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Symbiotic Fungi: Principles and Practice (Soil Biology)

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290 X.H. He et al.<br />

17.3.1.1 Comments<br />

The 15 N enrichment or labeling method has been widely used to investigate N<br />

transfer between plants (Stern 1993; Chalk 1998; He et al. 2003). However, the<br />

addition of 15 N to quantify N-transfer has limitations, especially in fertile soils.<br />

Compared to the large volume of soil N, which is available to plants along with the<br />

additional 15 N-labeled N, the amount of N transferred from the N-donor to the<br />

N-receiver may be much less than the amount of N taken up from the soil. Also, root<br />

depth <strong>and</strong> pattern of N uptake of the N-receiver may be dissimilar in mixed species<br />

<strong>and</strong> single species situations, which may cause the ratio of unlabeled-to-labeled N<br />

to be different in such growth systems. Either can invalidate the assumptions for<br />

estimating N transfer.<br />

Furthermore, the above experimental design <strong>and</strong> calculations to study two-way<br />

or net N transfer are based on separate, reciprocal, experiments — one where 15 Nis<br />

applied to plant 1, another where 15 N is applied to plant 2. A more accurate<br />

detection of net N transfer would be to use dual isotopes so that concurrent N<br />

transfer could be investigated in a single experiment. Obviously, 15 N is not suitable,<br />

since no technique is available to distinguish the side from which the detected 15 N<br />

comes. One possible method is to supply the stable isotope of 15 N to one plant <strong>and</strong><br />

the radioactive isotope of 13 N to another, but the short half-life time (10 min) of 13 N<br />

may not be suitable for tracing N translocation in long-term plant <strong>and</strong>/or fungal<br />

biological processes (Knowles <strong>and</strong> Blackburn 1993).<br />

17.4 Conclusions<br />

Both 15 N enrichment <strong>and</strong> 15 N natural abundance have been employed to study N<br />

transfer between plants linked by mychorrhizal networks. By tracing translocation<br />

of external high-enriched 15 N from one plant to another over short-term periods<br />

(days to weeks), the above experimental design allows a two-way or net N transfer<br />

through CMNs to be investigated by the 15 N enrichment method. Considering that<br />

N movement is crucial in most terrestrial ecosystems (Vitousek et al. 1997; Moffat<br />

1998; Sanchez 2002; Graham <strong>and</strong> Vance 2003; Nosengo 2003), <strong>and</strong> that the<br />

potential benefits of N transfer are great, particularly in N-limited environments<br />

(Chalk 1998; Sanchez 2002; Sprent, 2005; Forrester et al. 2006; Nara 2006), more<br />

research is warranted on two-way N transfers mediated by mychorrhizal networks<br />

with different species <strong>and</strong> under field conditions.<br />

References<br />

Chalk PM (1998) Dynamics of biologically fixed N in legume–cereal rotations: a review. Aust J<br />

Agric Res 49:303–316<br />

Faber BA, Zasoski RJ, Munns DN (1991) A method for measuring hyphal nutrient <strong>and</strong> water<br />

uptake in mychorrhizal plants. Can J Bot 69:87–94

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