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.     

Differences in AM fungal root colonization between populations of perennial Aster species have genetic reasons

We tested the hypothesis whether differences between plant populations in root colonization by arbuscular mycorrhizal (AM) fungi could be caused by genetic differentiation between populations. In addition, we investigated whether the response to AM fungi differs between plants from different populations and if it is affected by the soil in which the plants are cultivated. We used Aster amellus, which occurs in fragmented dry grasslands, as a model species and we studied six different populations from two regions, which varied in soil nutrient concentration.We found significant differences in the degree of mycorrhizal colonization of plant roots between regions in the field. To test if these differences were due to phenotypic plasticity or had a genetic basis, we performed a greenhouse experiment. The results suggested that Aster amellus is an obligate mycotrophic plant species with a high dependency upon mycorrhiza. Plant biomass was affected only by soil, and not by population or the interaction between the population and the soil. Mycorrhizal colonization was significantly affected by all three factors (soil, population, interaction of soil and population). Plants from the population originating from the soil with lower nutrient availability developed more mycorrhiza even when grown in soil with higher nutrient availability. The correspondence between mycorrhizal colonization of plants in the field and in both soils in the pot experiment suggests that the observed differences in root colonization have a genetic basis.     

Functional ecology of ecotypic differentiation in the Californian serpentine sunflower (Helianthus exilis)

* Here, we examined phenotypic differences between locally adapted serpentine and riparian populations of the serpentine sunflower Helianthus exilis from northern California, USA. * Within a common environment, plants from serpentine and riparian sites were grown in regular potting soil or serpentine soil. Physiology, morphology, phenology and fitness-related traits were measured. * Overall, riparian plants grew more rapidly, attained a larger final size, produced larger leaves, and smaller flowering heads. Riparian plants also invested less in root biomass and were more water-use-efficient than the serpentine plants. Serpentine and riparian plants also differed in leaf concentrations of boron, magnesium, sodium and molybdenum. * These ecotypic differences suggest contrasting adaptive strategies to cope with either edaphic stress in serpentine sites or intense above-ground competition at riparian sites. There was a significant population origin x soil type crossing interaction in one fitness trait (average dry weight) that mirrored local adaptation previously documented for these riparian and serpentine ecotypes. However, because all other fitness traits did not exhibit this crossing interaction in our common garden study, it is possible that phenotypic differences underlying local adaptation may be amplified in the field as a result of biotic and abiotic interactions.     

Responses of arbuscular mycorrhizal symbionts to contrasting environments: field evidence along a Tibetan elevation gradient

Plant adaptation to alpine ecosystems is not fully explained by plant physiological and morphological traits. Arbuscular mycorrhizal (AM) associations may be involved in mediating plant performance in response to environmental differences. Little is known, however, as to whether or not a close relationship exists between plant performance and arbuscular mycorrhizal fungus status across environmental gradients. We conducted a field investigation of the performance of six plant species and their associated AM fungi along higher and lower elevation gradients on Mount Segrila in Tibet. In most of our species, we observed higher shoot and inflorescence biomass production and a lower root-to-shoot ratio in the populations at those sites where the species was dominant (intermediate elevation sites) than in populations sampled at the limits of the distribution. The elevation pattern of root colonization differed with plant species on both gradients, and the extraradical hypha development of most species showed a unimodal pattern as did plant growth. The relationship between plant and fungus traits shows that AM fungus development generally matched host plant performance on the lower elevation gradient but not on the higher elevation gradient. This study provides evidence that plant distribution and productivity were significantly related to root and soil colonization by AM fungi, especially under less physically stressful conditions.     

Phylogenetic trait conservatism and the evolution of functional trade-offs in arbuscular mycorrhizal fungi

The diversity of functional and life-history traits of organisms depends on adaptation as well as the legacy of shared ancestry. Although the evolution of traits in macro-organisms is well studied, relatively little is known about character evolution in micro-organisms. Here, we surveyed an ancient and ecologically important group of microbial plant symbionts, the arbuscular mycorrhizal (AM) fungi, and tested hypotheses about the evolution of functional and life-history traits. Variation in the extent of root and soil colonization by AM fungi is constrained to a few nodes basal to the most diverse groups within the phylum, with relatively little variation associated with recent divergences. We found no evidence for a trade-off in biomass allocated to root versus soil colonization in three published glasshouse experiments; rather these traits were positively correlated. Partial support was observed for correlated evolution between fungal colonization strategies and functional benefits of the symbiosis to host plants. The evolution of increased soil colonization was positively correlated with total plant biomass and shoot phosphorus content. Although the effect of AM fungi on infection by root pathogens was phylogenetically conserved, there was no evidence for correlated evolution between the extent of AM fungal root colonization and pathogen infection. Variability in colonization strategies evolved early in the diversification of AM fungi, and we propose that these strategies were influenced by functional interactions with host plants, resulting in an evolutionary stasis resembling trait conservatism.     

Temperature‐mediated local adaptation alters the symbiotic function in arbuscular mycorrhiza

Variation in the symbiotic function of arbuscular mycorrhizal fungi (AM fungi) has been demonstrated among distinct biotic and abiotic interactions. However, there is little knowledge on how local temperature conditions influence the functional divergence of AM symbionts in alpine ecosystems. Here, we conduct a reciprocal inoculation experiment to explore the three‐way interactions among plants, AM fungal inoculum and temperature at sites of contrasting elevation. Evidence of local adaptation of plant growth was found only under low temperature conditions, with no consistent local versus foreign effect found in AM fungal performance. The origin of either the plant or the inoculum relative to the temperature was important in explaining symbiotic function. Specifically, when inoculum and temperature were sympatric but allopatric to the plant, poor adaptation by the plant to the novel environment was clearly found under both temperature conditions. Further analysis found that the symbiotic function was inversely related to fungal diversity under high temperature conditions. These results suggest that local adaptation represents a powerful factor in the establishment of novel combinations of plant, inoculum and temperature, and confirms the importance of taking into account both biotic and abiotic interactions in the prediction of the response of symbionts to global environmental change.     

Mycorrhiza and seedling establishment in a subarctic meadow: Effects of fertilization and defoliation

Abstract. Question: How does changing resource availability induced by fertilization and defoliation affect seedling establishment and mycorrhizal symbiosis in a subarctic meadow? Location: 610 m a.s.l., Kilpisjärvi (69°03’N, 20°50’E), Finland. Methods: A short‐term full‐factorial experiment was established, with fertilization and defoliation of natural established vegetation as treatments. Seeds of two perennial herbs Solidago virgaurea and Gnaphalium norvegicum were sown in natural vegetation and their germination and growth followed. At the final harvest we measured the response in terms of arbuscular mycorrhizal (AM) colonization, biomass and nitrogen concentration of the seedlings and the established vegetation. Results: Germination rate was negatively affected by defoliation in the unfertilized plots. The shoot biomass of S. virgaurea seedlings was reduced by the defoliation and fertilization treatments, but not affected by their interaction. In G. norvegicum, the germination rate and the seedling shoot biomass were negatively correlated with moss biomass in the plots. In the established plants the arbuscular colonization rate was low and defoliation and fertilization treatments either increased or did not affect the colonization by AM fungi. In the seedlings, the colonization rate by AM fungi was high, but it was not affected by treatments. Both seedlings and established plants were colonized by dark septate fungi. Conclusions: Reduction of plant biomass by herbivores can have different effects on seedling growth in areas of high and low soil nutrient availability. The weak response of AM colonization to defoliation and fertilization suggests that AM symbiosis is not affected by altering plant resource availability under the conditions employed in this study.     

Arbuscular mycorrhizal adaptation,spore germination,root colonization,and host plant growth and mineral acquisition at low pH

Arbuscular mycorrhizal (AM) fungi colonize plant roots and often enhance host plant growth and mineral acquisition, particularly for plants grown under low nutrient and mineral stress conditions. Information about AM fungi and mycorrhizal ( +AM) host plant responses at low pH ( < 5) is limited. Acaulospora are widely reported in acid soil, and Gigaspora sp. appear to be more common in acid soils than Glomus sp. Spores of some AM fungi are more tolerant to acid conditions and high Al than others; t Acaulospora sp., Gigaspora sp., and Glomus manihotis are particularly tolerant. Root colonization is generally less in low than in high pH soils. Percentage root colonization is generally not related to dry matter (DM) produced. Maximum enhancement of plant growth in acid soil varies with AM fungal isolate and soil pH, indicating adaptation of AM isolates to edaphic conditions. Acquisition of many mineral nutrients other than P and Zn is enhanced by +AM plants in acid soil, and the minerals whose concentration is enhanced are those commonly deficient in acid soils (Ca, Mg, and K). Some AM fungal isolates are effective in overcoming soil acidity factors, especially Al toxicity, that restrict plant growth at low pH.     

Effect of arbuscular mycorrhizal fungi, organic fertilizer and soil sterilization on maize growth

The effects of inoculation with arbuscular mycorrhizal (AM) fungi, organic fertilizer (F) applications, and soil sterilization on maize growth were evaluated in a pot experiment. The experiment was in a completely randomized factorial design (2 × 4 × 2) with six replicates for each treatment. There were two soil treatments (sterilized soil, SS and unsterilized soil, US), four organic fertilizer treatments (0.0, 0.5, 1.0 and 2.0 g kg^?1 soil), and two AM fungi treatments (inoculation with Glomus mosseae, +AM and uninoculated control, ?AM). Inoculated plants generally had greater AM colonization, plant height, dry weight and phosphorus (P) uptake than uninoculated controls, and these parameters were significantly increased as the organic fertilizer application increased up to 0.5 g kg^?1 but decreased or had no significant effect compared to the uninoculated plants at the highest fertilizer rate (2.0 g kg^?1). Plant growth, P uptake and AM colonization of root system were significantly higher in sterilized soil compared to the unsterilized control. Our results indicated that the inoculation of AM fungi in field soil with optimal organic fertilizer application greatly improved maize growth and nutrient uptake, and the effect was greater under sterilized soil condition.     

Effect of arbuscular mycorrhizal fungi, organic fertilizer and soil sterilization on maize growth ☆

The effects of inoculation with arbuscular mycorrhizal (AM) fungi, organic fertilizer (F) applications, and soil sterilization on maize growth were evaluated in a pot experiment. The experiment was in a completely randomized factorial design (2 × 4 × 2) with six replicates for each treatment. There were two soil treatments (sterilized soil, SS and unsterilized soil, US), four organic fertilizer treatments (0.0, 0.5, 1.0 and 2.0 g kg^-1 soil), and two AM fungi treatments (inoculation with Glomus mosseae, +AM and uninoculated control, －AM). Inoculated plants generally had greater AM colonization, plant height, dry weight and phosphorus (P) uptake than uninoculated controls, and these parameters were significantly increased as the organic fertilizer application increased up to 0.5 g kg^-1 but decreased or had no significant effect compared to the uninoculated plants at the highest fertilizer rate (2.0 g kg^-1). Plant growth, P uptake and AM colonization of root system were significantly higher in sterilized soil compared to the unsterilized control. Our results indicated that the inoculation of AM fungi in field soil with optimal organic fertilizer application greatly improved maize growth and nutrient uptake, and the effect was greater under sterilized soil condition.     

Patterns of diversity and adaptation in Glomeromycota from three prairie grasslands

Arbuscular mycorrhizal (AM) fungi are widespread root symbionts that often improve the fitness of their plant hosts. We tested whether local adaptation in mycorrhizal symbioses would shape the community structure of these root symbionts in a way that maximizes their symbiotic functioning. We grew a native prairie grass (Andropogon gerardii) with all possible combinations of soils and AM fungal inocula from three different prairies that varied in soil characteristics and disturbance history (two native prairie remnants and one recently restored). We identified the AM fungi colonizing A. gerardii roots using PCR amplification and cloning of the small subunit rRNA gene. We observed 13 operational taxonomic units (OTUs) belonging to six genera in three families. Taxonomic richness was higher in the restored than the native prairies with one member of the Gigaspora dominating the roots of plants grown with inocula from native prairies. Inoculum source and the soil environment influenced the composition of AM fungi that colonized plant roots. Correspondingly, host plants and AM fungi responded significantly to the soil–inoculum combinations such that home fungi often had the highest fitness and provided the greatest benefit to A. gerardii. Similar patterns were observed within the soil–inoculum combinations originating from two native prairies, where five sequence types of a single Gigaspora OTU were virtually the only root colonizers. Our results indicate that indigenous assemblages of AM fungi were adapted to the local soil environment and that this process occurred both at a community scale and at the scale of fungal sequence types within a dominant OTU.     

Niche differentiation between diploid and hexaploid <Emphasis Type="Italic">Aster amellus</Emphasis>

The maintenance of separated diploid and polyploid populations within a contact zone is possible due to both prezygotic and postzygotic isolation mechanisms. Niche differentiation between two cytotypes may be an important prezygotic isolating mechanism and can be studied using reciprocal transplant experiments. We investigated niche differentiation between diploid and hexaploid Aster amellus in their contact zone in the Czech Republic. Diploid populations are confined to habitats with low productivity, whereas hexaploid populations occur in habitats with both low and high productivity. Thus, we chose three diploid populations and six hexaploid populations, three in each of the two different habitat types. We analyzed habitat characteristics and carried out reciprocal transplant experiments in the field using both seeds and adult plants. Sites of diploid and hexaploid populations differed significantly in vegetation and soil properties. The mean number of juveniles was higher at sites of home ploidy level than at sites of foreign ploidy level, suggesting niche differentiation between the two cytotypes. On the other hand, transplanted adult plants survived at all sites and juvenile plants were able to establish at some sites of the foreign cytotype. Furthermore, the mean number of juveniles, survival, and flowering percentages were higher at home sites than at foreign sites, indicating local adaptation. We conclude that niche differentiation between the two cytotypes and local adaptation within each cytotype may contribute to the maintenance of diploid and hexaploid populations of A. amellus in their contact zone. Moreover, further factors, such as differences in flowering phenology and exclusion of minority cytotypes, should also be considered.     

Land-use change impact on mycorrhizal symbiosis in female and male plants of wild <Emphasis Type="Italic">Carica papaya</Emphasis> (Caricaceae)

It has been acknowledged that land-use change has negative effects on genetic diversity and sex ratio in dioecious species, but less attention has been paid on the influence that land-use change has on the biotic interactions, especially between dioecious species and arbuscular mycorrhizal (AM) fungi. AM mutualism involves reciprocal transfer of carbohydrates and mineral nutrients between the host plant’s roots and these fungi. Here, we report spatial and temporal variation in AM colonization in dioecious wild Carica papaya plants growing in sites with different land use intensity. We tagged, recorded the basal stem circumference and collected roots of reproductive female and male Carica papaya plants in three wild sites during dry and rainy seasons of western Mexico. We also collected soil samples in each site to conduct soil chemical analyses. The sexes of C. papaya did not show significant differences in the frequency (percentage of root colonized by intraradical fungal structures) and abundance (length of intraradical hyphae) of AM fungi but the higher AM colonization was observed during the dry season, and in the site with the lowest disturbance. There was no relationship between soil chemistry and AM colonization. Overall, our findings suggest that land-use intensity has a negative effect on AM colonization and we discuss the consequences of habitat loss for the reproductive female and male plants, the implications of decreasing AM colonization for wild Carica papaya plants an important species that provides a source of genetic variation for the C. papaya varieties.     

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     

Cu2+污染土壤接种AM真菌对紫云英生长的影响

通过5个土壤Cu2 水平(0,20,50,100,150mgkg-1)的盆栽试验,研究了不同土壤Cu2 水平接种AM真菌对紫云英生长的影响。结果表明:(1)随着土壤Cu2 水平升高,紫云英生物量下降,与未接种相比,接种AM真菌明显提高了紫云英的生物量,接种G.intraradices对紫云英生物量的提高比接种G.mosseae更为明显,两者间呈显著性差异。(2)随着土壤Cu2 水平升高,紫云英根段浸染率下降,菌丝琥珀酸脱氢酶、碱性磷酸酶活性也下降。(3)在相同土壤Cu2 水平接种不同的AM真菌,紫云英根段浸染率有显著差异,接种G.intraradices的紫云英根段浸染率显著高于接种G.mosseae的处理,其菌丝琥珀酸脱氢酶活性及碱性磷酸酶活性也显著高于接种G.mosseae的处理。(4)接种G.intraradices能显著抑制Cu2 从紫云英地下部分向地上部分的运转,降低Cu2 的毒害,接种G.mosseae相对促进了Cu2 的运转。以上结果显示,Cu2 污染土壤中接种G.intraradices对紫云英生长具有促进作用。     

干旱胁迫下AM真菌对油蒿叶片保护系统的影响

基利用盆栽试验在正常水分和干旱胁迫条件下研究了灭菌土接种AM真菌摩西球囊霉(Glomus mosseae)和土著AM真菌对油(蒿Artemisia ordosica)生长及叶片保护系统的影响。结果表明,干旱胁迫显著抑制了土著AM真菌对油蒿的侵染,但对G.mosseae的侵染影响较小。正常水分和干旱胁迫条件下,接种AM真菌显著增加了油蒿生物量和干重以及根系含磷量;提高了叶绿素、可溶性糖、可溶性蛋白含量并降低了脯氨酸和丙二醛含量;显著增强了过氧化氢酶(CAT)和过氧化物酶(POD)活性,增强了油蒿对干旱的防御能力。     

Sex‐specific interaction between arbuscular mycorrhizal and dark septate fungi in the dioecious plant Antennaria dioica (Asteraceae)

Male and female plants of dioecious species often differ in their resource demands and this has been linked to secondary sexual dimorphism, including sex‐specific interactions with other organisms such as herbivores and pollinators. However, little is known about the interaction between dioecious plants and fungal root endophytes. Plants may be simultaneously colonised by arbuscular mycorrhizal (AM) and dark septate (DS) fungi. While it is well established that AM mutualism involves reciprocal transfer of photosynthates and mineral nutrients between roots of host plants and these fungi, the role of DS fungi remains controversial. Here, we report the temporal and spatial variation in AM and DS fungi in female, male and non‐reproductive Antennaria dioica plants in three natural populations in Finland during flowering and after seed production. Females had higher colonisation by AM fungi, but lower colonisation by DS fungi than male and non‐reproductive plants. The higher AM colonisation was observed during flowering, and this difference varied among populations. Our results suggest that females and males of A. dioica interact with AM and DS fungi differently and that this relationship is dependent on soil fertility.     

Grazing tolerance and mycorrhizal colonization: Effects of resource manipulation and plant size in biennial Gentianella campestris

As herbivory usually leads to loss of photosynthesizing biomass, its consequences for plants are often negative. However, in favorable conditions, effects of herbivory on plants may be neutral or even beneficial. According to the compensatory continuum hypothesis plants can tolerate herbivory best in resource-rich conditions. Besides herbivory, also primarily positive biotic interactions like mycorrhizal symbiosis, bear carbon costs. Tritrophic plant–fungus–herbivore interaction further complicates plant's cost-benefit balance, because herbivory of the host plant is expected to cause decline in mycorrhizal colonization under high availability of soil nutrients when benefits of symbiosis decline in relation to costs. To gain insight into above interactions we tested the effects of plant size and resource manipulation (simulated herbivory and fertilization) on both above-ground performance and on root fungal colonization of the biennial Gentianella campestris.Clipping caused allocation shift from height growth to branches in all groups except in large and fertilized plants. For large plants nutrient addition may have come too late, as the number of meristems was most likely determined already before the fertilization. Clipping decreased the amount of DSE (dark septate endophytic) fungi which generally are not considered to be mycorrhizal. The effect of clipping on total fungal colonization and colonization by arbuscular mycorrhizal (AM) fungal coils were found to depend on host size and resource level. Dissimilar mycorrhizal response to simulated herbivory in small vs. large plants could be due to more intensive light competition in case of small plants. Carbon limited small plants may not be able to maintain high mycorrhizal colonization, whereas large clipped plants allocate extra resources to roots and mycorrhizal fungi at the expense of above-ground parts. Our results suggest that herbivory may increase carbon limitation that leads re-growing shoots and fungal symbionts to function as competing sinks for the limited carbon reserves.     

Colonization potential of in vitro-produced arbuscular mycorrhizal fungus spores compared with a root-segment inoculum from open pot culture

 A reliable inoculum, free from other microorganisms, to produce arbuscular mycorhizal (AM) plants is of the greatest importance when studying the interaction between AM plants and soil microorganisms. We investigated the colonization of leeks from monoxenic in vitro-produced Glomus intraradices spores. The isolated spores were produced using a two-compartment in vitro growth system previously described. A spore suspension was used as inoculum and was compared to the inoculum potential of endomycorrhizal root segments of pot-grown leek (Allium porrum L.) plants. The leeks were grown in a controlled environment and two types of sterilized growth media were tested: calcined montmorillonite clay and a soil mix. Root colonization progressed faster in the soil mix than in the clay. However, in this medium, after an initial delay, root colonization from in vitro-produced spores was essentially the same as that observed with the root-segment inoculum, reaching 44% and 58% respectively, after 16 weeks. Leek roots colonized by the monoxenically-produced spores harbored only the studied AMF fungi while the roots colonized from the root segments were substantially contaminated by other fungi. Accepted: 25 December 1998     

Environmental and genetic effects on the formation of ectomycorrhizal and arbuscular mycorrhizal associations in cottonwoods

Although both environment and genetics have been shown to affect the mycorrhizal colonization of host plants, the impacts of these factors on hosts that can be dually colonized by both ectomycorrhizal (EM) and arbuscular mycorrhizal (AM) fungi are less understood. We examined the influence of environment and host crosstype on the EM and AM colonization of cottonwoods (Populus angustifolia and natural hybrids) by comparing levels of colonization of trees growing in common gardens that differed in elevation and soil type. We also conducted a supplemental watering experiment to determine the influence of soil moisture on AM and EM colonization. Three patterns emerged. First, garden location had a significant impact on mycorrhizal colonization, such that EM colonization was 30% higher and AM colonization was 85% lower in the higher elevation garden than the lower elevation garden. Second, crosstype affected total (EM + AM) colonization, but did not affect EM or AM colonization. Similarly, a significant garden × crosstype interaction was found for total colonization, but not for EM or AM colonization. Third, experimental watering resulted in 33% higher EM colonization and 45% lower AM colonization, demonstrating that soil moisture was a major driver of the mycorrhizal differences observed between the gardens. We conclude that environment, particularly soil moisture, has a larger influence on colonization by AM versus EM fungi than host genetics, and suggest that environmental stress may be a major determinant of mycorrhizal colonization in dually colonized host plants.