Arbuscular mycorrhizal(AM) fungi are obligate symbionts that colonize the roots of more than 80% of land plants.Experiments on the relationship between the host plant and AM in soil or in sterile root-organ culture have provided clear evidence that the extraradical mycelia of AM fungi uptake various forms of nitrogen(N) and transport the assimilated N to the roots of the host plant.However,the uptake mechanisms of various forms of N and its translocation and transfer from the fungus to the host are virtually unknown.Therefore,there is a dearth of integrated models describing the movement of N through the AM fungal hyphae.Recent studies examined Ri T-DNA-transformed carrot roots colonized with AM fungi in 15 N tracer experiments.In these experiments,the activities of key enzymes were determined,and expressions of genes related to N assimilation and translocation pathways were quantified.This review summarizes and discusses the results of recent research on the forms of N uptake,transport,degradation,and transfer to the roots of the host plant and the underlying mechanisms,as well as research on the forms of N and carbon used by germinating spores and their effects on amino acid metabolism.Finally,a pathway model summarizing the entire mechanism of N metabolism in AM fungi is outlined.
The effects of carbon (C) and nitrogen (N) sources on N utilization and biosynthesis of amino acids were examined in the germinating spores of the arbuscular mycorrhizal (AM) fungus Glomus intraradices Schenck & Smith after exposure to various N substrates,CO2,glucose,and/or root exudates.The N uptake and de novo biosynthesis of amino acids were analyzed using stable isotopic labeling with mass spectrometric detection.High-performance liquid chromatography-based analysis was used to measure amino acid levels.In the absence of exogenous N sources and in the presence of 25 mL L-1 CO2,the germinating AM fungal spores utilized internal N storage as well as C skeletons derived from the degradation of storage lipids to biosynthesize the free amino acids,in which serine and glycine were produced predominantly.The concentrations of internal amino acids increased gradually as the germination time increased from 0 to 1 or 2 weeks.However,asparagine and glutamine declined to the low levels;both degraded to provide the biosynthesis of other amino acids with C and N donors.The availability of exogenous inorganic N (ammonium and nitrate) and organic N (urea,arginine,and glutamine) to the AM fungal spores using only CO2 for germination generated more than 5 times more internal free amino acids than those in the absence of exogenous N.A supply of exogenous nitrate to the AM fungal spores with only CO2 gave rise to more than 10 times more asparagine than that without exogenous N.In contrast,the extra supply of exogenous glucose to the AM fungal spores generated a significant enhancement in the uptake of exogenous N sources,with more than 3 times more free amino acids being produced than those supplied with only exogenous CO2.Meanwhile,arginine was the most abundant free amino acid produced and it was incorporated into the proteins of AM fungal spores to serve as an N storage compound.
