Fire shapes fungal guild diversity and composition through direct and indirect pathways.

Fire has shaped global ecosystems for millennia by directly killing organisms and indirectly altering habitats and resources. All terrestrial ecosystems, including fire-prone ecosystems, rely on soil-inhabiting fungi, where they play vital roles in ecological processes. Yet our understanding of how fire regimes influence soil fungi remains limited and our knowledge of these interactions in semiarid landscapes is virtually absent. We collected soil samples and vegetation measurements from sites across a gradient in time-since-fire ages (0-75 years-since-fire) and fire frequency (burnt 0-5 times during the recent 29-year period) in a semiarid heathland of south-eastern Australia. We characterized fungal communities using ITS amplicon-sequencing and assigned fungi taxonomically to trophic guilds. We used structural equation models to examine direct, indirect and total effects of time-since-fire and fire frequency on total fungal, ectomycorrhizal, saprotrophic and pathogenic richness. We used multivariate analyses to investigate how total fungal, ectomycorrhizal, saprotrophic and pathogenic species composition differed between post-fire successional stages and fire frequency classes. Time-since-fire was an important driver of saprotrophic richness; directly, saprotrophic richness increased with time-since-fire, and indirectly, saprotrophic richness declined with time-since-fire (resulting in a positive total effect), mediated through the impact of fire on substrates. Frequently burnt sites had lower numbers of saprotrophic and pathogenic species. Post-fire successional stages and fire frequency classes were characterized by distinct fungal communities, with large differences in ectomycorrhizal species composition. Understanding the complex responses of fungal communities to fire can be improved by exploring how the effects of fire flow through ecosystems. Diverse fire histories may be important for maintaining the functional diversity of fungi in semiarid regions.

Impacts of elevated atmospheric CO2 on arbuscular mycorrhizal fungi and their role in moderating plant allometric partitioning.

Elevated atmospheric CO2 concentration (eCO2) effects on plants depend on several factors including plant photosynthetic physiology (e.g. C3, C4), soil nutrient availability and plants' co-evolved soil-dwelling fungal symbionts, namely arbuscular mycorrhizal (AM) fungi. Complicated interactions among these components will determine the outcomes for plants. Therefore, clearer understanding is needed of how plant growth and nutrient uptake, along with root-colonising AM fungal communities, are simultaneously impacted by eCO2. We conducted a factorial growth chamber experiment with a C3 and a C4 grass species (± AM fungi and ± eCO2). We found that eCO2 increased plant biomass allocation towards the roots, but only in plants without AM fungi, potentially associated with an eCO2-driven increase in plant nutrient requirements. Furthermore, our data suggest a difference in the identities of root-colonising fungal taxa between ambient CO2 and eCO2 treatments, particularly in the C4 grass species, although this was not statistically significant. As AM fungi are ubiquitous partners of grasses, their response to increasing atmospheric CO2 is likely to have important consequences for how grassland ecosystems respond to global change.

Targeted plant defense: silicon conserves hormonal defense signaling impacting chewing but not fluid-feeding herbivores.

Plants deploy an arsenal of chemical and physical defenses against arthropod herbivores, but it may be most cost efficient to produce these only when attacked. Herbivory activates complex signaling pathways involving several phytohormones, including jasmonic acid (JA), which regulate production of defensive compounds. The Poaceae also have the capacity to take up large amounts of silicon (Si), which accumulates in plant tissues. Si accumulation has antiherbivore properties, but it is poorly understood how Si defenses relate to defense hormone signaling. Here we show that Si enrichment causes the model grass Brachypodium distachyon to show lower levels of JA induction when attacked by chewing herbivores. Triggering this hormone even at lower concentrations, however, prompts Si uptake and physical defenses (e.g., leaf hairs), which negatively impact chewing herbivores. Removal of leaf hairs restored performance. Crucially, activation of such Si-based defense is herbivore-specific and occurred only in response to chewing and not fluid-feeding (aphid) herbivores. This aligned with our meta-analysis of 88 studies that showed Si defenses were more effective against chewing herbivores than fluid feeders. Our results suggest integration between herbivore defenses in a model Si-accumulating plant, which potentially allows it to avoid unnecessary activation of other costly defenses.

Benefits from Below: Silicon Supplementation Maintains Legume Productivity under Predicted Climate Change Scenarios.

Many studies demonstrate that elevated atmospheric carbon dioxide concentrations (eCO2) can promote root nodulation and biological nitrogen fixation (BNF) in legumes such as lucerne (Medicago sativa). But when elevated temperature (eT) conditions are applied in tandem with eCO2, a more realistic scenario for future climate change, the positive effects of eCO2 on nodulation and BNF in M. sativa are often much reduced. Silicon (Si) supplementation of M. sativa has also been reported to promote root nodulation and BNF, so could potentially restore the positive effects of eCO2 under eT. Increased nitrogen availability, however, could also increase host suitability for aphid pests, potentially negating any benefit. We applied eCO2 (+240 ppm) and eT (+4°C), separately and in combination, to M. sativa growing in Si supplemented (Si+) and un-supplemented soil (Si-) to determine whether Si moderated the effects of eCO2 and eT. Plants were either inoculated with the aphid Acyrthosiphon pisum or insect-free. In Si- soils, eCO2 stimulated plant growth by 67% and nodulation by 42%, respectively, whereas eT reduced these parameters by 26 and 48%, respectively. Aphids broadly mirrored these effects on Si- plants, increasing colonization rates under eCO2 and performing much worse (reduced abundance and colonization) under eT when compared to ambient conditions, confirming our hypothesized link between root nodulation, plant growth, and pest performance. Examined across all CO2 and temperature regimes, Si supplementation promoted plant growth (+93%), and root nodulation (+50%). A. pisum abundance declined sharply under eT conditions and was largely unaffected by Si supplementation. In conclusion, supplementing M. sativa with Si had consistent positive effects on plant growth and nodulation under different CO2 and temperature scenarios. These findings offer potential for using Si supplementation to maintain legume productivity under predicted climate change scenarios without making legumes more susceptible to insect pests.

The role of silicon in plant biology: a paradigm shift in research approach.

Background: Silicon (Si) is known to have numerous beneficial effects on plants, alleviating diverse forms of abiotic and biotic stress. Research on this topic has accelerated in recent years and revealed multiple effects of Si in a range of plant species. Available information regarding the impact of Si on plant defence, growth and development is fragmented, discipline-specific, and usually focused on downstream, distal phenomena rather than underlying effects. Accordingly, there is a growing need for studies that address fundamental metabolic and regulatory processes, thereby allowing greater unification and focus of current research across disciplines. Scope and Conclusions: Silicon is often regarded as a plant nutritional 'non-entity'. A suite of factors associated with Si have been recently identified, relating to plant chemistry, physiology, gene regulation and interactions with other organisms. Research to date has typically focused on the impact of Si application upon plant stress responses. However, the fundamental, underlying mechanisms that account for the manifold effects of Si in plant biology remain undefined. Here, the known effects of Si in higher plants relating to alleviation of both abiotic and biotic stress are briefly reviewed and the potential importance of Si in plant primary metabolism is discussed, highlighting the need for a unifying research framework targeting common underlying mechanisms. The traditional approach of discipline-specific work on single stressors in individual plant species is currently inadequate. Thus, a holistic and comparative approach is proposed to assess the mode of action of Si between plant trait types (e.g. C3, C4 and CAM; Si accumulators and non-accumulators) and between biotic and abiotic stressors (pathogens, herbivores, drought, salt), considering potential pathways (i.e. primary metabolic processes) highlighted by recent empirical evidence. Utilizing genomic, transcriptomic, proteomic and metabolomic approaches in such comparative studies will pave the way for unification of the field and a deeper understanding of the role of Si in plants.

Trade-Offs between Silicon and Phenolic Defenses may Explain Enhanced Performance of Root Herbivores on Phenolic-Rich Plants.

Phenolic compounds play a role in plant defense against herbivores. For some herbivorous insects, particularly root herbivores, host plants with high phenolic concentrations promote insect performance and tissue consumption. This positive relationship between some insects and phenolics, however, could reflect a negative correlation with other plant defenses acting against insects. Silicon is an important element for plant growth and defense, particularly in grasses, as many grass species take up large amounts of silicon. Negative impact of a high silicon diet on insect herbivore performance has been reported aboveground, but is unreported for belowground herbivores. It has been hypothesized that some silicon accumulating plants exhibit a trade-off between carbon-based defense compounds, such as phenolics, and silicon-based defenses. Here, we investigated the impact of silicon concentrations and total phenolic concentrations in sugarcane roots on the performance of the root-feeding greyback canegrub (Dermolepida albohirtum). Canegrub performance was positively correlated with root phenolics, but negatively correlated with root silicon. We found a negative relationship in the roots between total phenolics and silicon concentrations. This suggests the positive impact of phenolic compounds on some insects may be the effect of lower concentrations of silicon compounds in plant tissue. This is the first demonstration of plant silicon negatively affecting a belowground herbivore.

Belowground Ecology of Scarabs Feeding on Grass Roots: Current Knowledge and Future Directions for Management in Australasia.

Many scarab beetles spend the majority of their lives belowground as larvae, feeding on grass roots. Many of these larvae are significant pests, causing damage to crops and grasslands. Damage by larvae of the greyback cane beetle (Dermolepida albohirtum), for example, can cause financial losses of up to AU$40 million annually to the Australian sugarcane industry. We review the ecology of some scarab larvae in Australasia, focusing on three subfamilies; Dynastinae, Rutelinae, and Melolonthinae, containing key pest species. Although considerable research on the control of some scarab pests has been carried out in Australasia, for some species, the basic biology and ecology remains largely unexplored. We synthesize what is known about these scarab larvae and outline key knowledge gaps to highlight future research directions with a view to improve pest management. We do this by presenting an overview of the scarab larval host plants and feeding behavior; the impacts of abiotic (temperature, moisture, and fertilization) and biotic (pathogens, natural enemies, and microbial symbionts) factors on scarab larvae and conclude with how abiotic and biotic factors can be applied in agriculture for improved pest management, suggesting future research directions. Several host plant microbial symbionts, such as arbuscular mycorrhizal fungi and endophytes, can improve plant tolerance to scarabs and reduce larval performance, which have shown promise for use in pest management. In addition to this, several microbial scarab pathogens have been isolated for commercial use in pest management with particularly promising results. The entomopathogenic fungus Metarhizium anisopliae caused a 50% reduction in cane beetle larvae while natural enemies such as entomopathogenic nematodes have also shown potential as a biocontrol. Key abiotic factors, such as soil water, play an important role in affecting both scarab larvae and these control agents and should therefore feature in future multi-factorial experiments. Continued research should focus on filling knowledge gaps including host plant preferences, attractive trap crops, and naturally occurring pathogens that are locally adapted, to achieve high efficacy in the field.