Investment in Fine Roots and Arbuscular Mycorrhizal Fungi Decrease During Succession in Three Brazilian Ecosystems

The functional groups of plants that characterize different phases of succession are expected to show differences in root distribution, fine‐root traits and degrees of association with arbuscular mycorrhizal (AM) fungi. The relationship involving fine‐root traits and AM fungi that regulate the nutrient acquisition potential among different plant functional groups are still not well understood. We assessed fine‐root morphology, AM fungal variables and soil fertility in grassland, secondary forest and mature forest in Atlantic, Araucaria and Pantanal ecosystems in Brazil. Soil cores were collected at 0–10 and 10–20 cm depths. Fine roots were extracted from soil by sieving and root morphological traits and AM colonization were determined. The AM spores were extracted from soil and counted. In all ecosystems, soil fertility, fine‐root mass and root diameter increased with the succession, while root length, specific root length, root‐hair length, root‐hair incidence, AM colonization and AM spore density decreased. These results suggest that plant species from early stages of tropical succession with inherent rapid growth invest in fine roots and maintain a high degree of AM colonization in order to increase the capacity for nutrient acquisition. Conversely, fine root morphological characteristics and low degree of AM colonization exhibited by plants of the later stages of succession lead toward a low nutrient uptake capacity that combine with their typical low growth rates. Abstract in Portuguese is available at http://www.blackwell‐synergy.com/loi/btp .     

Changes in arbuscular mycorrhizal associations and fine root traits in sites under different plant successional phases in southern Brazil

Fine root morphological traits and distribution, arbuscular mycorrhizal (AM) fungi, soil fertility, and nutrient concentration in fine root tissue were compared in sites under different successional phases: grass plants, secondary forest, and mature forest in Londrina county, Paraná state, southern Brazil. Soil cores were collected randomly at the 0-10- and 10-20-cm depths in three quadrants (50 m2) in each site. Plants from the different successional stages displayed high differences in fine root distribution, fine root traits, and mycorrhizal root colonization. There were increases in the concentration of nutrients both in soil and fine roots and decrease of bulk soil density along the succession. The fine root biomass and diameter increased with the succession progress. The total fine root length, specific root length, root hair length, and root hair incidence decreased with the succession advance. Similarly, the mycorrhizal root colonization and the density of AM fungi spores in the soil decreased along the succession. Mycorrhizal root colonization and spore density were positively correlated with fine root length, specific root length, root hair length, root hair incidence, and bulk density and negatively correlated with fine root diameter and concentration of some nutrients both in soil and root tissues. Nutrient concentration in root tissue and in soil was positively correlated with fine root diameter and negatively correlated with specific root length, root hair length, and root hair incidence. These results suggest different adaptation strategies of plant roots for soil exploration and mineral acquisition among the different successional stages. Early successional stages displayed plants with fine root morphology and AM fungi colonization to improve the root functional efficiencies for uptake of nutrients and faster soil resource exploration. Late successional stages displayed plants with fine root morphology and mycorrhizal symbiosis for both a lower rate of soil proliferation and soil exploration capacity to acquire nutrients.     

Effect of drought and season on arbuscular mycorrhizal fungi in a subtropical secondary forest

Arbuscular mycorrhizal (AM) fungi in both soil and roots were examined in May (summer) and December (winter) under a 4-y drought experiment in a Chinese subtropical secondary forest. Drought significantly decreased AM fungal extra-radical hyphal density, spore density, and root colonization rate in both seasons. These AM parameters were significantly higher in summer than in winter in the control treatment, but only AM fungal extra-radical hyphal density exhibited the same seasonal trend in the drought treatment. In total, 45 AM fungal operational taxonomic units (OTUs) were obtained at a 97% sequence similarity level using Illumina sequencing of 18S rDNA. Drought and season had no significant effects on AM fungal OTU richness in soil and roots. AM fungal community composition in soil and roots was significantly affected by season but not by drought. This finding enhances our understanding of the response of AM fungi to global climate change in subtropical forest ecosystems.     

Mycorrhiza of plants in different vegetation types in tropical ecosystems of Xishuangbanna,southwest China

We examined plants growing in four tropical vegetation types (primary forest, secondary forest, limestone forest and a slash and burn field) in Xishuangbanna, southwest China for mycorrhizal associations. Of the 103 plant species examined (belonging to 47 families), 81 had arbuscular mycorrhizal (AM) associations, while three species possessed orchid mycorrhiza. AM colonization levels ranged between 6% and 91% and spore numbers ranged between 1.36 spores and 25.71 spores per 10 g soil. Mean AM colonization level was higher in primary and secondary forest species than in plant species from limestone forests and a slash and burn field. In contrast, mean AM fungal spore numbers of the primary and limestone forest were lower than in the secondary forest or the slash and burn field. AM fungal spores belonging to Glomus and Acaulospora were the most frequent in soils of Xishuangbanna. AM fungal colonization and spore numbers were significantly correlated to each other and were significantly influenced by vegetation type.     

Relationships among soil properties, plant nutrition and arbuscular mycorrhizal fungi–plant symbioses in a temperate grassland along hydrologic, saline and sodic gradients

Temporal variations in the relationships among plant nutrient concentrations, soil properties and arbuscular-mycorrhizal (AM) fungal dynamics were studied along a topographic and saline gradient in a temperate grassland soil. Soil and plant ( Lotus tenuis , Paspalum vaginatum , Stenotaphrum secundatum ) samples were collected on four seasonally based occasions. The morphology of AM root colonization had a similar pattern in the plants studied. Maximum arbuscular colonization occurred at the beginning of the growing season in late winter and was minimal in late summer, but maximal vesicular colonization occurred in summer and was minimal in winter, suggesting a preferential production of these morphological phases by the fungus with respect to season. The greatest arbuscular colonization was associated with the highest N and P concentrations in plant tissue, suggesting a correspondence with increases in the rate of nutrient transfer between the symbiotic partners. Water content, salinity and sodicity in soil were positively associated with AM root colonization and arbuscule colonization in L. tenuis , but negatively so in the grasses. There were distinct seasonally related effects with respect to both spore density and AM colonization, which were independent of particular combinations of plant species and soil sites.     

Relationships among soil properties, plant nutrition and arbuscular mycorrhizal fungi-plant symbioses in a temperate grassland along hydrologic, saline and sodic gradients

Temporal variations in the relationships among plant nutrient concentrations, soil properties and arbuscular-mycorrhizal (AM) fungal dynamics were studied along a topographic and saline gradient in a temperate grassland soil. Soil and plant (Lotus tenuis, Paspalum vaginatum, Stenotaphrum secundatum) samples were collected on four seasonally based occasions. The morphology of AM root colonization had a similar pattern in the plants studied. Maximum arbuscular colonization occurred at the beginning of the growing season in late winter and was minimal in late summer, but maximal vesicular colonization occurred in summer and was minimal in winter, suggesting a preferential production of these morphological phases by the fungus with respect to season. The greatest arbuscular colonization was associated with the highest N and P concentrations in plant tissue, suggesting a correspondence with increases in the rate of nutrient transfer between the symbiotic partners. Water content, salinity and sodicity in soil were positively associated with AM root colonization and arbuscule colonization in L. tenuis, but negatively so in the grasses. There were distinct seasonally related effects with respect to both spore density and AM colonization, which were independent of particular combinations of plant species and soil sites.     

荒漠植物根际AM真菌的空间分布和定殖

 通过分析以色列荒漠地区Zygophyllum dumosum, Hammada scoparia, Artemisia herba-alba 和 Atriplex halimus 等4种灌木根际AM真菌的空间分布和定殖程度,研究了AM真菌分布和定殖与植物种类和土壤因子间的相关性。样品分别从0～10 cm,10～20 cm,20～30 cm,30～40 cm和40～50 cm等5个土层中采取,土样过2 mm筛。收集的根样切成1 cm根段,经染色后,根据感染长度确定AM真菌不同结构的定殖率;用湿筛倾析法和蔗     

Fungal root colonization responses in natural grasslands after long-term exposure to elevated atmospheric CO2

Arbuscular mycorrhizae, ubiquitous mutualistic symbioses between plant roots and fungi in the order Glomales, are believed to be important controllers of plant responses to global change, in particular to elevated atmospheric CO₂. In order to test if any effects on the symbiosis can persist after long-term treatment, we examined root colonization by arbuscular mycorrhizal (AM) and other fungi of several plant species from two grassland communities after continuous exposure to elevated atmospheric CO₂ for six growing seasons in the field. For plant species from both a sandstone and a serpentine annual grassland there was evidence for changes in fungal root colonization, with changes occurring as a function of plant host species. We documented decreases in percentage nonmycorrhizal fungal root colonization in elevated CO₂ for several plant species. Total AM root colonization (%) only increased significantly for one out of the five plant species in each grassland. However, when dividing AM fungal hyphae into two groups of hyphae (fine endophyte and coarse endophyte), we could document significant responses of AM fungi that were hidden when only total percentage colonization was measured. We also documented changes in elevated CO₂ in the percentage of root colonized by both AM hyphal types simultaneously. Our results demonstrate that changes in fungal root colonization can occur after long-term CO₂ enrichment, and that the level of resolution of the study of AM fungal responses may have to be increased to uncover significant changes to the CO₂ treatment. This study is also one of the first to document compositional changes in the AM fungi colonizing roots of plants grown in elevated CO₂. Although it is difficult to relate the structural data directly to functional changes, possible implications of the observed changes for plant communities are discussed.     

Arbuscular mycorrhizas of plants growing in the Western Ghats region, Southern India

A survey of the arbuscular mycorrhizal (AM) status of plants growing in the Western Ghats region of Southern India was undertaken. Root and soil samples of plants growing in the four vegetation types forest, grassland, scrub, and cultivated land or plantation were examined. Of the 329 species (representing 61 families) examined, 174 were mycorrhizal. AM association was recorded in 81 species for the first time, including species from several families assumed to be non-mycorrhizal, e.g. Amaranthaceae, Capparaceae, Commelinaceae, Cyperaceae and Portulacaceae. AM fungal spores of 35 species belonging to Acaulospora, Gigaspora, Glomus, Sclerocystis and Scutellospora were recorded. AM fungal species richness was found to be highest in scrub and lowest in agricultural and plantation soils. Mean colonization levels were dependent on plant life-form, life-cycle pattern and vegetation type. Accepted: 26 October 1999     

Dynamics of vesicular-arbuscular mycorrhizae during old field succession

Summary The species composition of vesicular-arbuscular mycorrhizal (VAM) fungal communities changed during secondary succession of abandoned fields based on a field to forest chronosequence. Twenty-five VAM fungal species were identified. Seven species were clearly early successional and five species were clearly late successional. The total number of VAM fungal species did not increase with successional time, but diversity as measured by the Shannon-Wiener index tended to increase, primarily because the community became more even as a single species, Glomus aggregatum, became less dominant in the older sites. Diversity of the VAM fungal community was positively correlated with soil C and N. The density of VAM fungi, as measured by infectivity and total spore count, first increased with time since abandonment and then decreased in the late successional forest sites. Within 12 abandoned fields, VAM fungal density increased with increasing soil pH, H₂O soluble soil C, and root biomass, but was inversely related to extractable soil P and percent cover of non-host plant species. The lower abundance of VAM fungi in the forest sites compared with the field sites agrees with the findings of other workers and corresponds with a shift in the dominant vegetation from herbaceous VAM hosts to woody ectomycorrhizal hosts.     

Changes in arbuscular mycorrhizal fungus community along an exotic plant Eupatorium adenophorum invasion in a chinese secondary forest

Knowledge of the changes in arbuscular mycorrhizal (AM) fungi is fundamental for understanding the success of exotic plant invasions in natural ecosystems. In this study, AM fungal colonization and spore community were examined along an invasive gradient of the exotic plant Eupatorium adenophorum in a secondary forest in southwestern China. With increasing E. adenophorum invasion, the density of arbuscules in the roots of E. adenophorum significantly increased, but the AM root colonization rate and the densities of vesicles and hyphal coils in roots of E. adenophorum were not significantly different. A total of 29 AM fungi belonging to nine genera were identified based on spore morphology. Claroideoglomus etunicatum, Funneliformis geosporus, and Glomus aggregatum were the most common AM fungal species. The E. adenophorum invasion significantly decreased the AM fungal spore density in the soil. Furthermore, with increasing of E. adenophorum invasion the spore densities of C. etunicatum, G. aggregatum, and G. arenarium significantly decreased, whereas F. geosporus significantly increased. Nonmetric multidimensional scaling demonstrated that the AM fungus community composition was significantly different (P=0.003) in the different invasive levels of E. adenophorum, and significantly correlated with plant species richness, soil total P, and soil NO₃ ^?-N. The results suggest that the alteration in AM fungus community might be caused by E. adenophorum invasion via changing the local plant community and soil properties in a Chinese secondary forest ecosystem.     

Effect of edaphic factors and seasonal variation on spore density and root colonization of arbuscular mycorrhizal fungi in sugarcane fields

Sugarcane fields in 14 different study sites were analyzed for the presence of different arbuscular mycorrhizal fungal (AMF) spores. A total of 23 AMF species representing four genera were identified, among which Glomus fasciculatum and G. mosseae were the dominant species. The mean spore density in the root-zone soils of sugarcane plants varied from 119 to 583 per 100 g of soil, and the mean percentage root colonization varied from 60 to 89 %. A study of the effect of edaphic factors on AM spore density and percentage root colonization revealed a positive correlation between pH and AMF spore density and root colonization and a negative correlation between electrical conductivity, nitrogen, and phosphorus. A positive correlation was observed between AMF spore density and root colonization. Season was also found to play a vital role in determining AMF spore density and percentage root colonization, with high spore density and root colonization observed during the summer season and lower spore densities and root colonization during the winter season.     

Arbuscular mycorrhizal dynamics in a chronosequence of Caragana korshinskii plantations

Arbuscular mycorrhizal (AM) fungi in a chronosequence of 5–42-year-old Caragana korshinskii plantations in the semi-arid Loess Plateau region of northwestern China were investigated. AM fungi colonization, spore diversity and PCR-denatured gradient gel electrophoresis-based AM fungal SSU rRNA gene sequences were analyzed. AM fungi colonization [measured as the percent of root length (%RLC), vesicular (%VC) and arbuscular (%AC) colonization] and spore density were significantly correlated with sampling month, but not with plant age, except for %RLC. The percent of vesicular colonization was negatively correlated with soil total nitrogen and organic carbon, and spore density was negatively correlated with soil moisture and available phosphorus. Ten distinguishable AM fungal spore morphotypes, nine Glomus and one Scutellospora species, were found. Nine AM fungal Glomus phylotypes were identified by sequencing, but at each sampling time only four to six AM fungal phylotypes were detected. The AM fungal community was significantly seasonal, whereas the AM fungal species richness did not increase with plantation age. A significant change in AM fungal colonization and community composition over an annual cycle was observed in this study, and our results suggest that the changes of AM are the product of the interaction between host phenology, soil characteristics and habitat. Understanding these interactions is essential if habitat restoration is to be effective.     

Availability and function of arbuscular mycorrhizal and ectomycorrhizal fungi during revegetation of dewatered reservoirs left after dam removal

Revegetation following dam removal projects may depend on recovery of arbuscular mycorrhizal (AM) and ectomycorrhizal (EM) fungal communities, which perform valuable ecosystem functions. This study assessed the availability and function of AM and EM fungi for plants colonizing dewatered reservoirs following a dam removal project on the Elwha River, Olympic Peninsula, Washington, United States. Availability was assessed via AM fungal spore density in soils and EM root tip colonization of Salix sitchensis (Sitka willow) in an observational field study. The effect of mycorrhizal fungi from 4 sources (reservoir soils, commercial inoculum, and 2 mature plant community soils) on growth and nutrient status of S. sitchensis was quantified in a greenhouse study. AM fungal spores and EM root tips were present in all field samples. In the greenhouse, plants receiving reservoir soil inoculum had only incipient mantle formation, while plants receiving inoculum from mature plant communities had fully formed EM root tips. EM formation corresponded with alleviation of phosphorus stress in plants (lower shoot nitrogen:phosphorus). Thus, revegetating plants have access to AM and EM fungi following dam removal, and EM formation may be especially important for plant P uptake in reservoir soils. However, availability of mycorrhizal fungi declines with distance from established plant communities. Furthermore, EM fungal communities in recently dewatered reservoirs may not be as effective at forming beneficial mycorrhizae as those from mature plant communities. Whole soil inoculum from mature plant communities may be important for the success of revegetating plants and recovery of mycorrhizal fungal communities.     

A preliminary survey of the arbuscular mycorrhizal status of grassland plants in southern Tibet

We report for the first time the arbuscular mycorrhizal (AM) status of native plant species and AM fungal diversity in the grasslands of southern Tibet. A total of 51 soil samples were collected from the rhizospheres of the dominant plant species, and AM fungal structures were observed in 18 (82%) of 22 plant species examined. Vesicles and aseptate hyphae were the structures most frequently observed in the plant roots. After trap culture for 5 months, 25 AM fungal taxa were identified in the soil samples collected, of which nine belonged to Glomus, ten to Acaulospora, one to Entrophospora and five to Scutellospora. The frequency of occurrence of different genera and species varied greatly. Glomus was the dominant genus, and the most frequent and abundant species was Glomus mosseae. Over the whole sampling area, spore density in the rhizosphere soil of different host plant species ranged from 2 to 66 per 20 g air-dried soil. Overall AM fungal species richness was 2.10 and species diversity was 2.35. AM fungal diversity was also compared among the four different land use types (farmland and normal, disturbed and highly disturbed montane scrub grassland). Spore densities in the farmland and normal grassland were much higher than in the grasslands that had been degraded to varying degrees. The species richness in normal grassland was the highest of the four land use types examined. Species diversity varied from 1.99 to 0.94 and was highest in normal grassland, intermediate in degraded grassland and farmland, and lowest in the highly disturbed grassland.     

Arbuscular mycorrhizas in cycads of southern India

Root and soil samples of three potted or ground-grown cycads ( Cycas circinalis, C. revoluta, Zamiasp.) were collected between November 1999 and June 2000 and surveyed for arbuscular mycorrhizal (AM) colonization and spore populations. AM fungi were associated with all root systems and rhizosphere samples examined. Root colonization was of a typical Arum type and AM colonization levels differed significantly between species and between potted and ground-grown cycads. Mycorrhizal colonization levels were inversely related to root hair number and length. Spores of nine morphotypes belonging to three genera ( Acaulospora, Glomus, Scutellospora) were extracted from soil. The percentage root length colonized by AM fungi was not related to soil factors, but total AM fungal spore numbers in the rhizosphere soil were inversely related to soil nitrogen and phosphorus levels. AM fungal spore numbers in the soil were linearly related to root length colonized. The co-occurrence of septate non-mycorrhizal fungi was recorded for the first time in cycads. These observations and the relationship between plant mycorrhizal status and soil nutrients are discussed.     

Increasing flavonoid concentrations in root exudates enhance associations between arbuscular mycorrhizal fungi and an invasive plant

Many invasive plants have enhanced mutualistic arbuscular mycorrhizal (AM) fungal associations, however, mechanisms underlying differences in AM fungal associations between introduced and native populations of invasive plants have not been explored. Here we test the hypothesis that variation in root exudate chemicals in invasive populations affects AM fungal colonization and then impacts plant performance. We examined flavonoids (quercetin and quercitrin) in root exudates of native and introduced populations of the invasive plant Triadica sebifera and tested their effects on AM fungi and plant performance. We found that plants from introduced populations had higher concentrations of quercetin in root exudates, greater AM fungal colonization and higher biomass. Applying root exudates more strongly increased AM fungal colonization of target plants and AM fungal spore germination when exudate donors were from introduced populations. The role of root exudate chemicals was further confirmed by decreased AM fungal colonization when activated charcoal was added into soil. Moreover, addition of quercetin into soil increased AM fungal colonization, indicating quercetin might be a key chemical signal stimulating AM fungal associations. Together these results suggest genetic differences in root exudate flavonoids play an important role in enhancing AM fungal associations and invasive plants’ performance, thus considering root exudate chemicals is critical to unveiling mechanisms governing shifting plant-soil microbe interactions during plant invasions.Subject terms: Population dynamics, Community ecology, Plant ecology     

绢蒿荒漠放牧草地丛枝菌根真菌数量特征的时空变化

以绢蒿荒漠草地的优势种绢蒿、羊茅(针茅)、苔草为研究材料,探究其根际AM真菌和根系侵染率及季节性动态变化对放牧的响应。结果表明:随放牧压力的增大,AM真菌孢子密度、物种丰富度和根系侵染率显著降低(P<0.05),多样性指数在过度放牧的条件下显著(P<0.05)降低;在同一放牧条件下,0~10 cm土壤层的孢子密度、物种丰富度和根系侵染率均显著(P<0.05)高于10~20 cm和20~30 cm土壤层,多样性指数随土壤深度的增大差异性不显著(P>0.05);不同的宿主植物对AM真菌的依赖性不同,即侵染率高低依次为绢蒿>羊茅>苔草;AM真菌孢子密度、物种丰富度和植物根系侵染率随季节的变化表现出一定的变化规律,三者均呈现"双峰"现象,在6月和9月分别达到峰值,与绢蒿荒漠草地植被的生长节律几乎同步;温度对AM真菌孢子密度、丰富度和侵染率的影响不显著,但降雨量对植物根系的侵染率有极显著(P<0.01)的影响,随降雨量的增加侵染率极显著升高。     

Arbuscular mycorrhizal fungal community divergence within a common host plant in two different soils in a subarctic Aeolian sand area

There is rising awareness that different arbuscular mycorrhizal (AM) fungi have different autoecology and occupy different soil niches and that the benefits they provide to the host plant are dependent on plant-AM fungus combination. However, the role and community composition of AM fungi in succession are not well known and the northern latitudes remain poorly investigated ecosystems. We studied AM fungal communities in the roots of the grass Deschampsia flexuosa in two different, closely located, successional stages in a northern Aeolian sand area. The AM fungal taxa richness in planta was estimated by cloning and sequencing small subunit ribosomal RNA genes. AM colonization, shoot δ ¹³C signature, and %N and %C were measured. Soil microbial community structure and AM fungal mycelium abundance were estimated using phospholipid (PLFA) and neutral lipid (NLFA) analyses. The two successional stages were characterized by distinct plant, microbial, and fungal communities. AM fungal species richness was very low in both the early and late successional stages. AM frequency in D. flexuosa roots was higher in the early successional stage than in the late one. The AM fungal taxa retrieved belonged to the genera generally adapted to Arctic or extreme environments. AM fungi seemed to be important in the early stage of the succession, suggesting that AM fungi may help plants to better cope with the harsh environmental conditions, especially in an early successional stage with more extreme environmental fluctuations.     

Arbuscular mycorrhizal status of spring ephemerals in the desert ecosystem of Junggar Basin, China

A survey was made of the arbuscular mycorrhizal (AM) status of 73 spring ephemeral plant species that grow in the desert ecosystem of Junggar Basin, northwest China. The proportion of AM colonization ranged from 7 to 73% with a mean value of 30%. A total of 65 plant species studied were AM with coils/arbuscules or vesicles and the remaining eight species were possibly AM with no coils/arbuscules or vesicles but with fungal mycelia in the root cortex. AM fungal spores were isolated from rhizosphere samples of all 73 plant species and identified. The mean spore density was 22 per 20 ml of air-dried soil, ranging from 0 to 120. Colonization and spore density of perennials were slightly higher than of annuals and varied among different plant families. A total of 603 AM fungal spore (or sporocarp) specimens were isolated belonging to six genera, Acaulospora, Archaeopora, Entrophospora, Glomus, Paraglomus, and Scutellospora; Glomus was the dominant genus. We conclude that spring ephemerals may be highly dependent on AM associations for survival in the very infertile and arid soils of this desert ecosystem.Electronic supplementary material Supplementary material is available for this article at