The Life Seam · a live disagreement, handled honestly

The Fraction That Reaches the Tree

A tree fixes carbon in its leaves. Weeks later, a labelled atom of it turns up beside a neighbour of another species. That much is real, measured, and repeatable. The famous wood-wide web argument is not about whether carbon moves — it does. It is a fight over one number nobody has measured: of the carbon a scientist detects at the neighbour, how much is inside the neighbour plant, and how much is sitting in fungal tissue running its own economy? Partition the real fluxes yourself. The mixing arithmetic is exact. The fractions are the open question — and this page picks no winner.

In 1997 Suzanne Simard and colleagues sealed paper-birch and Douglas-fir seedlings in the ground in British Columbia, fed one species ¹³C and the other ¹⁴C as carbon dioxide, waited, and harvested. Carbon labelled in the birch showed up in the fir, and vice versa. The transfer was bidirectional, and it netted out toward the fir: the fir's net gain averaged 6% of the carbon isotope it took up through its own photosynthesis, rising when the fir was shaded. A non-mycorrhizal western-red-cedar control barely picked up any label — evidence the carbon moved mainly through the fungal threads, not loose through the soil.1

Read this first: this is not a debunk. Underground carbon transfer between plants is established. Simard's result stands; it has been reproduced and extended. What is genuinely unsettled — and what this page is about — is the magnitude, the ecological significance, how common mapped networks actually are, and the popular mother-tree narrative built on top. Establishing that a thing happens is not the same as establishing that it matters, or how much.

Two numbers set the stakes. Simard's 6% net, fir-ward is a modest, careful field figure — and note the wording exactly: 6% of the isotope the fir itself fixed, not 6% of the fir's whole carbon budget, and a net figure hiding a larger two-way gross flow. Nineteen years later, Tamir Klein and colleagues labelled the canopy of a 40-metre spruce for five years with heavy-carbon-depleted CO₂ and estimated that tree-to-tree transfer amounted to ~40% of fine-root carbon — about 280 kg of carbon per hectare per year. A huge number. But whether that carbon truly crossed through the fungal network, rather than through overlapping roots and soil, is itself contested.2

Simard et al. · Nature 1997

~6%

net gain, birch → fir

Of the fir's own isotope uptake. Bidirectional; net fir-ward; non-EM cedar control barely labelled.

transfer: established

Klein et al. · Science 2016

~40%

of fine-root carbon

≈280 kg C ha⁻¹ yr⁻¹, 40 m spruce, 5-yr canopy label. Large — but the via-network reading is disputed.

magnitude: contested

Karst et al. · Nat. Ecol. Evol. 2023

0

field studies for mother-tree transfer

Zero peer-reviewed field studies support preferential resource/defence transfer to kin offspring. Seedling-benefit studies split.

that claim: unsupported

Follow a hundred labelled atoms into the ground

Here is the honest core. A field isotope experiment does not watch carbon travel. It labels a donor, waits, harvests tissue near a receiver, and measures the isotope excess in that sample. To read that excess as "the neighbour tree was fed," you have to assume where the labelled carbon ended up. So make the assumption explicit. Below, the donor pushes 100 units of labelled carbon below ground. You decide how it splits between four destinations. The bar always sums to 100 — that part is bookkeeping, and it is exact.

The partitioner · where the label goes100 units fixed
◀ donor treefungal networkneighbour tree ▶

Partition of 100 labelled carbon units across four compartments; live values in the key below.

stayed in donor 40 in fungal tissue 35 to neighbour via soil 5 into neighbour plant 20

Roots, respiration, retained biomass — carbon that never left.

The mycorrhizal fungus's own hyphae & mantle. Its economy, not the plant's.

Exudates, mass flow, direct root contact — reaches the plant, but not through the network.

Carbon crossing the network into the neighbour's own tissue. The claim's real payload.

When you harvest the neighbour's fine roots, this fraction of the fungal tissue comes up attached — inseparable from plant tissue in the sample. This is the number the whole debate hides.

What a harvest of the neighbour detects

42.5 label units

Measured ¹³C in that sample

1.2398 atom%

Two-source isotope mixing — the exact method. The sample is 1000 units of ordinary carbon at natural abundance (1.08 atom% ¹³C, a C3 leaf's baseline) plus 42.5 units of labelled carbon at 5.00 atom%:

ameas = (1000·1.08 + 42.5·5.00) / (1000 + 42.5) = 1.23981 atom%

Invert the same equation — the only thing a mass spectrometer can give you back:

M = 1000·(1.23981 − 1.08) / (5.00 − 1.23981) = 42.5000 units

✓ forward → invert returns the input exactly (identity holds to 1e-9)

But look what the inversion returns: a single number, 42.5 units — the label that came up in the sample. It is the sum of carbon in the neighbour plant (25) and fungal tissue co-harvested with it (17.5). The isotope method gives you the sum. It cannot, by construction, give you the split.

That last sentence is the entire controversy in miniature. In 1999 David Robinson and Alastair Fitter made the point sharply: much of the label recovered "in" a receiver may sit in fungal tissue, not the plant — the mycorrhizal fungus is a large living thing running its own carbon economy, and a plant-to-plant reading quietly credits the plant with the fungus's biomass. Their argument reframed the naive question. Not did carbon move? — yes. But what fraction reached the neighbour plant, versus stayed in the fungus? That is a measurable quantity. It has not been measured in the field.3


One measurement. Two stories.

So freeze the measurement. Whatever the partition above, the isotope reading collapses to one number: the label detected at the neighbour. Call it 100% of the signal. Now slide the one dial nobody has pinned — the share of that signal that is really fungal tissue, not plant — and watch the same, unchanged datum tell opposite stories.

The crux · the unmeasured dial the argument turns on

Of the detected signal, a live percentage fed the neighbour plant and the remainder stayed in fungal tissue.

◀ wood-wide webmycocentric ▶

No field study has measured this. Every value below is compatible with the same isotope reading.

Read as: the neighbour tree was fed

59%

of the detected label is in the plant

Read as: the fungus kept it

41%

is fungal biomass co-harvested with the roots

Same datum. At this dial, the signal supports a genuine but modest plant subsidy — neither the lavish web of the popular telling nor a pure fungal sink.

The dial is not a rhetorical trick. It is a real, physical quantity — the fraction of detected isotope that resides in fungal rather than plant tissue — and it is genuinely unknown for field networks. You cannot dissect a metre of intermingled hyphae and fine root at the resolution the question demands, and bulk digestion and mass spectrometry only ever hand back the sum. Push the dial left and Simard's 6% becomes a real subsidy between trees. Push it right and Robinson & Fitter are correct that the plant barely sees it. The published data are consistent with both. That is not a failure of anyone's experiment; it is where the evidence actually stops.


One sentence, three different amounts of evidence

The popular claim — "trees talk and share food through an underground fungal web, and mother trees nurture their young" — is really three claims welded together. Karst, Jones and Hoeksema separated them in 2023 and asked what the peer-reviewed literature actually supports. The three answers are not the same.4

1 · Mycorrhizal networks link trees, and carbon can move between themestablished

Directly demonstrated — Simard 1997 and many since. The mechanism is real. (Even here, Karst notes networks are rarely actually mapped in the field; linkage is usually inferred, not traced.)

2 · These networks commonly benefit the receiving seedlingmixed / split

Across the field/greenhouse studies Karst reviewed, seedling outcomes split roughly evenly between improved, no effect, and harmed — with no effect the most common. "Benefit" is not the default finding.

3 · Mother trees preferentially send resources / defence signals to their kin offspringno field evidence

Karst et al. found zero peer-reviewed field studies supporting preferential transfer to related seedlings, or network-borne defence signalling, in forests. The most cited part of the story is the least supported.

They also documented a positive citation bias: papers were increasingly cited as if they had shown benefit or communication even when their own results were neutral or negative — the story outrunning the data. (A minor 2023 author correction to their paper removed four stray references and renumbered; it changed none of this.4)


The argument, with no winner picked

This is a live scientific disagreement between serious researchers, and it would be dishonest to resolve it for you. Here are the positions in their own register.

Simard et al. · 2024

Transfer is real, bidirectional, and shaped by source–sink relations; the networks and their ecological role are worth taking seriously. The critics, they argue, set an unreasonably high bar, understate the positive evidence, and caricature the framing.

Response to questions about CMNs, Front. For. Glob. Change (2024)

Klein et al. · 2023

"Facts, not fantasy." Belowground carbon transfer among trees is a documented fact; what remains genuinely elusive is its magnitude and ecological significance. Defend the phenomenon; stay disciplined about how much it matters.

Facts, not fantasy, Open Research Europe (2023)

Robinson & Fitter · 1999

The mycocentric reframe: net plant-to-plant transfer is likely small, and much detected label sits in fungal tissue. This is an interpretation, not a field measurement — which is exactly why the partition above is honest rather than settled.

Magnitude & control of C transfer, J. Exp. Bot. (1999)

Karst et al. · 2023

Don't confuse "happens" with "matters" with "nurtures kin." The three are supported by strong, mixed, and absent evidence respectively — and citation practice has blurred the difference into a single confident story.

Positive citation bias…, Nat. Ecol. Evol. (2023)

One boundary worth keeping straight. Everything above is about ectomycorrhizal (EM) symbioses — the fungi of conifers, birches, oaks — which is what Simard and Klein studied. The other great guild, arbuscular mycorrhizal (AM) fungi of most herbs and grasses, is a different system with different carbon rules; some work (e.g. Pfeffer et al. 2004) argues the AM fungus does not pass its carbon on to plant tissue at all. Don't let a headline blur the two.

The check — what is exact, and what is the open question

Exact, and recomputed live above. Two-source isotope mixing is linear in atom fraction: a sample of 1000 background units at 1.08 atom% ¹³C plus M labelled units at 5.00 atom% reads a = (1000·1.08 + M·5.00)/(1000+M), and that equation inverts to return M exactly. The four-way partition always sums to 100. The detected signal is M = P + S + φ·F. All of this is arithmetic; the verifier reproduces every displayed value to nine decimals.

Open — flagged, not hidden. The partition fractions (D, F, S, P) and the fungal co-harvest fraction (φ) are not known for field networks. No published field study measures how the detected isotope splits between neighbour-plant tissue and co-harvested fungal tissue. The slick readout does not make those numbers real — it makes visible that the same measurement is compatible with a wide range of them. Simard's 6% is a genuine net figure; Klein's 40% is a genuine estimate whose via-network attribution is disputed; the crux fraction is genuinely unmeasured. The mixing is the theorem; the fractions are the frontier.

Recompute it yourself: node research/the-fraction-that-reaches-the-tree/verify-the-fraction-that-reaches-the-tree.mjs — and the sources for every cited datum are in research/the-fraction-that-reaches-the-tree/README.md.

What's exactly true here, and what is a modelling choice

Exactly true. Isotope dilution / two-source mixing is the real, standard method, and it is exactly linear in atom fraction — so the forward equation and its inverse are an identity, reproduced live and in the verifier to machine precision. Carbon transfer between EM trees is established (Simard 1997), and the fir's net gain of ≈6% of its own isotope uptake is quoted from the paper, not rounded to flatter the story. The logical point the instrument makes — that a bulk harvest returns the sum P + S + φF and can't return the split — is a fact about the measurement, not an opinion.

Net vs gross. Simard's 6% is a net figure: birch and fir each sent carbon the other way; 6% is the fir-ward balance, and it is a fraction of isotope, not of the fir's total carbon budget. Popular retellings routinely drop all three qualifiers.

Modelling choices. The 5.00 atom% label enrichment and the 1000-unit background pool are illustrative round numbers; the exactness of the mixing identity does not depend on them (the verifier checks it across a random sweep of values). Klein's real experiment used the opposite sign — heavy-carbon-depleted CO₂, which lowers δ¹³C — but the mixing algebra is identical. The four compartments are a deliberately coarse cartoon of a continuous, messy belowground reality; their point is not precision but to show which quantity the disagreement actually lives in.

An interpretation, not a measurement. Robinson & Fitter's mycocentric reading is an argument about how to attribute detected label — which is exactly what makes the partition honest. If the fungal fraction had been measured in the field, there would be no dial to slide.