The concept of iso- vs. anisohydry has been used to describe the stringency of stomatal regulation of plant water potential (ψ). However, metrics that accurately and consistently quantify species' operating ranges along a continuum of iso- to anisohydry have been elusive. Additionally, most approaches to quantifying iso/anisohydry require labour-intensive measurements during prolonged drought. We evaluated new and previously developed metrics of stringency of stomatal regulation of ψ during soil drying in eight woody species and determined whether easily-determined leaf pressure-volume traits could serve as proxies for their degree of iso- vs. anisohydry. Two metrics of stringency of stomatal control of ψ, (1) a 'hydroscape' incorporating the landscape of ψ over which stomata control ψ, and (2) the slope of the daily range of ψ as pre-dawn ψ declined, were strongly correlated with each other and with the leaf osmotic potential at full and zero turgor derived from pressure-volume curves.
Because iso- and anisohydric species differ in stomatal regulation of the rate and magnitude of fluctuations in shoot water potential, they may be expected to show differences in the plasticity of their shoot water relations components, but explicit comparisons of this nature have rarely been made. We subjected excised shoots of co-occurring anisohydric Juniperus monosperma and isohydric Pinus edulis to pressure-volume analysis with and without prior artificial rehydration. In J. monosperma, the shoot water potential at turgor loss (Ψ(TLP)) ranged from -3.4 MPa in artificially rehydrated shoots to -6.6 MPa in shoots with an initial Ψ of -5.5 MPa, whereas in P. edulis mean Ψ(TLP) remained at ∼ -3.0 MPa over a range of initial Ψ from -0.1 to -2.3 MPa. The shoot osmotic potential at full turgor and the bulk modulus of elasticity also declined sharply with shoot Ψ in J. monosperma, but not in P. edulis. The contrasting behaviour of J. monosperma and P. edulis reflects differences in their capacity for homeostatic regulation of turgor that may be representative of aniso- and isohydric species in general, and may also be associated with the greater capacity of J. monosperma to withstand severe drought.
Species' differences in the stringency of stomatal control of plant water potential represent a continuum of isohydric to anisohydric behaviours. However, little is known about how quasi-steady-state stomatal regulation of water potential may relate to dynamic behaviour of stomata and photosynthetic gas exchange in species operating at different positions along this continuum. Here, we evaluated kinetics of light-induced stomatal opening, activation of photosynthesis and features of quasi-steady-state photosynthetic gas exchange in 10 woody species selected to represent different degrees of anisohydry. Based on a previously developed proxy for the degree of anisohydry, species' leaf water potentials at turgor loss, we found consistent trends in photosynthetic gas exchange traits across a spectrum of isohydry to anisohydry. More anisohydric species had faster kinetics of stomatal opening and activation of photosynthesis, and these kinetics were closely coordinated within species. Quasi-steady-state stomatal conductance and measures of photosynthetic capacity and performance were also greater in more anisohydric species. Intrinsic water-use efficiency estimated from leaf gas exchange and stable carbon isotope ratios was lowest in the most anisohydric species. In comparisons between gas exchange traits, species rankings were highly consistent, leading to species-independent scaling relationships over the range of isohydry to anisohydry observed.
Given future climate predictions of increased temperature, and frequency and intensity of heat waves in the tropics, suitable habitat to grow ecologically, economically, and socially valuable Coffea arabica is severely threatened. We investigated how leaf age and heat stress duration impact recovery from heat stress in C. arabica. Treated plants were heated in a growth chamber at 49°C for 45 or 90 min. Physiological recovery was monitored in situ using gas exchange, chlorophyll fluorescence (the ratio of variable to maximum fluorescence, F V/F M), and leaf nonstructural carbohydrate (NSC) on mature and expanding leaves before and 2, 15, 25, and 50 days after treatment. Regardless of leaf age, the 90‐min treatment resulted in greater F V/F M reduction 2 days after treatment and slower recovery than the 45‐min treatment. In both treatments, photosynthesis of expanding leaves recovered more slowly than in mature leaves. Stomatal conductance (g s) decreased in expanding leaves but did not change in mature leaves. These responses led to reduced intrinsic water‐use efficiency with increasing heat stress duration in both age classes. Based on a leaf energy balance model, aftereffects of heat stress would be exacerbated by increases in leaf temperature at low g s under full sunlight where C. arabica is often grown, but also under partial sunlight. Starch and total NSC content of the 45‐min group significantly decreased 2 days after treatment and then accumulated 15 and 25 days after treatment coinciding with recovery of photosynthesis and F V/F M. In contrast, sucrose of the 90‐min group accumulated at day 2 suggesting that phloem transport was inhibited. Both treatment group responses contrasted with control plant total NSC and starch, which declined with time associated with subsequent flower and fruit production. No treated plants produced flowers or fruits, suggesting that short duration heat stress can lead to crop failure.
Temperature and the frequency and intensity of heat waves are predicted to increase throughout the 21st century. Germinant seedlings are expected to be particularly vulnerable to heat stress because they are in the boundary layer close to the soil surface where intense heating occurs in open habitats. We quantified leaf thermotolerance and whole-plant physiological responses to heat stress in first-year germinant seedlings in two populations each of Pinus ponderosa P. and C. Lawson (PIPO) and Pseudotsuga menziesii (Mirb.) Franco (PSME) from climates with contrasting precipitation and temperature regimes. Thermotolerance of detached needles was evaluated using chlorophyll fluorescence (FV/FM, FO) and electrolyte leakage. PSME was more heat tolerant than PIPO according to both independent assessments of thermotolerance. Following exposure of whole seedlings to a simulated heat wave at 45 °C for 1 h in a growth chamber, we monitored FV/FM, photosynthesis, stomatal conductance, non-structural carbohydrates (NSCs) and carbon isotope ratios (δ13C) for 14 days. Heat treatment induced significant reductions in FV/FM in both species and a transient reduction in photosynthetic gas exchange only in PIPO 1 day after treatment. Heat treatment induced an increase in glucose + fructose concurrent with a decrease in starch in both species, whereas total NSC and sucrose were not affected by heat treatment. The negative relationship between glucose + fructose and starch observed in treated plants may be due to the conversion of starch to glucose + fructose to aid recovery from heat-induced damage. Populations from drier sites displayed greater δ13C values than those from wetter sites, consistent with higher intrinsic water-use efficiency and drought resistance of populations from drier climates. Thermotolerance and heat stress responses appeared to be phenotypically plastic and representative of the environment in which plants were grown, whereas intrinsic water-use efficiency appeared to reflect ecotypic differentiation and the climate of origin.
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