Dry inside: progressive unsaturation within leaves with increasing vapour pressure deficit affects estimation of key leaf gas exchange parameters.

Climate change not only leads to higher air temperatures but also increases the vapour pressure deficit (VPD) of the air. Understanding the direct effect of VPD on leaf gas exchange is crucial for precise modelling of stomatal functioning. We conducted combined leaf gas exchange and online isotope discrimination measurements on four common European tree species across a VPD range of 0.8-3.6 kPa, while maintaining constant temperatures without soil water limitation. In addition to applying the standard assumption of saturated vapour pressure inside leaves (ei), we inferred ei from oxygen isotope discrimination of CO2 and water vapour. ei desaturated progressively with increasing VPD, consistently across species, resulting in an intercellular relative humidity as low as 0.73 ± 0.11 at the highest tested VPD. Assuming saturation of ei overestimated the extent of reductions in stomatal conductance and CO2 mole fraction inside leaves in response to increasing VPD compared with calculations that accounted for unsaturation. In addition, a significant decrease in mesophyll conductance with increasing VPD only occurred when the unsaturation of ei was considered. We suggest that the possibility of unsaturated ei should not be overlooked in measurements related to leaf gas exchange and in stomatal models, especially at high VPD.

Impacts of elevated temperature and vapour pressure deficit on leaf gas exchange and plant growth across six tropical rainforest tree species.

Elevated air temperature (Tair) and vapour pressure deficit (VPDair) significantly influence plant functioning, yet their relative impacts are difficult to disentangle. We examined the effects of elevated Tair (+6°C) and VPDair (+0.7 kPa) on the growth and physiology of six tropical tree species. Saplings were grown under well-watered conditions in climate-controlled glasshouses for 6 months under three treatments: (1) low Tair and low VPDair, (2) high Tair and low VPDair, and (3) high Tair and high VPDair. To assess acclimation, physiological parameters were measured at a set temperature. Warm-grown plants grown under elevated VPDair had significantly reduced stomatal conductance and increased instantaneous water use efficiency compared to plants grown under low VPDair. Photosynthetic biochemistry and thermal tolerance (Tcrit) were unaffected by VPDair, but elevated Tair caused Jmax25 to decrease and Tcrit to increase. Sapling biomass accumulation for all species responded positively to an increase in Tair, but elevated VPDair limited growth. This study shows that stomatal limitation caused by even moderate increases in VPDair can decrease productivity and growth rates in tropical species independently from Tair and has important implications for modelling the impacts of climate change on tropical forests.

Dynamic responses of gas exchange and photochemistry to heat interference during drought in wheat and sorghum.

Drought and heat stress significantly affect crop growth and productivity worldwide. It is unknown how heat interference during drought affects physiological processes dynamically in crops. Here we focussed on gas exchange and photochemistry in wheat and sorghum in response to simulated heat interference via +15°C of temperature during ~2 week drought and re-watering. Results showed that drought decreased net photosynthesis (Anet), stomatal conductance (gs), maximum velocity of ribulose-1, 5-bisphosphate carboxylase/oxygenase carboxylation (Vcmax) and electron transport rate (J) in both wheat and sorghum. Heat interference did not further reduce Anet or gs. Drought increased non-photochemical quenching (Φnpq), whereas heat interference decreased Φnpq. The δ13C of leaf, stem and roots was higher in drought-treated wheat but lower in drought-treated sorghum. The results suggest that (1) even under drought conditions wheat and sorghum increased or maintained gs for transpirational cooling to alleviate negative effects by heat interference; (2) non-photochemical quenching responded differently to drought and heat stress; (3) wheat and sorghum responded in opposing patterns in δ13C. These findings point to the importance of stomatal regulation under heat crossed with drought stress and could provide useful information on development of better strategies to secure crop production for future climate change.

Effect of Vapor Pressure Deficit on Gas Exchange in Wild-Type and Abscisic Acid-Insensitive Plants.

Stomata control the gas exchange of terrestrial plant leaves, and are therefore essential to plant growth and survival. We investigated gas exchange responses to vapor pressure deficit (VPD) in two gray poplar (Populus × canescens) lines: wild type and abscisic acid-insensitive (abi1) with functionally impaired stomata. Transpiration rate in abi1 increased linearly with VPD, up to about 2 kPa. Above this, sharply declining transpiration was followed by leaf death. In contrast, wild type showed a steady or slightly declining transpiration rate up to VPD of nearly 7 kPa, and fully recovered photosynthetic function afterward. There were marked differences in discrimination against 13CO2 (Δ13C) and C18OO (Δ18O) between abi1 and wild-type plants. The Δ13C indicated that intercellular CO2 concentrations decreased with VPD in wild-type plants, but not in abi1 plants. The Δ18O reflected progressive stomatal closure in wild type in response to increasing VPD; however, in abi1, stomata remained open and oxygen atoms of CO2 continued to exchange with 18O enriched leaf water. Coupled measurements of Δ18O and gas exchange were used to estimate intercellular vapor pressure, e i In wild-type leaves, there was no evidence of unsaturation of e i, even at VPD above 6 kPa. In abi1 leaves, e i approached 0.6 times saturation vapor pressure before the precipitous decline in transpiration rate. For wild type, a sensitive stomatal response to increasing VPD was pivotal in preventing unsaturation of e i In abi1, after taking unsaturation into account, stomatal conductance increased with increasing VPD, consistent with a disabled active response of guard cell osmotic pressure.

Ternary effects on the gas exchange of isotopologues of carbon dioxide.

The ternary effects of transpiration rate on the rate of assimilation of carbon dioxide through stomata, and on the calculation of the intercellular concentration of carbon dioxide, are now included in standard gas exchange studies. However, the equations for carbon isotope discrimination and for the exchange of oxygen isotopologues of carbon dioxide ignore ternary effects. Here we introduce equations to take them into account. The ternary effect is greatest when the leaf-to-air vapour mole fraction difference is greatest, and its impact is greatest on parameters derived by difference, such as the mesophyll resistance to CO(2) assimilation, r(m) . We show that the mesophyll resistance to CO(2) assimilation has been underestimated in the past. The impact is also large when there is a large difference in isotopic composition between the CO(2) inside the leaf and that in the air. We show that this partially reconciles estimates of the oxygen isotopic composition of CO(2) in the chloroplast and mitochondria in the light and in the dark, with values close to equilibrium with the estimated oxygen isotopic composition of water at the sites of evaporation within the leaf.

Transpiration efficiency over an annual cycle, leaf gas exchange and wood carbon isotope ratio of three tropical tree species.

Variation in transpiration efficiency (TE) and its relationship with the stable carbon isotope ratio of wood was investigated in the saplings of three tropical tree species. Five individuals each of Platymiscium pinnatum (Jacq.) Dugand, Swietenia macrophylla King and Tectona grandis Linn. f. were grown individually in large (760 l) pots over 16 months in the Republic of Panama. Cumulative transpiration was determined by repeatedly weighing the pots with a pallet truck scale. Dry matter production was determined by destructive harvest. The TE, expressed as experiment-long dry matter production divided by cumulative water use, averaged 4.1, 4.3 and 2.9 g dry matter kg(-1) water for P. pinnatum, S. macrophylla and T. grandis, respectively. The TE of T. grandis was significantly lower than that of the other two species. Instantaneous measurements of the ratio of intercellular to ambient CO(2) partial pressures (c(i)/c(a)), taken near the end of the experiment, explained 66% of variation in TE. Stomatal conductance was lower in S. macrophylla than in T. grandis, whereas P. pinnatum had similar stomatal conductance to T. grandis, but with a higher photosynthetic rate. Thus, c(i)/c(a) and TE appeared to vary in response to both stomatal conductance and photosynthetic capacity. Stem-wood delta(13)C varied over a relatively narrow range of just 2.2 per thousand, but still explained 28% of variation in TE. The results suggest that leaf-level processes largely determined variation among the three tropical tree species in whole-plant water-use efficiency integrated over a full annual cycle.

Stem and leaf gas exchange and their responses to fire in a north Australian tropical savanna.

We measured stem CO2 efflux and leaf gas exchange in a tropical savanna ecosystem in northern Australia, and assessed the impact of fire on these processes. Gas exchange of mature leaves that flushed after a fire showed only slight differences from that of mature leaves on unburned trees. Expanding leaves typically showed net losses of CO2 to the atmosphere in both burned and unburned trees, even under saturating irradiance. Fire caused stem CO2 efflux to decline in overstory trees, when measured 8 weeks post-fire. This decline was thought to have resulted from reduced availability of C substrate for respiration, due to reduced canopy photosynthesis caused by leaf scorching, and to priority allocation of fixed C towards reconstruction of a new canopy. At the ecosystem scale, we estimated the annual above-ground woody-tissue CO2 efflux to be 275 g C m(-2) ground area year(-1) in a non-fire year, or approximately 13% of the annual gross primary production. We contrasted the canopy physiology of two co-dominant overstory tree species, one of which has a smooth bark on its branches capable of photosynthetic re-fixation (Eucalyptus miniata), and the other of which has a thick, rough bark incapable of re-fixation (Eucalyptus tetrodonta). Eucalyptus miniata supported a larger branch sapwood cross-sectional area in the crown per unit subtending leaf area, and had higher leaf stomatal conductance and photosynthesis than E. tetrodonta. Re-fixation by photosynthetic bark reduces the C cost of delivering water to evaporative sites in leaves, because it reduces the net C cost of constructing and maintaining sapwood. We suggest that re-fixation allowed leaves of E. miniata to photosynthesize at higher rates than those of E. tetrodonta, while the two invested similar amounts of C in the maintenance of branch sapwood.