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34 Positive interactions and insights Throughout their existence on Earth, fungi and plants have been intricately linked, and their interactions span a vast continuum from detrimental to beneficial [1–3] . Although negative interactions have received more attention, due in part to their impacts on crops, forests and horticulturally important plants, positive plant–fungal interactions are not only ubiquitous but also vitally important to sustaining life on this planet. In this chapter, we highlight what is currently known about the beneficial interactions resulting from fungi that form symbiotic [4,5] relationships with plants. PLANT–FUNGAL MUTUALISMS Fungi and bacteria are now widely acknowledged as playing a central role in the health of all organisms, including plants and humans. In plants, beneficial symbiotic fungi live at least partly within the plant and fall broadly within two groups – endophytic fungi (endophyte meaning ‘within plant’) and mycorrhizal fungi (mycorrhiza meaning ‘fungus root’). These symbiotic fungi, found growing within or between cells in plant shoots and/or roots, provide a variety of benefits to their host plants. These range from improved access to nutrients and increased plant growth to enhancing plant defence mechanisms to herbivory and disease. In return, the fungi obtain nutrition (sugars, lipids) from the plant and a host to live in. Both endophytes and mycorrhizal fungi are ancient symbioses between plants and fungi and are functionally and ecologically very diverse. Endophytes are found in roots and/or shoots, where they are often symptomless. Mycorrhizal fungi are found only in roots, where they form specialised structures for nutritional exchange with plants (see Box 1). 1. Endophytes. It appears that all plants harbour endophytic fungi [3] , from the Arctic to the tropics [6] . There are four classes of fungal endophytes, differentiated by host range, transmission and symbiotic functionality [3] . They are found between and within cells in plant shoots and/or roots [6–9] (see Figure 1) and can be beneficial to plant growth, especially in harsh environments [3,10] . For example, endophytic fungi such as Trichoderma (Ascomycota), used as a seed treatment in agriculture, can induce plant resistance to diseases, water deficits, salinity and also heat stress [11,12] . They do this by altering the expression of the genes involved in root growth, nutrient uptake or protection against oxidative damage [13] . 2. Mycorrhizas. Evidence from the fossil record suggests that mycorrhizas are found even in some of the earliest land plants from around 400 million years ago; the extinct spore-bearing plant Aglaophyton major from the Rhynie chert in Scotland is a characteristic example [14] . These plant–fungal mutualisms have arisen many times in the evolutionary history of different groups of plants and fungi [2] . It is estimated that around 90% of living plant species have mycorrhizal fungi associated with their roots. In contrast, less than 2% of fungal species enter into mycorrhizal partnerships [15,16] . Through becoming specialised to co-exist with plants, the fungi involved rely on their plant hosts for their supply of carbon, having lost the ability of their ancestors to decompose dead organic matter [17] . In return for their photosynthetic carbon, the plants receive water and mineral nutrients from the soil via the fungi. Most mycorrhizal fungi are dependent on their hosts for survival, just as many plants are dependent on their fungal partners. Different plants associate with different groups of fungi to form mycorrhizas (see Table 1), which can be broadly described as ectomycorrhizas (where the fungus remains between the plant root cells) or endomycorrhizas (where the fungus penetrates the root cells). Only around 2% of plants form ectomycorrhizas but they are mostly ecologically and economically important trees that dominate temperate, boreal and some tropical ecosystems (e.g. oaks (Quercus spp.), beech (Fagus spp.), pines (Pinus spp.), spruces (Picea spp.), eucalypts (Eucalyptus spp.) and dipterocarps (e.g. Shorea spp.)); it is therefore important to understand the relationships between these plant species and their fungal partners. Endomycorrhizas are subdivided into arbuscular mycorrhizas (forming tree-like and/or coiled structures within root cells), ericoid mycorrhizas (forming coils within root cells in members of the heather family, the Ericaceae), and orchid mycorrhizas (forming coils within root cells of orchids) (see Box 1). Approximately 80% of plant species from across the plant tree of life have interactions with arbuscular mycorrhizal fungi (e.g. most food crops, as well as grasses, herbs and trees like junipers (Juniperus spp.), maple (Acer spp.), apple (Malus spp.) and cherry (Prunus spp.)). In contrast, the other two groups of endomycorrhizas are plant–fungal associations specific to one plant family each (Ericaceae and Orchidaceae). The fungi forming these different relationships are specialised in different roles in the different ecosystems in which they occur, helping the plants they associate with to thrive (see Boxes 2 and 3). THE FOUNDATIONS OF TERRESTRIAL ECOSYSTEMS It is now known that mycorrhizal fungi provide a number of key benefits to terrestrial ecosystems, including: (i) enhanced nutrient uptake, by provisioning plants with mineral nutrients from soil; (ii) soil structure, through the formation of soil aggregates and pores; and (iii) carbon sequestration, acting as sinks for plant carbon and slowing down decomposition by competing with free-living decomposer organisms in soil for organic nitrogen, known as the Gadgil Effect [18–21] . Another benefit of the mycorrhizal association is that the enhanced acquisition of resources by mycorrhizal plants leads to larger external structures (such as flowers), increasing the attractiveness of plants to pollinators [22] (see Figure 1). Ectomycorrhizal fungi include some well-known edible species such as chanterelles (Cantharellus spp.), truffles (Tuber spp.), milkcaps (Lactarius spp.) and boletes (Boletus spp.), and all of them are specialised in the
Positive plant–fungal interactions 35 FIGURE 1: INTERACTIONS ABOVE AND BELOW GROUND PARASITOIDS Parasitoids are attracted by the elevated production of volatile organic compounds in mycorrhizal plants, increasing the rate of parasitism of herbivores. POLLINATORS Plant nutritional changes produced by mycorrhizal fungi can affect pollinators (e.g. increased growth can lead to a larger number of flowers and more pollinator visits). HERBIVORES Herbivores respond to endophyte chemicals and to increased defences caused by mycorrhizal fungi. APHIDS Aphids respond to mycorrhizalinduced changes in the plant vascular system – the vascular bundles are larger, helping aphids to locate the sap-carrying phloem. LEAF ENDOPHYTES ORCHID MYCORRHIZAS ERICOID MYCORRHIZAS Spores ARBUSCULAR MYCORRHIZAS ECTOMYCORRHIZAS ROOT ENDOPHYTES Nutritional changes produced by mycorrhizal fungi TRUFFLES xxxxxxxxxx Chemical changes elicited by endophytic fungi uptake of inorganic and organic nitrogen – often the most limiting nutrient in temperate and boreal forests. Crucially, ectomycorrhizas are known to mediate ecosystem processes such as carbon storage. In fact, ecosystems dominated by ectomycorrhizal plants contain 70% more carbon per unit of nitrogen than ecosystems dominated by arbuscular mycorrhizal plants [19] . They are also key players in nutrient cycling: in exchange for 15% of plant carbon, they provide water and up to 80% of the soil nitrogen needed by the plant [16,23] . Overall, ectomycorrhizal fungi are directly involved in plant nutrition, growth and survival [24–26] . PLANT PROTECTION – FUNGAL ‘BODYGUARDS’ Arbuscular mycorrhizal fungi (see Table 1) and fungal endophytes have another important interaction with plants and that is providing plants with increased resistance to plant pests and diseases. They are therefore often described as fungal ‘bodyguards’ [27] . Arbuscular mycorrhizal fungi are specialised in the uptake of phosphorus from the soil [2] and have a major role in the regulation of plant diversity [16,28,29] . Moreover, they can alter the chemical composition of their plant host, including increasing levels of plant chemical defences against insect herbivores [30] (see Figure 1).
Page 1: State of the World’s Fungi 2018
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Page 80 and 81: 78 Contributors and references Cont
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86 Contributors and references reve
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88 Acknowledgements Acknowledgement
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The staff and trustees of the Royal
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