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Calculating iWUE from carbon isotopes can overestimate the physiological response to rising atmospheric CO2

Posted by wrhorwath on 15 Nov 2013 at 02:28 GMT

The artifact is clearly demonstrated by the fact that the equation always leads to increases in iWUE. Simulated (random) δ13C values, used here to replace tree-ring measurements, are obviously independent from physiological responses and yet lead to increasing iWUE over time. This is explained by the fact that Ca is used as an independent variable in the calculation, artificially producing positive responses as shown in Figure 1.
Indeed according to Fick's law A = g (Ca − Ci) with CO2 and H2O related by a constant factor that links gas exchange of carbon and water vapor. However, in the scenario proposed in the comment above, Ci would have to follow exactly the same increase in Ca to produce a constant Ca − Ci, which is not realistic given that gas exchange and carboxylation continue to occur. The Ci/Ca ratio, on the other hand, reflects the balance between A and g, so the assumption is that the linear relationship between Ci/Ca and 13C discrimination (eq. 2) can be used to calculate iWUE (A/g).
Based on the recent literature, we consider our results in the context of expected physiological responses (such as increases in A or declines g), which should cause changes in 13C discrimination. We found, however, that increases in iWUE would occur in any analysis independently of such physiological responses and respective changes in discrimination. The final outcome of our model reflects (based on the existing equations) what would be observed if Ci increased proportionally with Ca. This is a conservative scenario compared with, for example, constant Ci which would represent much stronger increases in iWUE.
For these reasons, we maintain that previously reported trends in iWUE do not necessarily reflect coherent physiological responses to rising CO2. This conclusion is further supported by the lack of consistent growth responses across biomes (see references cited).

No competing interests declared.

RE: Calculating iWUE from carbon isotopes can overestimate the physiological response to rising atmospheric CO2

jrbrooks replied to wrhorwath on 15 Nov 2013 at 21:45 GMT

In response to your statement: "However, in the scenario proposed in the comment above, Ci would have to follow exactly the same increase in Ca to produce a constant Ca − Ci, which is not realistic given that gas exchange and carboxylation continue to occur".
According to Fick’s law (A/g = ca - ci), a constant photosynthetic rate and a constant conductance to CO2 would result in a constant drop in CO2 from outside to in the leaf, regardless of the CO2 concentration outside the leaf (ca). That is the law of diffusion. Thus, a A of 10, and a g of 0.1 results in a drop of 100 ppm (ca - ci). According to the law, the only way for the drawdown in CO2 inside the leaf to get larger would be for either A or g to change, no matter what the outside CO2 concentration is. If you look at your cited references to Farquhar’s original derivation of ~13~C discrimination, he uses Fick’s law. If you are declaring equation 3 Fick's law incorrect, then equation 2 would also have to be incorrect. So in response to your statement above, if physiology (A/g) does not change with rising CO2, then ci will follow the same increase as ca keeping ca - ci constant over time.

No competing interests declared.

RE: RE: Calculating iWUE from carbon isotopes can overestimate the physiological response to rising atmospheric CO2

LucascrSilva replied to jrbrooks on 15 Nov 2013 at 22:58 GMT

As explicitly stated in the paper, we agree with the theory and the well-established association found between δ13C and leaf-level physiological processes. However, our results show that the extrapolation of this association for the temporal reconstruction of iWUE should be reevaluated. The message is simple, increases in iWUE occur independently of changes in δ13C. Any tree-ring data set would inevitably generate positive responses over time, making it impossible to differentiate real from artificial effects.

No competing interests declared.

RE: RE: RE: Calculating iWUE from carbon isotopes can overestimate the physiological response to rising atmospheric CO2

jrbrooks replied to LucascrSilva on 15 Nov 2013 at 23:19 GMT

What you showed is that trees with a constant ci/ca over time with increasing CO2 (with 1 per mill variance around the mean) are increasing their iWUE. Not every tree ring data set would show this. See for example Marshall and Monserud (1996) where they find a constant (ca-ci). What I dispute is that you claim the gas-exchange physiology of trees with ci/ca of 0.5 at 300 and 400 ppm CO2 would be the same, and they are not.

Marshall, J. D. and R. A. Monserud. 1996. Homeostatic gas-exchange parameters inferred from 13C/12C in tree rings of conifers. Oecologia 105:13-21.

No competing interests declared.