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Assessing Functions of Soil Microbes with Isotopic Measurements 385<br />

atmospheric N2 ( 15 N/ 14 N = 0. 0036765), for sulfur, Canyon Diablo troilite<br />

( 34 S/ 32 S = 0. 0450045), and for hydrogen and oxygen, Vienna Standard<br />

Mean Ocean Water (V-SMOW, D/H = 0. 00015576, 18 O/ 16 O = 0. 00200520)<br />

(Hoefs 1997). Isotopic values for carbon generally range from 0‰ (carbonates)<br />

to −50‰ (methane in some systems), with mostδ 13 Cvaluesinsystems<br />

dominated by plants of the C3 photosynthetic pathway ranging from −20 to<br />

−35‰. In systems dominated by plants of the C4 photosynthetic pathway,<br />

values range from −10 to −20‰.Forδ 15 N, pools in terrestrial ecosystems<br />

are usually between 20 and −10‰. Incomparisonsamongsamples,samples<br />

with more of the heavy isotope are commonly referred to as isotopically<br />

enriched, or heavy, and samples with less of the heavy isotope are referred<br />

to as isotopically depleted, or light. Isotopic fractionation (∆)forreactions<br />

in open systems can be calculated based on the isotopic signature of the<br />

source and product according to (18.2).<br />

∆(δ n Xsource − δ n Xproduct)/(1 + δ n Xproduct) (18.2)<br />

2.1<br />

Fungi<br />

Fungi can be divided into two main life history strategies depending on<br />

their carbon source. Saprotrophic fungi obtain their carbon from the decay<br />

of dead organic matter, whereas mycorrhizal fungi form symbiotic<br />

associations with plants in which plant-supplied sugars are exchanged<br />

for nutrients obtained by fungi from the soil. Mycorrhizal fungi can be<br />

further divided into several types, of which the two most important are<br />

arbuscular mycorrhizal (AM) fungi (belonging to the Glomerales), symbiotic<br />

on most herbaceous plants and some trees, and ectomycorrhizal<br />

fungi, associated with many of the dominant tree families of temperate<br />

and boreal regions (e.g., Pinaceae, Betulaceae, Salicaceae, and Fagaceae),<br />

but also some families distributed in the tropics (e.g., Dipterocarpaceae<br />

and Myrtaceae). Higher fungi in the phyla Basidiomycota and Ascomycota<br />

have been tempting targets for isotopic measurements in recent years because<br />

many produce large, conspicuous fruiting bodies that often can be<br />

identified to species. The production of large fruiting bodies has allowed<br />

researchers to analyze isotopes at the species level in situ, without resorting<br />

to techniques such as DNA- or RNA-specific analyses. Such isotopic analyses<br />

are providing interesting insights into patterns of carbon and nitrogen<br />

cycling by both ectomycorrhizal and saprotrophic fungi in these phyla.

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