources between different plant individuals, preventing loss of resources from the entire ecosystem.     

The influence of N2-fixation on the carbon balance of leguminous plants

Biological dinitrogen fixation by legume-rhizobia symbiosis is very important both from the economic and from the ecological point of view. Theoretically, the reduction of the N₂-molecule to ammonia requires at least 16 ATP and 1.5 mg C per mg N fixed (N_fix). These values are difficult to determine in situ as this necessitates the determination of that part of root respiration which drives N₂-fixation. New approaches to such determinations and the results obtained are described. The values vary, depending on the plant species studied, the developmental stage of the plants and the genetic variability of macro- (and micro-?) symbionts. The values range between 1.5 and 4 mg C/mg N_fix. In some species (e.g.Vicia faba L. cv. Fribo), the apparent CO₂ assimilation is enhanced in order to meet this high energy need. In others (e.g.Pisum sativum L. cv. Grapis), root growth is restricted. Physiological criteria are discussed which allow an early diagnosis of the energetic efficiency of various combinations of macro-and microsymbionts as a basis for a selection in plant breeding.     

Interaction ofRhizobium loti strain and host on the biosynthesis of unusual amino acids in leguminous plants

Summary The effect of differentRhizobium loti strains on the biosynthesis of 2,3-diamino-butanoic acid and 2,4-diamino-3-methyl-butanoic acid in root nodules ofLotus tenuis, Anthyllis vulneraria andLupinus densiflorus has been investigated. Results suggest that biosynthesis isRhizobium strain dependent, that the bacteroid is the site of synthesis of the compounds and that their biosynthesis is confined to the symbiosis.