Hey! We have a new tree ring isotope paper out in preprint and this one’s on tricky iWUE, so I thought I’d do a quick background thread on trees ,co2 and water: the ultimate complicated threesome! 1/ https://nph.onlinelibrary.wiley.com/doi/10.1111/nph.16811
Trees change the size, geometry, density, & functioning of their stomata when there’s more CO2 available. This changes their water use & photosynthetic rate. There's a lot of trees, so this impacts the C & h20 cycles via plant physiological forcing. 2/ https://onlinelibrary.wiley.com/doi/full/10.1111/j.1365-2486.2010.02273.x?casa_token=gOl-NKVrHXcAAAAA%3AqHAmd31V_oKkNVeqlo5JWfw9xDW6-OTInQp9JwpI_CLcOm-l7Pa_vmvGtLMvwUYGYrQg3ezzkXB2Uw8
Some of the changes are physical & some are changes in the action of tree’s super clever two-way carbon-water system. Summary-Tree stomata; hypersensitive to CO2 since the Devonian https://www.sciencedirect.com/science/article/abs/pii/S0305736485711122.
adapted frm McCarroll & Loader 2004 3/

The stomata seesaw (with CO2 [in] on one end and water [out] on the other) can be studied over time by measuring d13c in tree rings and converting the values to intrinsic Water Use Efficiency, as a stomata-seesaw proxy. When we do that we see, unsurprisingly, a complex pic 4/
The iWUE proxy has always shown a complex story. With some trees showing rising iWUE with rising CO2, but reaching a plateau, some showing no response, and some a reduction in iWUE. https://onlinelibrary.wiley.com/doi/full/10.1111/j.1365-2486.2010.02273.x?casa_token=PkcxLROD0fgAAAAA%3ArkU0ULlsewDk8O8MCk_qritcosaYVI9662oS4ukJTclaJQaKoR72kY4xdgOcP65wBO4CQ_tUjyMuZP0 & evidence that the CO2 response is weakening https://onlinelibrary.wiley.com/doi/full/10.1111/gcb.15166?af=R 5/
So why don’t trees just grow more with more CO2? Most evolved when there was less CO2 so they’re already maxed out & living in a raised CO2 world ( https://besjournals.onlinelibrary.wiley.com/doi/full/10.1046/j.1365-2745.2003.00742). A standard hypothesis was that they would show a mix of passive and active responses. 6/
But, more importantly, rising iWUE, and even capturing more C, doesn’t mean more biomass. https://www.nature.com/articles/s41559-019-1090-0 and https://link.springer.com/article/10.1007/s10021-018-0309-2 and https://www.pnas.org/content/116/7/2749 Summary: More CO2 doesn’t mean more tree. 7/
Other factors limit tree growth when there’s enough CO2. Like moisture and nutrients. https://onlinelibrary.wiley.com/doi/abs/10.1111/gcb.14428#.W4XjjgzCiXg.twitter Although trees may increase their WUE in droughts, https://www.nature.com/articles/s41561-018-0212-7 that doesn’t mean they’re laying down more carbon. 8/
And actually, climate change, and the way we manage forests, is making them dominated by trees that are younger and smaller, over time https://science.sciencemag.org/content/368/6494/eaaz9463 9/
and the latest climate models are aligning around things getting drier as temps rise further https://agupubs.onlinelibrary.wiley.com/doi/10.1029/2020GL087820 “Future drought changes are larger and more consistent in CMIP6 compared to CMIP5“. So rising iWUE in trees, sure, more tree & robust forest, probably not. 10/
So all this makes for a tricky set of variables to consider when it comes to modeling the likely physiological forcing response to future CO2 levels. 11/ https://pubmed.ncbi.nlm.nih.gov/23504999/
So what did we explore in our new paper? https://nph.onlinelibrary.wiley.com/doi/abs/10.1111/nph.16811 We sought to contribute to moving 'iWUE from d13c’ forwards to a point where we can explore how dynamic the response of tree carbon-water balance is to rising CO2 - via their tree ring histories. 12/
The trees we studied had adjusted their water-gas exchange dynamically in response to CO2, moisture, stand dynamics, site conditions. They shifted iWUE between passive/active (and neutral-ish) ci/ca behaviors in response to multiple variables, not in a simple one-way shift. 13/