Impact of above- and below-ground invertebrates on temporal and spatial stability of grassland of different diversity

1. Recent theoretical studies suggest that the stability of ecosystem processes is not governed by diversity per se, but by multitrophic interactions in complex communities. However, experimental evidence supporting this assumption is scarce.2. We investigated the impact of plant diversity and the presence of above- and below-ground invertebrates on the stability of plant community productivity in space and time, as well as the interrelationship between both stability measures in experimental grassland communities.3. We sampled above-ground plant biomass on subplots with manipulated above- and below-ground invertebrate densities of a grassland biodiversity experiment (Jena Experiment) 1, 4 and 6 years after the establishment of the treatments to investigate temporal stability. Moreover, we harvested spatial replicates at the last sampling date to explore spatial stability.4. The coefficient of variation of spatial and temporal replicates served as a proxy for ecosystem stability. Both spatial and temporal stability increased to a similar extent with plant diversity. Moreover, there was a positive correlation between spatial and temporal stability, and elevated plant density might be a crucial factor governing the stability of diverse plant communities.5. Above-ground insects generally increased temporal stability, whereas impacts of both earthworms and above-ground insects depended on plant species richness and the presence of grasses. These results suggest that inconsistent results of previous studies on the diversity–stability relationship have in part been due to neglecting higher trophic-level interactions governing ecosystem stability.6. Changes in plant species diversity in one trophic level are thus unlikely to mirror changes in multitrophic interrelationships. Our results suggest that both above- and below-ground invertebrates decouple the relationship between spatial and temporal stability of plant community productivity by differently affecting the homogenizing mechanisms of plants in diverse plant communities.7.Synthesis. Species extinctions and accompanying changes in multitrophic interactions are likely to result not only in alterations in the magnitude of ecosystem functions but also in its variability complicating the assessment and prediction of consequences of current biodiversity loss.

A below-ground herbivore shapes root defensive chemistry in natural plant populations

Plants display extensive intraspecific variation in secondary metabolites. However, the selective forces shaping this diversity remain often unknown, especially below ground. Using Taraxacum officinale and its major native insect root herbivore Melolontha melolontha, we tested whether below-ground herbivores drive intraspecific variation in root secondary metabolites. We found that high M. melolontha infestation levels over recent decades are associated with high concentrations of major root latex secondary metabolites across 21 central European T. officinale field populations. By cultivating offspring of these populations, we show that both heritable variation and phenotypic plasticity contribute to the observed differences. Furthermore, we demonstrate that the production of the sesquiterpene lactone taraxinic acid β-d-glucopyranosyl ester (TA-G) is costly in the absence, but beneficial in the presence of M. melolontha, resulting in divergent selection of TA-G. Our results highlight the role of soil-dwelling insects for the evolution of plant defences in nature.

Arbuscular mycorrhizal fungi mediate below-ground plant-herbivore interactions: a phylogenetic study

Ecological interactions are complex networks, but have typically been studied in a pairwise fashion. Examining how third-party species can modify the outcome of pairwise interactions may allow us to better predict their outcomes in realistic systems. For instance, arbuscular mycorrhizal fungi (AMF) can affect plant interactions with other organisms, including below-ground herbivores, but the mechanisms underlying these effects remain unclear. Here, we use a comparative, phylogenetically controlled approach to test the relative importance of mycorrhizal colonization and plant chemical defences (cardenolides) in predicting plant survival and the abundance of a generalist below-ground herbivore across 14 species of milkweeds (Asclepias spp.). Plants were inoculated with a mixture of four generalist AMF species or left uninoculated. After 1month, larvae of Bradysia sp. (Diptera: Sciaridae), a generalist below-ground herbivore, colonized plant roots. We performed phylogenetically controlled analyses to assess the influence of AMF colonization and toxic cardenolides on plant growth, mortality and infestation by fungus gnats. Overall, plants inoculated with AMF exhibited greater survival than did uninoculated plants. Additionally, surviving inoculated plants had lower numbers of larvae in their roots and fewer non-AM fungi than surviving uninoculated plants. In phylogenetic controlled regressions, gnat density in roots was better predicted by the extent of root colonized by AMF than by root cardenolide concentration. Taken as a whole, AMF modify the effect of below-ground herbivores on plants in a species-specific manner, independent of changes in chemical defence. This study adds to the growing body of literature demonstrating that mycorrhizal fungi may improve plant fitness by conferring protection against antagonists, rather than growth benefits. In addition, we advocate using comparative analyses to disentangle the roles of shared history and ecology in shaping trait expression and to better predict the outcomes of complex multitrophic interactions.

Modelling mean above and below ground litter production based on yield tables

Selective papers of the workshop on "Development of models and forest soil surveys for monitoring of soil carbon", Koli, Finland, April 5-9 2006.

Effect of soil waterlogging on below-ground biomass allometric relations in Norway spruce

An increasing importance is assigned to the estimation and verification of carbon stocks in forests. Forestry practice has several long-established and reliable methods for the assessment of aboveground biomass; however we still miss accurate predictors of belowground biomass. A major windthrow event exposing the coarse root systems of Norway spruce trees allowed us to assess the effects of contrasting soil stone and water content on belowground allocation. Increasing stone content decreases root/shoot ratio, while soil waterlogging leads to an increase in this ratio. We constructed allometric relationships for belowground biomass prediction and were able to show that only soil waterlogging significantly impacts model parameters. We showed that diameter at breast height is a reliable predictor of belowground biomass and, once site-specific parameters have been developed, it is possible to accurately estimate belowground biomass in Norway spruce.

The role of soil microbes in the global carbon cycle: tracking the below-ground microbial processing of plant-derived carbon for manipulating carbon dynamics in agricultural systems.

It is well known that atmospheric concentrations of carbon dioxide (CO2) (and other greenhouse gases) have increased markedly as a result of human activity since the industrial revolution. It is perhaps less appreciated that natural and managed soils are an important source and sink for atmospheric CO2 and that, primarily as a result of the activities of soil microorganisms, there is a soil-derived respiratory flux of CO2 to the atmosphere that overshadows by tenfold the annual CO2 flux from fossil fuel emissions. Therefore small changes in the soil carbon cycle could have large impacts on atmospheric CO2 concentrations. Here we discuss the role of soil microbes in the global carbon cycle and review the main methods that have been used to identify the microorganisms responsible for the processing of plant photosynthetic carbon inputs to soil. We discuss whether application of these techniques can provide the information required to underpin the management of agro-ecosystems for carbon sequestration and increased agricultural sustainability. We conclude that, although crucial in enabling the identification of plant-derived carbon-utilising microbes, current technologies lack the high-throughput ability to quantitatively apportion carbon use by phylogentic groups and its use efficiency and destination within the microbial metabolome. It is this information that is required to inform rational manipulation of the plant–soil system to favour organisms or physiologies most important for promoting soil carbon storage in agricultural soil.

Above- and below-ground biomass and carbon dynamics in Brazilian Cerrado wet grasslands

Questions: Grasslands are usually neglected as potential carbon stocks, partially due to the lack of studies on biomass and carbon dynamics in tropical grasslands. What is the importance of Brazilian tropical wet grasslands as carbon sinks? Does fire frequency and season affect biomass and carbon allocation in Brazilian wet grasslands? Location: Wet grasslands, tropical savanna, Jalapão, Tocantins, northern Brazil. Methods: We determined biomass above- and below-ground, estimated carbon stocks in biennially burned plots (B2) and plots excluded from fire for 4 yr (B4). Moreover, we determined biomass in both rainy and dry seasons. Samples were 0.25 m × 0.25 m × 0.2 m (eight samples per treatment, applying a nested design, total of 48 samples). The biomass was classified in above-ground graminoids, forbs and dead matter, and below-ground roots and other below-ground organs. We used ANOVA to compare variables between treatments and seasons. Results: More than 40% of the total biomass and carbon stocks were located below-ground, mostly in roots. A high proportion of dead biomass (B4) was found in the above-ground material, probably due to low decomposition rates and consequent accumulation over the years. Although these grasslands do not experience water stress, we found significant evidence of resource re-allocation from below-ground organs to the above-ground biomass in the rainy season. Conclusions: We found more dead biomass in the rainy season, probably due to low decomposition rates, which can increase fire risk in these grasslands during the following dry season. These tropical wet grasslands stored high amounts of carbon (621 to 716 g C.m-2), mostly in the roots. Thus, policymakers should consider tropical grasslands as potential carbon stocks, since they are one of the most threatened and unprotected ecosystems in Brazil. © 2012 International Association for Vegetation Science.

Conversion of sub-tropical native vegetation to introduced conifer forest: Impacts on below-ground and above-ground carbon pools

Land-use change can have a major influence on soil organic carbon (SOC) and above-ground C pools. We assessed a change from native vegetation to introduced Pinus species plantations on C pools using eight paired sites. At each site we determined the impacts on 0–50 cm below-ground (SOC, charcoal C, organic matter C, particulate organic C, humic organic C, resistant organic C) and above-ground (litter, coarse woody debris, standing trees and woody understorey plants) C pools. In an analysis across the different study sites there was no significant difference (P > 0.05) in SOC or above-ground tree C stocks between paired native vegetation and pine plantations, although significant differences did exist at specific sites. SOC (calculated based on an equivalent soil mass basis) was higher in the pine plantations at two sites, higher in the native vegetation at two sites and did not differ for the other four sites. The site to site variation in SOC across the landscape was far greater than the variation observed with a change from native vegetation to introduced Pinus plantation. Differences between sites were not explained by soil type, although tree basal area was positively correlated with 0–50 cm SOC. In fact, in the native vegetation there was a significant linear relationship between above-ground biomass and SOC that explained 88.8% of the variation in the data. Fine litter C (0–25 mm diameter) tended to be higher in the pine forest than in the adjacent native vegetation and was significantly higher in the pine forest at five of the eight paired sites. Total litter C (0–100 mm diameter) increased significantly with plantation age (R2 = 0.64). Carbon stored in understorey woody plants (2.5–10 cm DBH) was higher in the native vegetation than in the adjacent pine forest. Total site C varied greatly across the study area from 58.8 Mg ha−1 at a native heathland site to 497.8 Mg ha−1 at a native eucalypt forest site. Our findings suggest that the effects of change from native vegetation to introduced Pinus sp. forest are highly site-specific and may be positive, negative, or have no influence on various C pools, depending on local site characteristics (e.g. plantation age and type of native vegetation).

Differences between root traits of early- and late-successional trees influence below-ground competition and seedling establishment.

The competitive influence of the root system of the exotic grass Urochloa brizantha and the widespread forb Leonotis nepetifolia on the emergence, survival and early growth of the seedlings of eight tropical heliophilous herbaceous species, six early-successional woody species and five late-successional woody species from Brazil, grown in 3500-cm3 pots and in greenhouse without light restriction were assessed. The density of fine-root systems produced by the forb and the grass in pots were 6.8 cm cm-3 soil and 48.1 cm cm-3 soil, respectively. Seedlings survival of the heliophilous herbaceous, early- and late-successional woody species were 86%, 70% and 100% in presence of the forb root system and 12%, 14% and 100% in competition with grass root system, respectively. The competitive pressure applied by the grass root system on seedling growth of the heliophilous herbaceous, early- and late-successional woody species were 2.4, 1.9 and 1.4 times greater than the forb root system. Total root length of the heliophilous herbaceous, early- and late-successional woody species grown without competitors were 13, 33 and 5 times greater than in competition with forb, and were 66, 54 and 6 times greater than in competition with grass root system, respectively. The averages of fine-root diameter of plants grown without competitors were 209 microm for the heliophilous herbaceous, 281 microm for early-successional trees and 382 microm for late-successional trees. The root system of the forb did not avoid seedling establishment of most plant species, but the grass root system hampered more the establishment of heliophilous herbaceous and early-successional woody species than the seedling establishment of late-successional woody species. The different density of root systems produced in soil by the forb and the grass, and the distinct root traits (e.g. root diameter and root tissue density) of the early- and late-successional plant species can explain the differences in the establishment of seedlings of plant species belonging to different groups of tropical succession when exposed to below-ground competition.

Wing polymorphism and dispersal of Scaptocoris carvalhoi (Hemiptera : Cydnidae)

Functional wing polymorphism is commonly observed it) insects, and it may confer an important adaptive value to populations that bear this trait, because it allows dispersal and the location to more favorable habitats for their survival and reproduction. According to the oogenesis-flight syndrome theory, such wing polymorphism may imply differences in the locomotion Capacity of individuals, which is a factor induced by adverse environmental conditions during muscle development in immatures. Scaptocoris carvalhoi Becker (Hemiptera: Cydnidae) is an important agriculture pest in Brazil, and it has burrowing habits. The adults swarm in the beginning of the rainy season after a prolonged drought period in the Brazilian cerrado region. In these swarms, part of the population leaves the soil, performing long flights until locations with more abundant vegetation. In this study, we characterized wing polymorphism in S. carvalhoi, this being the first description in a species of Cydnidae. Brachypterous and macropterous males and females were observed, which showed positive and significant correlations between body length and hindwing length. Macropterous individuals demonstrated greater locomotion capacity than brachypterous individuals. In addition, only long-winged adults could fly, showing wing mobility and flight reaction. The increased number of macropterous individuals inside the soil during the swarming season and in the beginning of the rainy period suggests that wing polymorphism in this population occurs in seasonal cycles and that factors related to the scarcity of rains influence the development of immatures and the formation of polymorphic adults.

An investigation into the possible inhibitory effects of white cypress pine (Callitris glaucophylla) litter on the germination and growth of associated ground cover species

White cypress-pine stands typically support sparse densities of shrubs and grasses. The commonly held opinion is that leaching of allelopathic chemical compounds from cypress-pine litter partly facilitates this exclusion. Germination and growth of cypress pine seedlings do not appear to be similarly affected. This study set out to determine whether cypress litter had a differential effect on germination and growth of cypress-pine seedlings and on associated ground-cover species. Glasshouse trials comparing seedling emergence under cypress- and artificial-litter layers were undertaken. Cypress-pine litter did not have an inhibitory effect on the germination or growth of ground-cover species. In most cases, seedling emergence was facilitated by the application of cypress-pine litter due to its ability to increase the water holding capacity of the underlying soil. Cypress litter did not promote growth of its own seedlings over its competitors except on coarse-textured soils where it provided an ameliorative function to water stress due to the soil's reduced water holding capacity. The inhibition of ground-cover species' germination and growth in pure cypress stands was suggested to be the result of high below-ground resource competition due to the pine's expansive root morphology.

Direct and indirect root defences of milkweed (Asclepias syriaca): trophic cascades, trade-offs and novel methods for studying subterranean herbivory

P>1. Entomopathogenic nematodes can function as indirect defence for plants that are attacked by root herbivores. By releasing volatile organic compounds (VOCs), plants signal the presence of host insects and thereby attract nematodes.2. Nonetheless, how roots deploy indirect defences, how indirect defences relate to direct defences, and the ecological consequences of root defence allocation for herbivores and plant biomass are essentially unknown.3. We investigate a natural below-ground tritrophic system, involving common milkweed, a specialist root-boring beetle and entomopathogenic nematodes, and asked whether there is a negative genetic correlation between direct defences (root cardenolides) and indirect defences (emission of volatiles in the roots and nematode attraction), and between constitutive and inducible defences.4. Volatiles of roots were analysed using two distinct sampling methods. First, we collected emissions from living Asclepias syriaca roots by dynamic headspace sampling. This method showed that attacked A. syriaca plants emit five times higher levels of volatiles than control plants. Secondly, we used a solid phase micro-extraction (SPME) method to sample the full pool of volatiles in roots for genetic correlations of volatile biosynthesis.5. Field experiments showed that entomopathogenic nematodes prevent the loss of biomass to root herbivory. Additionally, suppression of root herbivores was mediated directly by cardenolides and indirectly by the attraction of nematodes. Genetic families of plants with high cardenolides benefited less from nematodes compared to low-cardenolide families, suggesting that direct and indirect defences may be redundant. Although constitutive and induced root defences traded off within each strategy (for both direct and indirect defence, cardenolides and VOCs, respectively), we found no trade-off between the two strategies.6. Synthesis. Constitutive expression and inducibility of defences may trade off because of resource limitation or because they are redundant. Direct and indirect defences do not trade off, likely because they may not share a limiting resource and because independently they may promote defence across the patchiness of herbivore attack and nematode presence in the field. Indeed, some redundancy in strategies may be necessary to increase effective defence, but for each strategy, an economy of deployment reduces overall costs.

Identity and combinations of arbuscular mycorrhizal fungal isolates influence plant resistance and insect preference

1. Accumulating evidence indicates that plant resistance against above-ground herbivores can be affected by the presence of arbuscular mycorrhizal fungi (AMF) in association with the host plant. Little is known, however, about how AMF composition can influence herbivore choice to feed on a particular plant. 2. Unravelling the preference-performance hypothesis in a multitrophic context is needed to expand our knowledge of complex multitrophic interactions in natural systems. If given mycorrhizal fungal genotypes increase attractiveness for a herbivore (reduced plant resistance), then the benefits of increased unpalatability provided by the mycorrhizal fungi (increased plant resistance) might be outweighed by the increased herbivore recruitment. 3. This was addressed by designing three experiments to test the effects of different AMF genotypes, inoculated either alone or in combination, to measure intraspecific AMF effects on plant resistance and insect herbivore preference. Using strawberry (Fragaria vesca L.) plants that were colonised by eight different combinations of Rhizophagus irregularis isolates, we measured effects on plant growth, insect growth and survival, as well as feeding preferences of a generalist herbivore caterpillar (Spodoptera littoralis Boisduval). 4. Overall, it was found that: (i) AMF influenced plant resistance in an AMF genotype-specific manner; (ii) some AMF inoculations decreased insect performance; (iii) insects preferentially chose to feed more on leaves originating from non-mycorrhizal plants; but also that (iv) in a whole plant bioassay, insects preferentially chose the biggest plant, regardless of their mycorrhizal status. 5. Therefore, AMF-mediated trade-offs between growth and resistance against herbivores have been shown. Such trade-offs, particularly driven by plant attractiveness to herbivores, buffer the positive effects of the mycorrhizal symbiosis on enhanced plant growth.

Why don`t our stands grow even faster? Control of production and carbon cycling in eucalypt plantations

The growth of Eucalyptus stands varies several fold across sites, under the influence of resource availability, stand age and stand structure. We describe a series of related studies that aim to understand the mechanisms that drive this great range in stand growth rates. In a seven-year study in Hawaii of Eucalyptus saligna at a site that was not water limited, we showed that nutrient availability differences led to a two-fold difference in stand wood production. Increasing nutrient supply in mid-rotation raised productivity to the level attained in continuously fertilised plots. Fertility affected the age-related decline in wood and foliage production; production in the intensive fertility treatments declined more slowly than in the minimal fertility treatments. The decline in stem production was driven largely by a decline in canopy photosynthesis. Over time, the fraction of canopy photosynthesis partitioned to below-ground allocation increased, as did foliar respiration, further reducing wood production. The reason for the decline in photosynthesis was uncertain, but it was not caused by nutrient limitation, a decline in leaf area or in photosynthetic capacity, or by hydraulic limitation. Most of the increase in carbon stored from conversion of the sugarcane plantation to Eucalyptus plantation was in the above-ground woody biomass. Soil carbon showed no net change. This study and other studies on carbon allocation showed that resource availability changes the fraction of annual photosynthesis used below-ground and for wood production. High resources (nutrition or water) decrease the partitioning below-ground and increase partitioning to wood production. Annual foliage and wood respiration and foliage production as a fraction of annual photosynthesis was remarkably constant across a wide range of fertility treatments and forest age. In the Brazil Eucalyptus Productivity Project, stand structure was manipulated by planting clonal Eucalyptus all at once or in three groups at three-monthly intervals, producing a stand where trees did not segregate into dominants and one that had strong dominance. The uneven stand structure reduced production 10-15% throughout the rotation.