Phosphorus and intraspecific density alter plant responses to arbuscular mycorrhizas

We investigated how phosphorus availability, intraspecific density, and their interaction affect plant responses to arbuscular mycorrhizas. Four facultatively mycotrophic species: chile, cilantro, tomato, and corn were examined separately in pot experiments that employed a tropical phosphorus-immobilizing soil. Each experiment comprised nine soluble phosphorus additions, two levels of intraspecific plant density, and inoculation with arbuscular mycorrhizal fungi or not. High phosphorus signi- ficantly diminished mycorrhizal colonization of corn, cilantro, and tomato, but not chile, which was highly variably colonized. Corn roots were colonized by other root-inhabiting fungi, and mycorrhizas significantly reduced colonization by these potential root parasites. High phosphorus significantly increased relative growth rates (RGR) of all species, and high density significantly reduced RGR of cilantro, tomato, and corn. Chile showed little growth at any but the highest phosphorus additions, and consequently had no RGR response to density or mycorrhizas. Mycorrhiza inoculation caused transient depression of corn growth during the first month, but mycorrhizas increased corn RGR during the second month of growth. Both RGR and dry weights at harvest, cilantro, tomato, and corn benefited from mycorrhizas at low phosphorus availability, but this benefit diminished or changed to disadvantage as phosphorus availability increased. At low phosphorus availability, high density increased the dry weight of mycorrhizal cilantro and thereby amplified the benefit of mycorrhizas. At high phosphorus availability, increased density diminished the effects of mycorrhizas on dry weight, reducing mycorrhiza benefit to tomato and chile and reducing mycorrhiza detriment to cilantro. This study demonstrates that for three of the four plant species examined, phosphorus availability, intraspecific density, and their interaction significantly modify plant responses to arbuscular mycorrhizas.     

Anatomy and ultrastructure of mycorrhizal associations of neotropical Ericaceae

Ericaceae are obligatory associated with symbiotic fungi forming several, distinctive categories of mycorrhizas. While ericoid, arbutoid, and monotropoid mycorrhizas are known since many years from ericads of the northern hemisphere and the ericoid mycorrhiza also from Australia, a further mycorrhizal category with hyphal sheath, Hartig net, and intracellular colonization was described by us recently and termed cavendishioid mycorrhiza because it was found on Cavendishia nobilis, a species belonging to the Andean clade (Vaccinioideae) of Ericaceae. As the previous findings indicated a correlation between the mycorrhizal category and the systematic position of Ericaceae, we tested the hypothesis that other ericads of the Andean clade might also form cavendishioid mycorrhizas, while ericads occurring in the same area but not belonging to the Andean clade might not. Mycorrhizas of 20 different ericaceous species, 15 belonging to the Andean clade and 5 to other Vaccinioideae or Ericoideae, were sampled in the tropical mountain rain forest area of South Ecuador and investigated by light and electron microscopy. All the 15 members of the Andean clade ericads displayed a hyphal sheath, as well as inter- and intracellular colonization by hyphae as was found on Cavendishia previously. The five species not belonging to the Andean clade ericads displayed only intracellular colonization by hyphae and hence were typical ericoid mycorrhizal. Ultrastructural studies revealed Sebacinales and ascomycetes as mycorrhiza formers in both associations even within one single cell. The results thus support the hypothesis that the Andean clade of Ericaceae forms mycorrhizas distinct from the arbutoid category and most likely presents an independent evolutionary line in the Ericaceae derived from the ericoid mycorrhizas, justifying the new term “cavendishioid mycorrhiza”.     

Paris-type mycorrhizas in Smilax aspera L. growing in a Mediterranean scherophyllous wood

Paris- type mycorrhiza is described in Smilax aspera L., an evergreen climbing plant of Mediterranean sclerophyllous woods. Wild plants were sampled from a protected area inside the Regional Natural Park Migliarino-S.Rossore-Massaciuccoli, on the northwestern coast of Italy, near Pisa. Mycorrhizas formed by S. aspera were identified as a variation of Paris-type arbuscular mycorrhizas. Detailed observations on stained roots and on fresh root sections showed that, after forming the appressorium, the fungus colonized the root by penetrating individual cells, growing intracellularly from cell to cell, and producing many coils and terminal arbuscules. S. aspera seedlings inoculated with the arbuscular mycorrhizal fungi Glomus mosseae and G. viscosum, which are known to form Arum-type mycorrhizas in many plant species, produced the same Paris-type-like mycorrhizas found in nature. This confirms that the type of arbuscular mycorrhizal infection is largely governed by the plant host genotype. Plants of S. aspera inoculated with G. mosseae and G. viscosum had larger growth increments than uninoculated plants. Thus Paris-type mycorrhizas produce growth responses comparable to those of Arum-type mycorrhizas. Accepted: 11 January 2000     

基于SCI文献分析我国菌根学研究现状和发展方向

基于美国科学情报研究所(ISI)科学引文索引(Science Citation Index, SCI)数据库, 对1989~2007年期间我国发表的菌根研究论文进行检索, 并采用文献计量方法对所获资料进行统计和趋势分析。结果表明, 1989~2007年间, 我国菌根学研究呈现不断上升趋势, 尤其是2000年之后, 增长速度明显加快。但只有5.22%的论文发表于影响因子大于5的刊物, 在研究深度上仍需进一步加强。半数以上的研究论文与丛枝菌根研究有关, 研究重点主要集中在菌根对植物的生理效应, 菌根与植物抗性     

Phylogenetic distribution and evolution of mycorrhizas in land plants

A survey of 659 papers mostly published since 1987 was conducted to compile a checklist of mycorrhizal occurrence among 3,617 species (263 families) of land plants. A plant phylogeny was then used to map the mycorrhizal information to examine evolutionary patterns. Several findings from this survey enhance our understanding of the roles of mycorrhizas in the origin and subsequent diversification of land plants. First, 80 and 92% of surveyed land plant species and families are mycorrhizal. Second, arbuscular mycorrhiza (AM) is the predominant and ancestral type of mycorrhiza in land plants. Its occurrence in a vast majority of land plants and early-diverging lineages of liverworts suggests that the origin of AM probably coincided with the origin of land plants. Third, ectomycorrhiza (ECM) and its derived types independently evolved from AM many times through parallel evolution. Coevolution between plant and fungal partners in ECM and its derived types has probably contributed to diversification of both plant hosts and fungal symbionts. Fourth, mycoheterotrophy and loss of the mycorrhizal condition also evolved many times independently in land plants through parallel evolution.     

Environmental modulation of plant mycorrhizal traits in the global flora

Mycorrhizal symbioses are known to strongly influence plant performance, structure plant communities and shape ecosystem dynamics. Plant mycorrhizal traits, such as those characterising mycorrhizal type (arbuscular (AM), ecto-, ericoid or orchid mycorrhiza) and status (obligately (OM), facultatively (FM) or non-mycorrhizal) offer valuable insight into plant belowground functionality. Here, we compile available plant mycorrhizal trait information and global occurrence data ($\sim$100 million records) for 11,770 vascular plant species. Using a plant phylogenetic mega-tree and high-resolution climatic and edaphic data layers, we assess phylogenetic and environmental correlates of plant mycorrhizal traits. We find that plant mycorrhizal type is more phylogenetically conserved than plant mycorrhizal status, while environmental variables (both climatic and edaphic; notably soil texture) explain more variation in mycorrhizal status, especially FM. The previously underestimated role of environmental conditions has far-reaching implications for our understanding of ecosystem functioning under changing climatic and soil conditions.     

Mycorrhizas and soil structure

In addition to their well-recognized roles in plant nutrition and communities, mycorrhizas can influence the key ecosystem process of soil aggregation. Here we review the contribution of mycorrhizas, mostly focused on arbuscular mycorrhizal fungi (AMF), to soil structure at various hierarchical levels: plant community; individual root; and the soil mycelium. There are a suite of mechanisms by which mycorrhizal fungi can influence soil aggregation at each of these various scales. By extension of these mechanisms to the question of fungal diversity, it is recognized that different species or communities of fungi can promote soil aggregation to different degrees. We argue that soil aggregation should be included in a more complete 'multifunctional' perspective of mycorrhizal ecology, and that in-depth understanding of mycorrhizas/soil process relationships will require analyses emphasizing feedbacks between soil structure and mycorrhizas, rather than a uni-directional approach simply addressing mycorrhizal effects on soils. We finish the discussion by highlighting new tools, developments and foci that will probably be crucial in further understanding mycorrhizal contributions to soil structure.     

Mycorrhizas influence functional traits of two tallgrass prairie species

Over the past decade, functional traits that influence plant performance and thus, population, community, and ecosystem biology have garnered increasing attention. Generally lacking, however, has been consideration of how ubiquitous arbuscular mycorrhizas influence plant allometric and stoichiometric functional traits. We assessed how plant dependence on and responsiveness to mycorrhizas influence plant functional traits of a warm‐season, C4 grass, Andropogon gerardii Vitman, and the contrasting, cool‐season, C3 grass, Elymus canadensis L. We grew both host species with and without inoculation with mycorrhizal fungi, across a broad gradient of soil phosphorus availabilities. Both host species were facultatively mycotrophic, able to grow without mycorrhizas at high soil phosphorus availability. A. gerardii was most dependent upon mycorrhizas and E. canadensis was weakly dependent, but highly responsive to mycorrhizas. The high dependence of A. gerardii on mycorrhizas resulted in higher tissue P and N concentrations of inoculated than noninoculated plants. When not inoculated, E. canadensis was able to take up both P and N in similar amounts to inoculated plants because of its weak dependence on mycorrhizas for nutrient uptake and its pronounced ability to change root‐to‐shoot ratios. Unlike other highly dependent species, A. gerardii had a high root‐to‐shoot ratio and was able to suppress colonization by mycorrhizal fungi at high soil fertilities. E. canadensis, however, was unable to suppress colonization and had a lower root‐to shoot ratio than A. gerardii. The mycorrhiza‐related functional traits of both host species likely influence their performance in nature: both species attained the maximum responsiveness from mycorrhizas at soil phosphorus availabilities similar to those of tallgrass prairies. Dependence upon mycorrhizas affects performance in the absence of mycorrhizas. Responsiveness to mycorrhizal fungi is also a function of the environment and can be influenced by both mycorrhizal fungus species and soil fertility.     

Asymmetric symbiont adaptation to Arctic conditions could explain why high Arctic plants are non-mycorrhizal

Mycorrhizal symbiosis generally improves nutrient and water acquisition of the host plant. Furthermore, mycorrhizal fungi affect plant herbivory and pathogen resistance. The symbiotic condition of land plants is evolutionarily ancient and the functions performed by the fungal symbiont are thought to be pivotal to successful plant life. Although most land plants are mycorrhizal, the extreme high Arctic habitats are dominated by plant species and genera characteristically free of mycorrhiza. In this paper, previous attempts to explain the lack of mycorrhizas in high Arctic are reviewed as well as a new idea of asymmetric symbiont adaptation is proposed as potential explanation why high Arctic plants are non-mycorrhizal.     

The significance of host-fungus combinations in ectomycorrhizal symbioses for the chemical quality of willow foliage

The majority of plants interacts with mycorrhizal fungi, which predominantly provide mutual benefits, but also costs. We tested the hypothesis that specific combinations of host plants (four commercial varieties of Salix spp.) and ectomycorrhizal partners (species of Laccaria, Paxillus, Tricholoma and Hebeloma) differ in their effects on the host foliar chemistry. Twenty specific host - mycorrhiza combinations were pot-grown outdoors under low-N conditions. Foliar concentrations of total phenolics and salicylic acid were decreased by mycorrhizas in S. schwerinii x S. viminalis and S. x dasyclados genotypes, but increased in S. viminalis. Mycorrhiza effects on host biomass production were positive, indifferent or negative, depending on the specific host—mycorrhiza combination. The host plant genotype influenced mainly the direction of mycorrhizal effects on foliar chemistry, whereas the magnitude of mycorrhizal effects varied mostly among the fungal genotypes. The results are relevant with respect to the possible interactions between mycorrhizas, plants and leaf herbivores.     

An economic approach to evaluate the role of mycorrhizas in managed ecosystems

The benefits of management of mycorrhizas in agricultural and horticultural croppiing systems remains problematic except where the indigenous fungal population is low. Most experiments have focused on the introduction of exotic fungal isolates. Promotion of plant growth by mycorrhizas can be enhanced by increasing the effectiveness of the indigenous fungi as well as by introducing more effective species. Lack of reliable methods for identification of fungal species colonizing roots is a major limitation to characterizing the change in mycorrhizal populations. Assessment of the role of mycorrhizas in commerical food production systems must include an economic analysis. To do so requires an evaluation of the response to increasing the effectiveness of the mycorrhizal symbiosis relative to increasing yield with addition of phosphorus fertilizer. Thus field experiments should be designed to measure the response to phosphorus addition with the existing mycorrhizal population as well as with the more effectively managed population. In this paper we discuss changes that may be induced in mycorrhizal fungi by management to increase their effectiveness in promoting plant growth. We then suggest an economic analysis approach to assessing the potential benefits of this increase in effectiveness. We conclude with a discussion of research approaches needed to determine, in a more objective manner, the role of mycorrhizas in managed ecosystems.     

Progress and Challenges in Agricultural Applications of Arbuscular Mycorrhizal Fungi

Arbuscular mycorrhiza is a widespread interaction between plant roots and mutualistic fungi that both promotes plant mineral nutrition and supports stability in many natural ecosystems. Given the demand in current agriculture for these ecosystem services, it appears desirable to further integrate this interaction in current management schemes. While two different approaches can be identified in this regard, a number of problems have limited the agricultural application of mycorrhiza to date. In this review, the sensitivity of arbuscular mycorrhizal fungi to certain agricultural management methods, the functional variability of arbuscular mycorrhizal fungi, and the variability of plant responsiveness will be discussed in particular. In addition we will focus on recent advances in the study of mycorrhiza in natural ecosystems, and illustrate how such knowledge might be used to further the integration of arbuscular mycorrhizal fungi in agricultural regimes, and so increase the sustainability and robustness of current practice.     

Occurrence of arbuscular mycorrhizal fungi in a phosphorus-poor wetland and mycorrhizal response to phosphorus fertilization

The presence of arbuscular mycorrhizas in fens has received little attention, but because fen plants are often phosphorus limited, the plant-fungus interaction could be an important factor in plant competition for phosphorus. In this field study, we determined mycorrhizal colonization rates for 18 fen plant species. Also in the field, we examined the effect of four different forms of phosphorus on the percentage colonization for one fen plant species, Solidago patula. We found that in a species-rich, phosphorus-poor wetland both mycorrhizal and nonmycorrhizal species were common. Nine of ten dicotyledonous species examined formed arbuscular mycorrhizas, while all monocotyledonous species were at most very weakly mycorrhizal. A morphological explanation for this pattern is that the monocots in our study have more extensive aerenchyma, especially in coarse roots. Therefore, monocots are able to transport oxygen to their roots more effectively than dicots. In the organic wetland soil, additional oxygen in the rhizosphere promotes phosphorus mineralization and availability. Two of the monocot species (Typha latifolia and Carex lasiocarpa), which have been described previously as mycorrhizal in other wetland types, are surprisingly nonmycorrhizal in our phosphorus-poor study site, suggesting that a mycorrhizal association would not offer improved phosphorus nutrition to these species. In contrast, our field phosphorus addition decreased mycorrhizal colonization in S. patula, suggesting that one benefit to S. patula of the mycorrhizas is phosphorus uptake.     

Cytoskeleton in mycorrhizal symbiosis

An understanding of the role played by the cytoskeleton in formation and function of mycorrhizas has been hampered by the technical difficulty of working with mycorrhizal material. Recently, however, improved labelling techniques suitable for both plant and fungal symbionts in combination with either epifluorescence microscopy or laser scanning confocal microscopy have resulted in new information. As well, molecular methods have made it possible to monitor changes of cytoskeletal elements during mycorrhiza development. Currently we know that the cytoskeletal systems of both plant and fungal partners undergo changes during both ecto- and endomycorrhizal symbiosis. However, little information is available concerning the regulatory factors or the cause and effect relationship of cytoskeletal changes and cellular events. In this article, research involving the cytoskeleton of mycorrhizas is reviewed in detail, whereas basic information of the cytoskeleton of plant and fungal cells is only briefly discussed as background. A brief comparison is also made between the information on mycorrhizas with that of biotrophic pathogenic fungi and the Rhizobium–legume symbiosis.     

Micronutrient transport in mycorrhizal symbiosis; zinc steals the show

Mycorrhizas are mutually beneficial associations between soil-borne fungi and plant roots. Mycorrhizal fungi provide their host plant with essential nutrients in exchange for sugars and/or lipids. Traditionally, transport and translocation of macronutrients, including nitrogen and phosphorus, throughout the fungal mycelium and towards the host plant are well studied. However, the regulation of nutrient exchange and their contribution in the morphogenesis and development of mycorrhizas remains unclear. In this Opinion, we argue that adding micronutrients in the current models of symbiotic transport is essential to fully understand the establishment, maintenance, and functioning of mycorrhizal associations. Homeostatic mechanisms at the cellular level and the first transport proteins involved have been recently documented for zinc (Zn) in arbuscular mycorrhizal, ectomycorrhizal, and ericoid mycorrhizal fungi. Mycorrhizal plants benefit from an improved Zn status in control conditions and are better protected when environmental Zn availability fluctuates. These recent progresses are paving the way for a better understanding of micronutrient allocation in mycorrhizas. Revising our vision on the role of micronutrients, particularly of Zn, in these interactions will allow a better use of mycorrhizal fungi in sustainable agriculture and forestry, and will increase management practices in waste land, as well as in agricultural and natural ecosystems.     

Arbuscular mycorrhizas modify plant responses to soil zinc addition

Zinc deficiency is one of the most commonly reported plant and human nutrient deficiencies worldwide. Conversely, Zn is also a common environmental contaminant, significantly reducing plant growth. These contrasting effects of Zn on plant growth and nutrition have been the focus of a considerable number of studies; however, most studies focus on plant responses to soil Zn concentration under either deficient or toxic concentrations, but not both. The formation of arbuscular mycorrhizas (AM) can increase plant Zn uptake under low soil Zn concentrations, and on the other hand, ‘protect’ plants against excessive Zn accumulation under high soil Zn conditions. Here we report the findings of an experiment in which we studied the response of AM formed by tomatoes under low, medium and high soil Zn conditions. To control for the formation of AM in this study we used a mycorrhiza defective tomato mutant and its mycorrhizal wildtype progenitor. While mycorrhizal colonization was not significantly impacted by soil Zn addition, the growth of plants and tissue Zn concentrations were. Together these data highlight the complex interactions between AM and Zn, and the utility of a genotypic approach for studying AM in this context.     

Mapping the potential mycorrhizal associations of the conterminous United States of America

Mycorrhizal associations are recognized as key symbioses in a changing world, yet our understanding of their geographic distribution and temporal dynamics remains limited. We combined data on mycorrhizal associations and historical dominant vegetation to map the pre-European Settlement mycorrhizal associations of the conterminous United States of America (USA). As a demonstration of the map's utility, we estimated changes in mycorrhizal associations due to urbanization, agriculture and the establishment of non-native species in two regions. We found that the conterminous USA was dominated by vegetation associated with arbuscular mycorrhizas, but that ∼40% of vegetation types included multiple mycorrhizal associations. Shifting land use to agriculture and the introduction of non-native species has disproportionately affected ectomycorrhizas, as did urbanization. These preliminary results set a baseline for mycorrhizal biogeography of the USA and illustrate how synthesis of available data can help us understand the impact of anthropogenic changes on an important mutualism.     

Dynamics of cytoskeletal proteins in developing pine ectomycorrhiza

 Mycorrhizal short roots of Pinus contorta Dougl. ex Loud colonized by Suillus variegatus (Sow. ex Fr.) O. Kuntze or Paxillus involutus (Batsch) Fr. were collected 1–>60 days after fungal contact. The proteins of the inoculated roots were extracted, electrophoretically separated, blotted and immunostained for α-tubulin and actin. The development of the mycorrhiza was also followed microscopically. The signal of plant α-tubulin was stronger than the signal of fungal α-tubulin during the first 5 days in S. variegatus mycorrhiza and was then exceeded by fungal α-tubulin. This correlated well with the increase of fungal mycelium in the mycorrhiza. A transient drop in both plant and fungal α-tubulin signals was observed 20 days after fungal contact, suggesting a change in the metabolism of the mycorrhiza. The signals for plant and fungal actins in the mycorrhiza increased steadily during early infection and then remained at a high level as the mycorrhiza matured. Similar trends were observed in P. contorta–P. involutus mycorrhiza. The data from P. contorta–S.variegatus mycorrhizas suggests that α-tubulin is a growth-related protein, subject to changes, while the amount of actin reflects the general metabolic activity of the mycorrhiza. In both mycorrhizal systems clear α-tubulin and actin signals were detected 60 days after colonization, which indicates that the mycorrhizas were metabolically active in spite of their withered appearance. Accepted: 6 May 1996