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The 18O enrichment (Δ18O) of cellulose (Δ18OCel) is recognized as a unique archive of past climate and plant function. However, there is still uncertainty regarding the proportion of oxygen in cellulose (pex) that exchanges post‐photosynthetically with medium water of cellulose synthesis. Particularly, recent research with C3 grasses demonstrated that the Δ18O of leaf sucrose (Δ18OSuc, the parent substrate for cellulose synthesis) can be much higher than predicted from daytime Δ18O of leaf water (Δ18OLW), which could alter conclusions on photosynthetic versus post‐photosynthetic effects on Δ18OCel via pex. Here, we assessed pex in leaves of perennial ryegrass (Lolium perenne) grown at different atmospheric relative humidity (RH) and CO2 levels, by determinations of Δ18OCel in leaves, Δ18OLGDZW (the Δ18O of water in the leaf growth‐and‐differentiation zone) and both Δ18OSuc and Δ18OLW (adjusted for εbio, the biosynthetic fractionation between water and carbohydrates) as alternative proxies for the substrate for cellulose synthesis. Δ18OLGDZW was always close to irrigation water, and pex was similar (0.53 ± 0.02 SE) across environments when determinations were based on Δ18OSuc. Conversely, pex was erroneously and variably underestimated (range 0.02–0.44) when based on Δ18OLW. The photosynthetic signal fraction in Δ18OCel is much more constant than hitherto assumed, encouraging leaf physiological reconstructions. Summary statement: The proportion of oxygen in leaf cellulose which is derived from leaf sucrose in Lolium perenne is much more constant and predictable than was hitherto assumed, encouraging reconstructions of photosynthetic information [ABSTRACT FROM AUTHOR]

The 13C isotope composition (δ13C) of leaf dry matter is a useful tool for physiological and ecological studies. However, how post-photosynthetic fractionation associated with respiration and carbon export influences δ13C remains uncertain. We investigated the effects of post-photosynthetic fractionation on δ13C of mature leaves of Cleistogenes squarrosa , a perennial C4 grass, in controlled experiments with different levels of vapour pressure deficit and nitrogen supply. With increasing leaf age class, the 12C/13C fractionation of leaf organic matter relative to the δ13C of atmosphere CO2 (ΔDM) increased while that of cellulose (Δcel) was almost constant. The divergence between ΔDM and Δcel increased with leaf age class, with a maximum value of 1.6‰, indicating the accumulation of post-photosynthetic fractionation. Applying a new mass balance model that accounts for respiration and export of photosynthates, we found an apparent 12C/13C fractionation associated with carbon export of –0.5‰ to –1.0‰. Different ΔDM among leaves, pseudostems, daughter tillers, and roots indicate that post-photosynthetic fractionation happens at the whole-plant level. Compared with ΔDM of old leaves, ΔDM of young leaves and Δcel are more reliable proxies for predicting physiological parameters due to the lower sensitivity to post-photosynthetic fractionation and the similar sensitivity in responses to environmental changes. [ABSTRACT FROM AUTHOR]

The 18O enrichment (Δ18O) of leaf water affects the Δ18O of photosynthetic products such as sucrose, generating an isotopic archive of plant function and past climate. However, uncertainty remains as to whether leaf water compartmentation between photosynthetic and nonphotosynthetic tissue affects the relationship between Δ18O of bulk leaf water (Δ18OLW) and leaf sucrose (Δ18OSucrose). We grew Lolium perenne (a C3 grass) in mesocosm‐scale, replicated experiments with daytime relative humidity (50% or 75%) and CO2 level (200, 400 or 800 μmol mol−1) as factors, and determined Δ18OLW, Δ18OSucrose and morphophysiological leaf parameters, including transpiration (Eleaf), stomatal conductance (gs) and mesophyll conductance to CO2 (gm). The Δ18O of photosynthetic medium water (Δ18OSSW) was estimated from Δ18OSucrose and the equilibrium fractionation between water and carbonyl groups (εbio). Δ18OSSW was well predicted by theoretical estimates of leaf water at the evaporative site (Δ18Oe) with adjustments that correlated with gas exchange parameters (gs or total conductance to CO2). Isotopic mass balance and published work indicated that nonphotosynthetic tissue water was a large fraction (~0.53) of bulk leaf water. Δ18OLW was a poor proxy for Δ18OSucrose, mainly due to opposite Δ18O responses of nonphotosynthetic tissue water (Δ18Onon‐SSW) relative to Δ18OSSW, driven by atmospheric conditions. Summary Statement: The 18O enrichment of water in the two functionally distinct water pools of grasses—photosynthetic and nonphotosynthetic tissue water—revealed opposite responses to atmospheric humidity in Lolium perenne. 18O enrichment of sucrose was close to equilibrium with evaporative site enrichment. [ABSTRACT FROM AUTHOR]