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1.
An ecological view of the formation of VA mycorrhizas   总被引:3,自引:0,他引:3  
In spite of the major advances in understanding the functioning of symbioses between plants and arbuscular mycorrhizal fungi, details of the ecology of mycorrhizal fungi are not well documented. The benefits of the association are related to the timing and extent of colonization of roots, and fungi differ in their contribution to plant growth and presumably to soil aggregation. Knowledge of the processes that lead to successful colonization of roots by beneficial fungi at appropriate times for the host plants will form the basis of guidelines for soil management to maximize the benefits from the symbiosis. Fungi differ in the manner and extent to which they colonize roots. They also differ in their capacity to form propagules. The importance of hyphae, spores and propagules within living or dead mycorrhizal roots also differs among species and for the same species in different habitats. The relationships between colonization of roots and propagule formation, and between propagule distribution and abundance and subsequent mycorrhiza formation, for different fungi in field environments, are not well understood. Methods for quantifying mycorrhizal fungi are not especially suitable for distinguishing among different fungi within roots. Consequenctly, the dynamics of colonization of roots by different fungi, within and between seasons, have been little studied. Research is required that focuses on the dynamics of fungi within roots as well as on changes in the abundance of propagules of different fungi within soil. Interactions between fungi during the colonization of roots, the colonization of soil by hyphae and sporulation are all poorly understood. Without knowledge of these processes, it will by difficult to predict the likely success of inoculation with introduced fungi. Such knowledge is also required for selecting soil management procedures to enhance growth and survival of key species within the population. The relative tolerance of various fungi to perturbations in their surroundings will provide a basis for identifying those fungi that are likely to persist in specific environments. The processes that influence mycorrhizal fungi in field soils can be identified in controlled studies. However, greater emphasis is required on studying these processes with mixed populations of fungi. The role played by diversity within populations of mycorrhizal fungi is virtually unexplored.  相似文献   

2.
Legume plants are able to establish root nodule symbioses with nitrogen-fixing bacteria, called rhizobia. Recent studies revealed that the root nodule symbiosis has co-opted the signaling pathway that mediates the ancestral mycorrhizal symbiosis that occurs in most land plants. Despite being unable to induce nodulation, rhizobia have been shown to be able to infect and colonize the roots of non-legumes such as rice. One fascinating question is whether establishment of such associations requires the common symbiosis (Sym) genes that are essential for infection of plant cells by mycorrhizal fungi and rhizobia in legumes. Here, we demonstrated that the common Sym genes are not required for endophytic colonization of rice roots by nitrogen-fixing rhizobia.  相似文献   

3.
植物菌根共生磷酸盐转运蛋白   总被引:1,自引:0,他引:1  
大多数植物能和丛枝菌根(arbuscular mycorrhiza, AM)真菌形成菌根共生体。AM能够促进植物对土壤中矿质营养的吸收,尤其是磷的吸收。磷的吸收和转运由磷酸盐转运蛋白介导。总结了植物AM磷酸盐转运蛋白及其结构特征,分析其分类及系统进化,并综述了AM磷酸盐转运蛋白介导的磷的吸收和转运过程及其基因的表达调控。植物AM磷酸盐转运蛋白属于Pht1家族成员,它不仅对磷的吸收和转运是必需的,而且对AM共生也至关重要,为进一步了解菌根形成的分子机理及信号转导途径提供了理论基础。  相似文献   

4.
Abstract

Colonization of plant roots by arbuscular mycorrhizal fungi can greatly increase the plant uptake of phosphorus and nitrogen. The most prominent contribution of arbuscular mycorrhizal fungi to plant growth is due to uptake of nutrients by extraradical mycorrhizal hyphae. Quantification of hyphal nutrient uptake has become possible by the use of soil boxes with separated growing zones for roots and hyphae. Many (but not all) tested fungal isolates increased phosphorus and nitrogen uptake of the plant by absorbing phosphate, ammonium, and nitrate from soil. However, compared with the nutrient demand of the plant for growth, the contribution of arbuscular mycorrhizal fungi to plant phosphorus uptake is usually much larger than the contribution to plant nitrogen uptake. The utilization of soil nutrients may depend more on efficient uptake of phosphate, nitrate, and ammonium from the soil solution even at low supply concentrations than on mobilization processes in the hyphosphere. In contrast to ectomycorrhizal fungi, nonsoluble nutrient sources in soil are used only to a limited extent by hyphae of arbuscular mycorrhizal fungi. Side effects of mycorrhizal colonization on, for example, plant health or root activity may also influence plant nutrient uptake.  相似文献   

5.
Zhu H  Riely BK  Burns NJ  Ané JM 《Genetics》2006,172(4):2491-2499
Most land plants can form a root symbiosis with arbuscular mycorrhizal (AM) fungi for assimilation of inorganic phosphate from the soil. In contrast, the nitrogen-fixing root nodule symbiosis is almost completely restricted to the legumes. The finding that the two symbioses share common signaling components in legumes suggests that the evolutionarily younger nitrogen-fixing symbiosis has recruited functions from the more ancient AM symbiosis. The recent advances in cloning of the genes required for nodulation and AM symbioses from the two model legumes, Medicago truncatula and Lotus japonicus, provide a unique opportunity to address biological questions pertaining to the evolution of root symbioses in plants. Here, we report that nearly all cloned legume genes required for nodulation and AM symbioses have their putative orthologs in nonlegumes. The orthologous relationship can be clearly defined on the basis of both sequence similarity and microsyntenic relationship. The results presented here serve as a prelude to the comparative analysis of orthologous gene function between legumes and nonlegumes and facilitate our understanding of how gene functions and signaling pathways have evolved to generate species- or family-specific phenotypes.  相似文献   

6.
Abstract

The importance of vesicular arbuscular mycorrhizas (VAM) in an agricultural crop production system depends largely on our ability, through soil management, to increase the effectiveness of the indigenous mycorrhizal fungal population. To do so requires a good understanding of the functional ecology of the symbiosis.

In this article, we discuss primarily our programs at Guelph, which have focused on two aspects of the symbiosis: the influence of phosphate (P) fertilization on colonization and the influence of soil disturbance by tillage on colonization and P absorption.

Although it is generally accepted that the level of colonization of roots by VAM fungi decreases with increased P availability, we have found that the decrease is not as marked as thought. A reasonable degree of colonization was observed at available P levels well above those required for maximum yield. We have also found that the reduction in colonization occurs to a greater extent in the roots growing in a fertilized zone than in those outside this zone. Thus, although a band application of fertilizer may markedly reduce colonization in the fertilized zone, the remainder of the root system would be well colonized, and have an increased ability to acquire phosphorus.

That soil disturbance by tillage reduces the effectiveness of the VAM symbiosis in maize was first observed in the early 1980s. Since then we have conducted numerous field and growth chamber experiments to determine the practical importance of this effect in crop production and to elucidate the mechanisms involved. Systems such as no-till and ridge-till have been shown to result in greater P absorption during early maize growth. They also result in earlier colonization in some, but not all, cases. We have not, as yet, observed a yield advantage as a result of the earlier P absorption. This may be because factors other than P nutrition are limiting yields with the reduced tillage systems. Our results do indicate quite clearly, however, that greater rates of P fertilizer are not required in reduced tillage systems, compared with systems that cause a greater degree of soil disturbance. We suggest that lesser rates of P fertilizer may be required, which would have both an economic and environmental impact.

Our studies to elucidate the mechanisms have led to the conclusion that the integrity of the extraradical mycelium from a previous crop is a critical factor in the greater early P absorption in undisturbed systems. It may also be important in rapid colonization of roots of newly developing seedlings. This latter effect, however, has been quite inconsistent in our studies. It is apparent that some unknown factor or factors, in addition to the integrity of the mycelium, is also critical.

The fact that a preexisting extraradical mycelium is important for early P nutrition under our field conditions led to the hypothesis that the mycelium from a previous crop retains its infectivity and is able to absorb and transport P to newly attached roots following an extensive period during which the soil is frozen. Experiments in which pouches containing soil with a root-free mycelium were exposed to freezing for differing lengths of time have provided evidence to support this hypothesis. Roots growing in undisturbed pouches removed from the field when the soil was frozen were rapidly colonized during a bioassay. Disturbance of the soil in these pouches invariably reduced P absorption by the bioassay plants and sometimes, but not always, reduced colonization.

A more thorough understanding of the mechanisms involved in the effect of disturbance on colonization would increase the potential for increasing the effectiveness of the VAM symbiosis in crop production through management practices.  相似文献   

7.
Associations between plants and arbuscular mycorrhizal (AM) fungi are widespread and well-studied. Yet little is known about the pattern of association between clonal plants and AM fungi. Here we report on the pattern of mycorrhizal association within the rhizome systems of mayapple, Podophyllum peltatum. Mayapple is a long-lived understory clonal herb that is classified as obligately mycorrhizal. We found that while all mayapple rhizome systems maintained mycorrhizal associations, the percent colonization of roots by AM fungi differed among ramets of different age. The highest concentrations of AM fungi were in the roots of intermediate-aged ramets, while roots beneath the youngest ramet were not colonized. This pattern of ramet age or position-dependent colonization was observed in two separate studies; each conducted in a different year and at a different site. The pattern of AM fungal colonization of mayapple rhizome systems suggests that the mycorrhizal relationship is facultative at the ramet level. This conclusion is reinforced by our observation that augmentation of soil phosphate lowers root colonization by AM fungi. We also found that soil phosphate concentrations were depleted by ca. 1% under the same ramet positions where roots bore the highest AM fungal loads. Three non-exclusive hypotheses are proposed regarding the mechanisms that might cause this developmentally dependent pattern of mycorrhizal association.  相似文献   

8.
Abstract

Members of the Australian native perennial Fabaceae have been little explored with regard to their root biology and the role played by arbuscular mycorrhizal (AM) fungi in their establishment, nutrition and long-term health. The ultimate goal of our research is to determine the dependency of native perennial legumes on their co-evolved AM fungi and conversely, the impact of AM fungal species in agricultural fields on the productivity of sown native perennial legume pastures. In this paper we investigate the colonisation morphology in roots and the AMF, identified by spores extracted from rhizosphere soil, from three replicate plots of each of the native legumes, Cullen australasicum, C. tenax and Lotus australis and the exotic legumes L. pedunculatus and Medicago sativa. The plants were grown in an agricultural field. The level and density of colonisation by AM fungi, and the frequency of intraradical and extraradical hyphae, arbuscules, intraradical spores and hyphal coils all differed between host plants and did not consistently differ between native and exotic species. However, there were strong similarities between species in the same genus. The three dominant species of AM fungi in rhizosphere soil also differed with host plant, but one fungus (Glomus mosseae) was always the most dominant. Sub-dominant AM species were the same between species in the same genus. No consistent differences in dominant spores were observed between the exotic and native Fabaceae species. Our results suggest that plant host influences the mycorrhizal community in the rhizosphere soil and that structural and functional differences in the symbiosis may occur at the plant genus level, not the species level or due to provenance.  相似文献   

9.
Phosphorus effect on phosphatase activity in endomycorrhizal maize   总被引:3,自引:0,他引:3  
Success of a mycorrhizal symbiosis is influenced by the availability of phosphorus (P) in the soil. Maize ( Zea mays L. cv. Great Lakes 586) plants were grown under five different levels of soil P, either in the presence or absence of formononetin or the vesicular‐arbuscular mycorrhizal (VAM) fungus Glomus intraradices Schenck and Smith. We detected physiological differences in mycorrhizal roots very early in the development of symbiosis, before the onset of nutrient‐dependent responses. Under low P levels, VAM roots accumulated a greater shoot dry weight (13%), root P concentration (15%) and protein concentration (30%) than non-VAM roots, although root growth was not statistically significantly different. At higher P levels, mycorrhizal roots weighed less than non-VAM roots (10%) without a concomitant host alteration of growth or root P concentration. Mycorrhizal colonization decreased as soil P increased. Formononetin-treatment enhanced colonization of the root by G. intraradices and partially overcame inhibition of VAM colonization by high soil P concentrations. This is the first report that formononetin improves root colonization under high levels of soil P. Acid phosphatase (ACP) and alkaline phosphatase (ALP) activities were closely related to the level of fungal colonization in corn roots. ACP activity in corn roots responded more to soil P availability than did ALP activity (38% more). These results suggest that ACP was involved in the increased uptake of P from the soil, while ALP may be linked to active phosphate assimilation or transport in mycorrhizal roots. Thus, soil P directly affected a number of enzymes essential in host-endophyte interplay, while formononetin enhanced fungal colonization.  相似文献   

10.
Versaw  Wayne K.  Chiou  Tzyy-Jen  Harrison  Maria J. 《Plant and Soil》2002,244(1-2):239-245
Most vascular plants acquire phosphate from their environment either directly, via the roots, or indirectly, via a symbiotic interaction with arbuscular mycorrhizal (AM) fungi. The symbiosis develops in the plant roots where the fungi colonize the cortex of the root to obtain carbon from the plant host, while assisting the plant with acquisition of phosphate and other mineral nutrients from the soil solution. As a first step toward understanding the molecular basis of the symbiosis and phosphate utilization, we have cloned and characterized phosphate transporter genes from the AM fungi Glomus versiforme and Glomus intraradices, and from the roots of a host plant, Medicago truncatula. Expression analyses and localization studies indicate that each of these transporters has a role in phosphate uptake from the soil solution.  相似文献   

11.
Plants from the Cyperaceae family (sedges), usually considered as non-mycorrhizal, constitute almost exclusively the herbaceous stratum of the ultramafic maquis in New Caledonia. These plants are pioneers and are important for the ecological restoration of mined areas. Costularia comosa, one of the most common sedges in this environment, was grown under field conditions on ultramafic soil, fertilized or not with phosphate and/or nitrogen. Results showed that the addition of phosphate to the soil induced a clear increase in mycorrhizal colonization of C. comosa and an increase in arbuscule abundance, reflecting the establishment of a functional mycorrhizal symbiosis. Significant positive correlations were found among mycorrhizal parameters and plant or soil phosphorus concentrations. Nitrogen fertilization did not affect mycorrhizal colonization of C. comosa. The improvement in mycorrhizal colonization by phosphate fertilization did not influence significantly nickel concentrations in the roots and shoots of plants. This study demonstrated that phosphate fertilization of ultramafic soil improved mycorrhizal colonization of C. comosa, with formation of a functional symbiosis under field conditions.  相似文献   

12.
植物中丛枝菌根形成的信号途径研究进展   总被引:1,自引:0,他引:1  
丛枝菌根(arbuscular mycorrhizal,AM)共生是丛枝菌根真菌与大多数陆地植物的根系之间形成的一种互利共生关系。植物给菌根真菌提供碳水化合物;作为回报,菌根真菌能够增强植物对矿质营养元素(尤其是磷)的吸收。菌根的形成过程是一系列信号交换和转导的结果,具有严格并且一致的顺序。本文以植物中菌根形成的信号途径为主线,对菌根真菌的形成过程和信号转导途径及其方式进行了分析和讨论。高等植物中菌根形成的信号途径与豆科植物的结瘤信号途径部分共享,并且与钙离子信号途径相关,但前者更为广泛。尽管该途径中很多过程目前还不十分清楚,但是相信在不久的将来就可以揭开菌根形成过程中的众多谜团。  相似文献   

13.
丛枝菌根(arbuscular mycorrhizal, AM)共生是丛枝菌根真菌与大多数陆地植物的根系之间形成的一种互利共生关系。植物给菌根真菌提供碳水化合物; 作为回报, 菌根真菌能够增强植物对矿质营养元素(尤其是磷)的吸收。菌根的形成过程是一系列信号交换和转导的结果, 具有严格并且一致的顺序。本文以植物中菌根形成的信号途径为主线, 对菌根真菌的形成过程和信号转导途径及其方式进行了分析和讨论。高等植物中菌根形成的信号途径与豆科植物的结瘤信号途径部分共享, 并且与钙离子信号途径相关, 但前者更为广泛。尽管该途径中很多过程目前还不十分清楚, 但是相信在不久的将来就可以揭开菌根形成过程中的众多谜团。  相似文献   

14.

This review highlights the key role that mycorrhizal fungi play in making phosphorus (Pi) more available to plants, including pathways of phosphorus absorption, phosphate transporters and plant-mycorrhizal fungus symbiosis, especially in conditions where the level of inorganic phosphorus (Pi) in the soil is low. Mycorrhizal fungi colonization involves a series of signaling where the plant root exudates strigolactones, while the mycorrhizal fungi release a mixture of chito-oligosaccharides and liposaccharides, that activate the symbiosis process through gene signaling pathways, and contact between the hyphae and the root. Once the symbiosis is established, the extraradical mycelium acts as an extension of the roots and increases the absorption of nutrients, particularly phosphorus by the phosphate transporters. Pi then moves along the hyphae to the plant root/fungus interface. The transfer of Pi occurs in the apoplectic space; in the case of arbuscular mycorrhizal fungi, Pi is discharged from the arbuscular to the plant’s root symplasm, in the membrane that surrounds the arbuscule. Pi is then absorbed through the plant periarbuscular membrane by plant phosphate transporters. Furthermore, plants can acquire Pi from soil as a direct absorption pathway. As a result of this review, several genes that codify for high-affinity Pi transporters were identified. In plants, the main family is Pht1 although it is possible to find others such as Pht2, Pht3, Pho1 and Pho2. As in plants, mycorrhizal fungi have genes belonging to the Pht1 subfamily. In arbuscular mycorrhizal fungi we found L1PT1, GiPT, MtPT1, MtPT2, MtPT4, HvPT8, ZmPht1, TaPTH1.2, GmosPT and LYCes. HcPT1, HcPT2 and BePT have been characterized in ectomycorrhizal fungi. Each gene has a different way of expressing itself. In this review, we present diagrams of the symbiotic relationship between mycorrhizal fungi and the plant. This knowledge allows us to design solutions to regional problems such as food production in soils with low levels of Pi.

  相似文献   

15.
16.
Many terrestrial plant species are able to form symbiotic associations with arbuscular mycorrhizal fungi. Here we have identified three cDNA clones representing genes whose expression is induced during the arbuscular mycorrhizal symbiosis formed between Medicago truncatula and an arbuscular mycorrhizal fungus, Glomus versiforme. The three clones represent M. truncatula genes and encode novel proteins: a xyloglucan endotransglycosylase-related protein, a putative arabinogalactan protein (AGP), and a putative homologue of the mammalian p110 subunit of initiation factor 3 (eIF3). These genes show little or no expression in M. truncatula roots prior to formation of the symbiosis and are significantly induced following colonization by G. versiforme. The genes are not induced in roots in response to increases in phosphate. This suggests that induction of expression during the symbiosis is due to the interaction with the fungus and is not a secondary effect of improved phosphate nutrition. In situ hybridization revealed that the putative AGP is expressed specifically in cortical cells containing arbuscules. The identification of two mycorrhiza-induced genes encoding proteins predicted to be involved in cell wall structure is consistent with previous electron microscopy data that indicated major alterations in the extracellular matrix of the cortical cells following colonization by mycorrhizal fungi.  相似文献   

17.
Gazey C  Abbott LK  Robson AD 《Mycorrhiza》2004,14(6):355-362
Arbuscular mycorrhizal (AM) fungi occur in all agricultural soils but it is not easy to assess the contribution they make to plant growth under field conditions. Several approaches have been used to investigate this, including the comparison of plant growth in the presence or absence of naturally occurring AM fungi following soil fumigation or application of fungicides. However, treatments such as these may change soil characteristics other than factors directly involving AM fungi and lead to difficulties in identifying the reason for changes in plant growth. In a glasshouse experiment, we assessed the contribution of indigenous AM fungi to growth of subterranean clover in undisturbed cores of soil from two agricultural field sites (a cropped agricultural field at South Carrabin and a low input pasture at Westdale). We used the approach of estimating the benefit of AM fungi by comparing the curvature coefficients ( C) of the Mitscherlich equation for subterranean clover grown in untreated field soil, in field soil into which inoculum of Glomus invermaium was added and in soil fumigated with methyl bromide. It was only possible to estimate the benefit of mycorrhizas using this approach for one soil (Westdale) because it was the only soil for which a Mitscherlich response to the application of a range of P levels was obtained. The mycorrhizal benefit ( C of mycorrhizal vs. non-mycorrhizal plants or C of inoculated vs. uninoculated plants) of the indigenous fungi corresponded with a requirement for phosphate by plants that were colonised by AM fungi already present in the soil equivalent to half that required by non-mycorrhizal plants. This benefit was independent of the plant-available P in the soil. There was no additional benefit of inoculation on plant growth other than that due to increased P uptake. Indigenous AM fungi were present in both soils and colonised a high proportion of roots in both soils. There was a higher diversity of morphotypes of mycorrhizal fungi in roots of plants grown in the Westdale soil than in the South Carrabin soil that had a history of high phosphate fertilizer use in the field. Inoculation with G. invermaium did not increase the level of colonisation of roots by mycorrhizal fungi in either soil, but it replaced approximately 20% of the root length colonised by the indigenous fungi in Westdale soil at all levels of applied P. The proportion of colonised root length replaced by G. invermaium in South Carrabin soil varied with the level of application of P to the soil; it was higher at intermediate levels of recently added soil P.  相似文献   

18.
Abstract

Legume plants enter two important endosymbioses – with soil fungi, forming phosphorus acquiring arbuscular mycorrhiza (AM), and with nitrogen-fixing bacteria, leading to the formation of nitrogen-fixing root nodules. Both symbioses have been studied extensively because these symbioses have great potential for agricultural applications. Although 80% of all living land plants form AM, the nitrogen-fixing root nodule symbiosis with rhizobia is almost exclusively restricted to legumes. Despite varying degree of differences in the morphological responses induced by both endosymbionts in the host plants, significant similarities in the development of both fungal and bacterial symbioses have been reported. The signal perception and signal transduction cascades that initiate nodulation and mycorrhization in legumes partially overlap. Legume genes have been identified that are required for the establishment of both AM and root nodule symbiosis and are referred to as the common SYM genes. Genetic dissection of the common SYM signal transduction pathway required for bacterial and fungal root endosymbiosis has not only unraveled the players involved but also provided a first glimpse at conservation and specialization of signaling cascades essential for nodulation and mycorrhiza development. Based on the observation of common signaling cascades, it is tempting to speculate that the root nodule symbiosis, where fossil records date back to the late Cretaceaous, adopted and subsequently modified more ancient signal transduction pathways leading to AM formation, having already been in place 400 million years ago. This review discusses the common aspects of recognition of mycorrhizal fungi and Rhizobium by the host, and further signal transduction that leads to an effective symbiosis.  相似文献   

19.
Most plant species form symbioses with arbuscular mycorrhizal (AM) fungi, which facilitate the uptake of mineral nutrients such as phosphate from the soil. Several transporters, particularly proton-coupled phosphate transporters, have been identified on both the plant and fungal membranes and contribute to delivering phosphate from fungi to plants. The mechanism of nutrient exchange has been studied in plants during mycorrhizal colonization, but the source of the electrochemical proton gradient that drives nutrient exchange is not known. Here, we show that plasma membrane H+-ATPases that are specifically induced in arbuscule-containing cells are required for enhanced proton pumping activity in membrane vesicles from AM-colonized roots of rice (Oryza sativa) and Medicago truncatula. Mutation of the H+-ATPases reduced arbuscule size and impaired nutrient uptake by the host plant through the mycorrhizal symbiosis. Overexpression of the H+-ATPase Os-HA1 increased both phosphate uptake and the plasma membrane potential, suggesting that this H+-ATPase plays a key role in energizing the periarbuscular membrane, thereby facilitating nutrient exchange in arbusculated plant cells.  相似文献   

20.
Soil nutrient availability and colonization by arbuscular mycorrhizal fungi are important and potentially interacting factors shaping vegetation composition and succession. We investigated the effect of carbon (C) addition, aimed at reducing soil nutrient availability, on arbuscular mycorrhizal colonization. Seedlings of 27 plant species with different sets of life-history traits (functional group affiliation, life history strategy and nitrophilic status) were grown in pots filled with soil from a nutrient-rich set-aside field and amended with different amounts of C. Mycorrhizal colonization was progressively reduced along the gradient of increasing C addition in 17 out of 27 species, but not in the remaining species. Grasses had lower colonization levels than forbs and legumes and the decline in AM fungal colonization was more pronounced in legumes than in other forbs and grasses. Mycorrhizal colonization did not differ between annual and perennial species, but decreased more rapidly along the gradient of increasing C addition in plants with high Ellenberg N values than in plants with low Ellenberg N values. Soil C addition not only limits plant growth through a reduction in available nutrients, but also reduces mycorrhizal colonization of plant roots. The effect of C addition on mycorrhizal colonization varies among plant functional groups, with legumes experiencing an overproportional reduction in AM fungal colonization along the gradient of increasing C addition. We therefore propose that for a better understanding of vegetation succession on set-aside fields one may consider the interrelationship between plant growth, soil nutrient availability and mycorrhizal colonization of plant roots.  相似文献   

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