Most plants entertain mutualistic interactions known as arbuscular mycorrhiza (AM) with soil fungi (Glomeromycota) which provide them with mineral nutrients in exchange for reduced carbon from the plant. Mycorrhizal roots represent strong carbon sinks in which hexoses are transferred from the plant host to the fungus. However, most of the carbon in AM fungi is stored in the form of lipids. The absence of the type I fatty acid synthase (FAS-I) complex from the AM fungal model species Rhizophagus irregularis suggests that lipids may also have a role in nutrition of the fungal partner. This hypothesis is supported by the concerted induction of host genes involved in lipid metabolism. We explore the possible roles of lipids in the light of recent literature on AM symbiosis.
Carbohydrates in AM Fungal NutritionAM fungi are strictly biotrophic, in other words they depend on their host to complete their life cycle. Although the basis of biotrophy is poorly understood, it has been suggested that it is due to the dependency of AM fungi on some essential nutritional factor(s) from the host [1]. Direct uptake measurements revealed that intraradical mycelium (IRM) can take up carbohydrates, whereas extraradical mycelium (ERM) did not acquire significant amounts of carbohydrates [2,3]. Perhaps this is because AM fungal genes involved in nutrient uptake are expressed only in the host, which would explain their biotrophy. Tracer studies on mycorrhizal roots, and respirometric analysis on isolated intraradical hyphae have established glucose as a central substrate for AM fungi [3][4][5], and indeed a monosaccharide transporter has been identified in the fungal partner [6]. Mycorrhizal roots represent strong carbon sinks [2,4,7], suggesting that they attract sucrose from photosynthetic source tissues. Sucrose is thought to be cleaved in the vicinity of the fungus by invertases and sucrose synthase [8], resulting in monosaccharides (glucose and fructose) that are released to the fungus and taken up by its IRM [5,6]. However, in the fungal storage compartments -the spores and the vesicles -carbon is stored primarily in the form of lipids, with minor amounts of the glucose polymer glycogen. We discuss here salient issues relating to carbohydrate and lipid metabolism in AM fungi and their significance for fungal nutrition during symbiosis.
Lipid Metabolism in AM Fungi -Open Questions and SurprisesAlthough the aforementioned carbon fluxes in AM fungi have been firmly established, several questions remain open. AM fungi store and transport most of their carbon in the form of lipids [9][10][11][12]. However, AM fungi cannot produce the basic fatty acid (FA) palmitate (C16) in the absence of their host (as shown in two distantly related species, Rhizophagus irregularis and Gigaspora rosea), while FA elongation and desaturation can occur independently of the plant [13]. This and similar results suggested that AM fungi can only synthesize C16 in the IRM [3,13]. Taken together, the available evidence suggests that AM fungi take up sugars (...