Paul Gosling

The role of local environment and geographical distance in determining community composition of arbuscular mycorrhizal fungi at the landscape scale

Arbuscular fungi have a major role in directing the functioning of terrestrial ecosystems yet little is known about their biogeographical distribution. The Baas-Becking hypothesis (‘everything is everywhere, but, the environment selects’) was tested by investigating the distribution of arbuscular mycorrhizal fungi (AMF) at the landscape scale and the influence of environmental factors and geographical distance in determining community composition. AMF communities in Trifolium repens and Lolium perenne roots were assessed in 40 geographically dispersed sites in Ireland representing different land uses and soil types. Field sampling and laboratory bioassays were used, with AMF communities characterised using 18S rRNA terminal-restriction fragment length polymorphism. Landscape-scale distribution of AMF was driven by the local environment. AMF community composition was influenced by abiotic variables (pH, rainfall and soil type), but not land use or geographical distance. Trifolium repens and L. perenne supported contrasting communities of AMF, and the communities colonising each plant species were consistent across pasture habitats and over distance. Furthermore, L. perenne AMF communities grouped by soil type within pasture habitats. This is the largest and most comprehensive study that has investigated the landscape-scale distribution of AMF. Our findings support the Baas-Becking hypothesis at the landscape scale and demonstrate the strong influence the local environment has on determining AMF community composition.

Contrasting arbuscular mycorrhizal communities colonizing different host plants show a similar response to a soil phosphorus concentration gradient

High soil phosphorus (P) concentration is frequently shown to reduce root colonization by arbuscular mycorrhizal (AM) fungi, but the influence of P on the diversity of colonizing AM fungi is uncertain. We used terminal restriction fragment length polymorphism (T-RFLP) of 18S rDNA and cloning to assess diversity of AM fungi colonizing maize (Zea mays), soybean (Glycene max) and field violet (Viola arvensis) at three time points in one season along a P gradient of 10–280 mg l−1 in the field. Percentage AM colonization changed between sampling time points but was not reduced by high soil P except in maize. There was no significant difference in AM diversity between sampling time points. Diversity was reduced at concentrations of P > 25 mg l−1, particularly in maize and soybean. Both cloning and T-RFLP indicated differences between AM communities in the different host species. Host species was more important than soil P in determining the AM community, except at the highest P concentration. Our results show that the impact of soil P on the diversity of AM fungi colonizing plants was broadly similar, despite the fact that different plants contained different communities. However, subtle differences in the response of the AM community in each host were evident.

Spatial scaling of arbuscular mycorrhizal fungal diversity is affected by farming practice

Evidence suggests that microbial communities show patterns of spatial scaling which can be driven by geographical distance and environmental heterogeneity. Here we demonstrate that human management can have a major impact on microbial distribution patterns at both the local and landscape scale. Mycorrhizal fungi are vital components of terrestrial ecosystems, forming a mutualistic symbiosis with plant roots which has a major impact on above ground ecology and productivity. We used contrasting agricultural systems to investigate the spatial scaling of the most widespread mycorrhizal fungus group, the arbuscular mycorrhizal fungi (AMF). Using multiple sampling sites with a maximum separation of 250 km we describe for the first time the roles which land management, environmental heterogeneity and geographical distance play in determining spatial patterns of microbial distribution. Analysis of AMF taxa-area relationships at each sampling site revealed that AMF diversity and spatial turnover was greater under organic relative to conventional farm management. At the regional scale (250 km) distance-decay analyses showed that there was significant change in AMF community composition with distance, and that this was greater under organic relative to conventional management. Environmental heterogeneity was found to be the major factor determining turnover of AMF taxa at the landscape scale. Overall we demonstrate that human management can play a key role in determining the turnover of microbial communities at both the local and regional scales.

Diversity of fungi associated with hair roots of ericaceous plants is affected by land use

Culture-independent molecular studies have provided new insights into the diversity of fungi associating with ericaceous plant roots. However, there is little understanding of the distribution of these fungi across landscapes, or the effects of environmental heterogeneity on ericoid mycorrhizal (ERM) fungal diversity and distribution. Terminal-restriction fragment length polymorphism and selective sequence analyses of the internal transcribed spacer regions of rDNA were used to infer fungal diversity of bait Vaccinium macrocarpon grown in soils from nine peatland sites in Ireland, representing three different land uses (bog, rough grazing and forest plantation) and the fungal communities of field-collected Calluna vulgaris for five of these nine sites. A diverse range of potential ERM fungi were found, and the sampling approach significantly affected the diversity of the fungal community. Despite significant site groupings of the fungal communities associated with V. macrocarpon and C. vulgaris, fungal communities were significantly dissimilar between sites with different land uses. Soil nitrogen content significantly explained 52% of the variation in the V. macrocarpon fungal communities. Evidence suggests that environmental heterogeneity has a role in shaping ERM fungal community composition at the landscape scale.

Distribution and diversity of Paraglomus spp. in tilled agricultural soils

Understanding of the ecology of arbuscular mycorrhizal fungi comes primarily from the order Glomerales, and relatively little is known of the ecology of other orders including the Paraglomerales. We investigated the distribution of the Paraglomerales across the English agricultural landscape under different management systems. Soils were collected from 11 tilled agricultural sites. Presence of Paraglomerales was assessed using PCR amplification of 18S/ITS region ribosomal DNA isolated from trap plants, terminal restriction fragment length polymorphism and cloning. Paraglomus spp. were detected in all samples from one location and sporadically in six more, but not at the other locations. Distribution was not related to soil physiochemical characteristics, but the Paraglomaceae were significantly more common in soils under organic management. Cloning of samples from three sites produced sequences closely related to Paraglomus laccatum but only distantly related to Paraglomus brasilianum and Paraglomus occultum. Individual sites had between 10 and 27 separate terminal restriction fragments (T-RFs). The large number of T-RFs reflected a significant sequence diversity in the ITS region. Paraglomerales were, therefore, widely distributed across the agricultural landscape, though with patchy distribution and low diversity. More intensive agricultural management appeared to impact negatively on Paraglomus spp.

Facilitation of Urtica dioica colonisation by Lupinus arboreus on a nutrient-poor mining spoil

Facilitation is an important process during succession. Legumes often play a significant role as facilitators, particularly in primary succession, enriching the soil with nitrogen (N). The leguminous shrub Lupinus arboreus (Sims) can fix significant N on acidic, nutrient-poor soils. An apparent association between L. arboreus and Urtica dioica (L), which requires high concentrations of soil N and phosphorus (P), suggested that L. arboreus might facilitate colonisation by Urtica of nutrient-poor soils by increasing both soil N and P. I measured significantly higher concentrations of extractable soil P and higher values of soil pH beneath L. arboreus canopy, compared with areas between bushes, occupied by herbaceous vegetation. Litter inputs beneath L. arboreus were more than two and a half times higher in terms of mass of material and P and three times higher in terms of N, than in areas between bushes. This high litter input accounted for the higher soil P concentration and higher pH. It did not lead to higher soil organic matter content however, probably because high nutrient concentration in L. arboreus litter leads to rapid decay. A glasshouse trial showed that Urtica grew poorly on soil collected from areas between bushes of L. arboreus without the addition of supplementary N and P fertiliser, indicting co-limitation, by both N and P. Growth of Urtica on soil from beneath L. arboreus was more than four times higher than on soil from between L. arboreus. Amendment of the latter soil with N significantly increased growth of Urtica, but amendment with P did not, indicating that Urtica growth on this soil was not P limited, even when amended with N. Facilitation of colonisation of this site by Urtica therefore can be attributed to increased soil N and P, derived from litter of L. arboreus. However, in the field Urtica was only found beneath dead and senescent L. arboreus, suggesting a period of inhibition caused by shading, before senescence of L. arboreus allows light penetration to the nutrient-rich soil below.

Evidence for functional redundancy in arbuscular mycorrhizal fungi and implications for agroecosystem management

Arbuscular mycorrhizal (AM) fungi provide benefits to host plants and show functional diversity, with evidence of functional trait conservation at the family level. Diverse communities of AM fungi ought therefore to provide increased benefits to the host, with implications for the management of sustainable agroecosystems. However, this is often not evident in the literature, with diversity saturation at low species number. Growth and nutrient uptake were measured in onions in the glasshouse on AM-free phosphorus (P)-poor soil, inoculated with between one and seven species of AM fungi in all possible combinations. Inoculation with AM fungi increased shoot dry weight as well as P and copper concentrations in shoots but reduced the concentration of potassium and sulphur. There was little evidence of increased benefit from high AM fungal diversity, and increasing diversity beyond three species did not result in significantly higher shoot weight or P or Cu concentrations. Species of Glomeraceae had the greatest impact on growth and nutrient uptake, while species of Acaulospora and Racocetra did not have a significant impact. Failure to show a benefit from high AM fungal diversity in this and other studies may be the result of experimental conditions, with the benefits of AM fungal diversity only becoming apparent when the host plant is faced with multiple stress factors. Replicating the complex interactions between AM fungi, the host plant and their environment in the laboratory in order to fully understand these interactions is a major challenge to AM research.

Converting highly productive arable cropland in Europe to grassland : – a poor candidate for carbon sequestration

Sequestration of atmospheric CO2 as organic carbon by agricultural soils (SOC) is promoted as a climate change mitigation option. IPCC provides guidelines for determining carbon stocks and sequestration potential, incentivising policy changes towards management of farmland for carbon sequestration. However, the basis of the assumption that agricultural soils can sequester significant atmospheric CO2 has been questioned. We sought to determine the potential for conversion of arable cropland to grassland to sequester carbon in the short to medium term and potential limiting factors. There were no differences in SOC stocks in the top 30 cm between grassland up to 17 years old and arable cropland at 14 sites across the UK. However, SOC showed different distribution patterns, being concentrated in the top 10 cm under grassland. Soil microbial communities were significantly different between arable and grassland, with higher biomass and lesser dominance by bacteria in grassland soils. A land use conversion experiment showed these changes occurred within one year of land use change. Failure of grassland soils to accumulate SOC was attributed to reduced available soil nitrogen, resulting in low productivity. The implications of these results for carbon sequestration in soils as a climate change mitigation strategy are discussed.