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We show that fungi that forage for wood do not conform to the paradigm of symmetric radial growth and grow asymmetrically by default. Asymmetry is further accentuated by contact with a resource that also partially polarizes growth in the direction of the resource. Despite marked changes at the perimeter, overall growth allocation on an area basis is, however, unchanged implying sophisticated regulation at the colony level. Using mathematical models, we show that this behaviour is best explained as a local response of the immediate segment contacting the resource. The model reveals that foraging behaviour is adaptive but only for resources that are clustered in space and is selectively neutral for randomly scattered resources. This clustered spatial distribution matches that found in the natural environment. Modelling also shows that the foraging strategy used by these fungi involves substantial risks as well as benefits., (© 2013 Society for Applied Microbiology and John Wiley & Sons Ltd.)

Transport networks are vital components of multi-cellular organisms, distributing nutrients and removing waste products. Animal cardiovascular and respiratory systems, and plant vasculature, are branching trees whose architecture is thought to determine universal scaling laws in these organisms. In contrast, the transport systems of many multi-cellular fungi do not fit into this conceptual framework, as they have evolved to explore a patchy environment in search of new resources, rather than ramify through a three-dimensional organism. These fungi grow as a foraging mycelium, formed by the branching and fusion of threadlike hyphae, that gives rise to a complex network. To function efficiently, the mycelial network must both transport nutrients between spatially separated source and sink regions and also maintain its integrity in the face of continuous attack by mycophagous insects or random damage. Here we review the development of novel imaging approaches and software tools that we have used to characterise nutrient transport and network formation in foraging mycelia over a range of spatial scales. On a millimetre scale, we have used a combination of time-lapse confocal imaging and fluorescence recovery after photobleaching to quantify the rate of diffusive transport through the unique vacuole system in individual hyphae. These data then form the basis of a simulation model to predict the impact of such diffusion-based movement on a scale of several millimetres. On a centimetre scale, we have used novel photon-counting scintillation imaging techniques to visualize radiolabel movement in small microcosms. This approach has revealed novel N-transport phenomena, including rapid, preferential N-resource allocation to C-rich sinks, induction of simultaneous bi-directional transport, abrupt switching between different pre-existing transport routes, and a strong pulsatile component to transport in some species. Analysis of the pulsatile transport component using Fourier techniques shows that as the colony forms, it self-organizes into well demarcated domains that are identifiable by differences in the phase relationship of the pulses. On the centimetre to metre scale, we have begun to use techniques borrowed from graph theory to characterize the development and dynamics of the network, and used these abstracted network models to predict the transport characteristics, resilience, and cost of the network.

Microorganisms display a range of oscillatory phenomena that operate over different temporal scales. Fourier analysis provides a compact description of such oscillations in terms of their frequency, magnitude and phase. However, in the majority of studies there is no explicit consideration of the spatial organisation of the oscillation. Here we describe procedures and a software package to map oscillatory phenomena in microorganisms in both the time and frequency domains. Key parameters of interest, such as frequency, phase or magnitude of the oscillations, are presented as pseudo-colour coded maps. This maintains the spatial information in the image and greatly facilitates understanding of potentially complex propagating waves or development of oscillatory domains with distinct behaviour. We illustrate the utility of this system with reference to spatial analysis of the pulsatile component to amino acid transport in mycelial systems of Phanerochaete velutina and Coniophora puteana, and actin-myosin based contractions in Physarum polycephalum.

Fungi play a central role in the nutrient cycles of boreal and temperate forests. In these biomes, the saprotrophic wood-decay fungi are the only organisms that can completely decompose woody plant litter. In particular, cord-forming basidiomycete fungi form extensive mycelial networks that scavenge scarce mineral nutrients and translocate them over long distances to exploit new food resources. Despite the importance of resource allocation, there is limited information on nutrient dynamics in these networks, particularly for nitrogen, as there is no suitable radioisotope available. We have mapped N-translocation using photon-counting scintillation imaging of the non-metabolised amino acid analogue, (14)C-aminoisobutyrate. We describe a number of novel phenomena, including rapid, preferential N-resource allocation to C-rich sinks, induction of simultaneous bi-directional N-transport, abrupt switching between different pre-existing transport routes, and emergence of locally synchronised, oscillatory phase domains. It is possible that such self-organised oscillatory behaviour is a mechanism to achieve global co-ordination in the mycelium.