Shade cloth can be used to protect grapevines from high temperatures. However, the resulting low light intensity is shown to reduce photosynthesis, leading to lower carbon allocation to vegetative growth and sugar accumulation. Protection from heat by shading is, therefore, costly for the carbon economy of the vines.
Background and aims
Covering whole vines with shade cloth is used to protect the vines from heat stress, but may have costs on vine productivity through reduced light availability. Our aim was to assess the carbon balance of vines growing with and without shade to quantify the impact of the covering.
Whole vines were covered with 70 % shade cloth, and shoot leaf area and leaf, stem and bunch growth were followed over two growing seasons. Photosynthesis was measured in situ in all leaves along selected shoots over the growing season. A carbon balance was constructed from the difference in acquisition of carbon and the sequestration of carbon as biomass across the growing seasons.
Shade covering had no initial impact on shoot growth but later reduced leaf growth and later still bunch growth. Stem growth was unaffected. Photosynthetic properties were characteristic of shade leaves, with lower rates and lower light saturation compared with well-exposed leaves. Overall, net photosynthesis was reduced by 40 % by the shade covering and was attributed to the reduced photon flux densities. From the carbon balance, vines were reliant on carbon reserves over 6 weeks after budbreak until current photosynthate increased sufficiently to supply the growth. Shade covering impacted most on biomass accumulation to leaves and bunches at the stage when the vines became autotrophic, consistent with the reduction in carbon acquisition. The markedly high carbon demand by bunches caused a mid-season negative carbon balance, implying that shoots had to draw further on reserves to supply the carbon.
Shade covering over whole grapevines exacerbated the imbalance between the supply of and demand for carbon and greatly reduced vine biomass, especially reproductive allocation. Covering vines with shade cloth to protect the vines from heat events, therefore, had major costs in the carbon economy.
Different spatial distributions of soil moisture were imposed on field-grown grapevines by applying the same irrigation volumes to the entire (DI; deficit irrigation) or part of the (PRD; partial root zone drying) root zone. Five treatments were applied: controls irrigated at 60% ETc (crop evapotranspiration) for the whole season (308 mm year−1); DI-1 and PRD-1 that received the same irrigation as controls before fruit set, 30% ETc from fruit set to harvest and 45% ETc post-harvest (192 mm year−1); and DI-2 and PRD-2 that were the same, except that 15% ETc was applied from fruit set to harvest (142 mm year−1). Compared with DI-1, PRD-1 maintained higher leaf area post-veraison and increased root water uptake, whole-plant hydraulic conductance, leaf transpiration, stomatal conductance, and photosynthesis, but decreased intrinsic gas exchange efficiency without causing differences in leaf xylem abscisic acid (ABA) concentration. Compared with DI-2, PRD-2 increased leaf xylem ABA concentration and decreased root water uptake, whole-plant hydraulic conductance, leaf transpiration, stomatal conductance, and photosynthesis, mainly at the beginning of PRD cycles. Distinctive PRD effects (e.g. greater stomatal closure) depended on the volumetric soil water content of the wet root zone, as predicted from a model of root-to-shoot ABA signalling.
ABA signalling; deficit irrigation; leaf water relations; partial root zone drying; root water uptake; soil water content
Background and Aims
Daytime root-zone temperature may be a significant factor regulating water flux through plants. Water flux can also occur during the night but nocturnal stomatal response to environmental drivers such as root-zone temperature remains largely unknown.
Here nocturnal and daytime leaf gas exchange was quantified in ‘Shiraz’ grapevines (Vitis vinifera) exposed to three root-zone temperatures from budburst to fruit-set, for a total of 8 weeks in spring.
Despite lower stomatal density, night-time stomatal conductance and transpiration rates were greater for plants grown in warm root-zones. Elevated root-zone temperature resulted in higher daytime stomatal conductance, transpiration and net assimilation rates across a range of leaf-to-air vapour pressure deficits, air temperatures and light levels. Intrinsic water-use efficiency was, however, lowest in those plants with warm root-zones. CO2 response curves of foliar gas exchange indicated that the maximum rate of electron transport and the maximum rate of Rubisco activity did not differ between the root-zone treatments, and therefore it was likely that the lower photosynthesis in cool root-zones was predominantly the result of a stomatal limitation. One week after discontinuation of the temperature treatments, gas exchange was similar between the plants, indicating a reversible physiological response to soil temperature.
In this anisohydric grapevine variety both night-time and daytime stomatal conductance were responsive to root-zone temperature. Because nocturnal transpiration has implications for overall plant water status, predictive climate change models using stomatal conductance will need to factor in this root-zone variable.
Water-use efficiency; leaf gas exchange; CO2 assimilation; grapevine; Vitis vinifera; stomatal conductance; root-zone temperature; ‘Shiraz’; grapevine; soil temperature
The heat event that occurred in many parts of Australia in 2009 was the worst on record for the past decade, with air temperatures exceeding 40°C for 14 days. Our aim was to assess the impacts of this heat event on vine performance, including ripening, yield, and gas exchange of Vitis vinifera cv. Semillon grown in a Riverina vineyard. To assess the affect of high temperatures on Semillon grapevines, the vines were covered with a protective layer to reduce radiant heating and were compared with vines exposed to ambient conditions. The heat event had major effects on ripening; reducing the rate of ripening by 50% and delaying harvest ripeness and causing a high incidence of berry shrivel and sunburn. Yield was not affected. Photosynthesis was reduced 35% by the heat event while transpiration increased nearly threefold and was accounted for by increased stomatal conductance. The conclusion of this study was that heat events delayed ripening in Semillon berries and caused a significant reduction in berry quality. Strategies to minimize the radiant load during heat events are required and this study has confirmed a protective layer can reduce canopy temperatures and enhance berry quality.
photosynthesis; rate of ripening; soluble solids concentration; stomatal conductance; transpiration; yield
Background and Aims
Leaf responses to environmental conditions have been frequently described in fruit trees, but differences among cultivars have received little attention. This study shows that parameters of Farquhar's photosynthesis and Jarvis' stomatal conductance models differed between two apple cultivars, and examines the consequences of these differences for leaf water use efficiency.
Leaf stomatal conductance (gsw), net CO2 assimilation rate (An), respiration (Rd) and transpiration (E) were measured during summer in 8-year-old ‘Braeburn’ and ‘Fuji’ apple trees under well-watered field conditions. Parameters of Farquhar's and Jarvis' models were estimated, evaluated and then compared between cultivars. Leaf carbon isotope discrimination (Δ13C) was measured at the end of the growing season.
A single positive relationship was established between VCmax (maximum carboxylation rate) and Na (leaf nitrogen concentration per unit area), and between Jmax (maximum light-driven electron transport rate) and Na. A higher leaf Rd was observed in ‘Fuji’. The gsw responded similarly to increasing irradiance and leaf temperature in both cultivars. gsw responded to lower vapour pressure deficit in ‘Fuji’ than in ‘Braeburn’. Maximal conductance (gswmax) was significantly smaller and An was more limited by gsw in ‘Braeburn’ than ‘Fuji’. Lower gsw, E and higher intrinsic water use efficiency were shown in ‘Braeburn’ and confirmed by smaller leaf Δ13C compared with ‘Fuji’ leaves.
The use of functional model parameters allowed comparison of the two cultivars and provided evidence of different water use ‘strategies’: ‘Braeburn’ was more conservative in water use than ‘Fuji’, due to stomatal limitation of An, higher intrinsic water use efficiency and lower Δ13C. These physiological traits need to be considered in relation to climate adaptation, breeding of new cultivars and horticultural practice.
Apple; carbon isotope discrimination; leaf nitrogen; leaf temperature; irradiance; Malus × domestica; modelling; photosynthesis; stomata; transpiration; vapour pressure deficit; water use efficiency
Models seldom consider the effect of leaf-level biochemical acclimation to temperature when scaling forest water use. Therefore, the dependence of transpiration on temperature acclimation was investigated at the within-crown scale in climatically contrasting genotypes of Acer rubrum L., cv. October Glory (OG) and Summer Red (SR). The effects of temperature acclimation on intracanopy gradients in transpiration over a range of realistic forest growth temperatures were also assessed by simulation. Physiological parameters were applied, with or without adjustment for temperature acclimation, to account for transpiration responses to growth temperature. Both types of parameterization were scaled up to stand transpiration (expressed per unit leaf area) with an individual tree model (MAESTRA) to assess how transpiration might be affected by spatial and temporal distributions of foliage properties. The MAESTRA model performed well, but its reproducibility was dependent on physiological parameters acclimated to daytime temperature. Concordance correlation coefficients between measured and predicted transpiration were higher (0.95 and 0.98 versus 0.87 and 0.96) when model parameters reflected acclimated growth temperature. In response to temperature increases, the southern genotype (SR) transpiration responded more than the northern (OG). Conditions of elevated long-term temperature acclimation further separate their transpiration differences. Results demonstrate the importance of accounting for leaf-level physiological adjustments that are sensitive to microclimate changes and the use of provenance-, ecotype-, and/or genotype-specific parameter sets, two components likely to improve the accuracy of site-level and ecosystem-level estimates of transpiration flux.
Intraspecific acclimation; MAESTRA; microclimate; modelling; red maple; temperature acclimation; transpiration
Reduction of hydraulic conductance to the canopy has been shown to result in stomatal responses to limit transpiration. To test for similar responses to perturbations of the hydraulic network in leaves, we simultaneously measured leaf gas exchange with spatially explicit chlorophyll-a fluorescence and leaf temperature to examine the effects of cutting a primary leaf vein in Helianthus annuus. We repeated the leaf treatment at each of three different vapor pressure deficits and monitored the short-term dynamics of gas exchange following the treatment. Immediately after treatment, photosynthesis and stomatal conductance (gs) showed a transient “wrong way” response in which photosynthesis declined despite increased gs. Comparisons of fluorescence and temperature across the leaf showed that both photosynthesis and gs were transiently patchy across the measured leaf area, but that the patchiness of the two processes did not correspond in space or time. This suggests that photosynthesis and gs respond to vein cutting-induced cavitation via different mechanisms. Because the stomatal response varied by vapor pressure difference condition but photosynthesis did not, it is likely that gs, but not photosynthesis, responded to a hydraulic signal. In contrast, we hypothesize that photosynthesis declined due to a wound-induced electrical signal that has recently been shown to transiently decrease mesophyll conductance to CO2. The interaction of epidermal hydraulics and the electrical signal across the leaf likely created a patchy pattern of chlorophyll fluorescence and leaf temperature that cannot be explained through the action of a single signal.
stomatal conductance; leaf hydraulic conductance; mesophyll conductance; stomatal patchiness; chlorophyll fluorescence imaging; photosynthesis; transpiration; cavitation
A spatially explicit mechanistic model, MAESTRA, was used to separate key parameters affecting transpiration to provide insights into the most influential parameters for accurate predictions of within-crown and within-canopy transpiration. Once validated among Acer rubrum L. genotypes, model responses to different parameterization scenarios were scaled up to stand transpiration (expressed per unit leaf area) to assess how transpiration might be affected by the spatial distribution of foliage properties. For example, when physiological differences were accounted for, differences in leaf width among A. rubrum L. genotypes resulted in a 25% difference in transpiration. An in silico within-canopy sensitivity analysis was conducted over the range of genotype parameter variation observed and under different climate forcing conditions. The analysis revealed that seven of 16 leaf traits had a ≥5% impact on transpiration predictions. Under sparse foliage conditions, comparisons of the present findings with previous studies were in agreement that parameters such as the maximum Rubisco-limited rate of photosynthesis can explain ∼20% of the variability in predicted transpiration. However, the spatial analysis shows how such parameters can decrease or change in importance below the uppermost canopy layer. Alternatively, model sensitivity to leaf width and minimum stomatal conductance was continuous along a vertical canopy depth profile. Foremost, transpiration sensitivity to an observed range of morphological and physiological parameters is examined and the spatial sensitivity of transpiration model predictions to vertical variations in microclimate and foliage density is identified to reduce the uncertainty of current transpiration predictions.
Boundary layer conductance; leaf width; modelling; sensitivity analysis; stomatal conductance; transpiration; water vapour transfer; wind
To understand the physiological basis of genetic variation and resulting quantitative trait loci (QTLs) for photosynthesis in a rice (Oryza sativa L.) introgression line population, 13 lines were studied under drought and well-watered conditions, at flowering and grain filling. Simultaneous gas exchange and chlorophyll fluorescence measurements were conducted at various levels of incident irradiance and ambient CO2 to estimate parameters of a model that dissects photosynthesis into stomatal conductance (g
s), mesophyll conductance (g
m), electron transport capacity (J
max), and Rubisco carboxylation capacity (V
cmax). Significant genetic variation in these parameters was found, although drought and leaf age accounted for larger proportions of the total variation. Genetic variation in light-saturated photosynthesis and transpiration efficiency (TE) were mainly associated with variation in g
s and g
m. One previously mapped major QTL of photosynthesis was associated with variation in g
s and g
m, but also in J
max and V
cmax at flowering. Thus, g
s and g
m, which were demonstrated in the literature to be responsible for environmental variation in photosynthesis, were found also to be associated with genetic variation in photosynthesis. Furthermore, relationships between these parameters and leaf nitrogen or dry matter per unit area, which were previously found across environmental treatments, were shown to be valid for variation across genotypes. Finally, the extent to which photosynthesis rate and TE can be improved was evaluated. Virtual ideotypes were estimated to have 17.0% higher photosynthesis and 25.1% higher TE compared with the best genotype investigated. This analysis using introgression lines highlights possibilities of improving both photosynthesis and TE within the same genetic background.
drought; genetic variation; mesophyll conductance; modelling; Oryza sativa L.; photosynthesis; rice; stomatal conductance
Background and Aims
Global climate models predict decreases in leaf stomatal conductance and transpiration due to increases in atmospheric CO2. The consequences of these reductions are increases in soil moisture availability and continental scale run-off at decadal time-scales. Thus, a theory explaining the differential sensitivity of stomata to changing atmospheric CO2 and other environmental conditions must be identified. Here, these responses are investigated using optimality theory applied to stomatal conductance.
An analytical model for stomatal conductance is proposed based on: (a) Fickian mass transfer of CO2 and H2O through stomata; (b) a biochemical photosynthesis model that relates intercellular CO2 to net photosynthesis; and (c) a stomatal model based on optimization for maximizing carbon gains when water losses represent a cost. Comparisons between the optimization-based model and empirical relationships widely used in climate models were made using an extensive gas exchange dataset collected in a maturing pine (Pinus taeda) forest under ambient and enriched atmospheric CO2.
Key Results and Conclusion
In this interpretation, it is proposed that an individual leaf optimally and autonomously regulates stomatal opening on short-term (approx. 10-min time-scale) rather than on daily or longer time-scales. The derived equations are analytical with explicit expressions for conductance, photosynthesis and intercellular CO2, thereby making the approach useful for climate models. Using a gas exchange dataset collected in a pine forest, it is shown that (a) the cost of unit water loss λ (a measure of marginal water-use efficiency) increases with atmospheric CO2; (b) the new formulation correctly predicts the condition under which CO2-enriched atmosphere will cause increasing assimilation and decreasing stomatal conductance.
Economics of gas exchange; free air CO2 enrichment; marginal water-use efficiency; photosynthesis; Pinus taeda; stomatal conductance; stomatal optimization
• Background and Aims Kaolin applications have been used to mitigate the negative effects of water and heat stress on plant physiology and productivity with variable results, ranging from increased to decreased yields and photosynthetic rates. The mechanisms of action of kaolin applications are not clear: although the increased albedo reduces leaf temperature and the consequent heat stress, it also reduces the light available for photosynthesis, possibly offsetting benefits of lower temperature. The objective of this study was to investigate which of these effects are prevalent and under which conditions.
• Methods A 6 % kaolin suspension was applied on well-irrigated and water-stressed walnut (Juglans regia) and almond (Prunus dulcis) trees. Water status (i.e. stem water potential, Ψs), gas exchange (i.e. light-saturated CO2 assimilation rate, Amax; stomatal conductance, gs), leaf temperature (Tl) and physiological relationships in treated and control trees were then measured and compared.
• Key Results In both species, kaolin did not affect the daily course of Ψs whereas it reduced Amax by 1–4 μmol CO2 m–2 s–1 throughout the day in all combinations of species and irrigation treatments. Kaolin did not reduce gs in any situation. Consequently, intercellular CO2 concentration (Ci) was always greater in treated trees than in controls, suggesting that the reduction of Amax with kaolin was not due to stomatal limitations. Kaolin reduced leaf temperature (Tl) by about 1–3 °C and leaf-to-air vapour pressure difference (VPDl) by about 0·1–0·7 kPa. Amax was lower at all values of gs, Tl and VPDl in kaolin-treated trees. Kaolin affected the photosynthetic response to the photosynthetically active radiation (PAR) in almond leaves: kaolin-coated leaves had similar dark respiration rates and light-saturated photosynthesis, but a higher light compensation point and lower apparent quantum yield, while the photosynthetic light-response curve saturated at higher PAR. When these parameters were used to model the photosynthetic response curve to PAR, it was estimated that the kaolin film allowed 63 % of the incident PAR to reach the leaf.
• Conclusions The main effect of kaolin application was the reduction, albeit minor, of photosynthesis, which appeared to be related to the shading of the leaves. The reduction in Tl and VPDl with kaolin did not suffice to mitigate the adverse effects of heat and water stress on Amax.
Juglans regia; kaolin particle film; photosynthesis; Prunus dulcis; stomatal conductance; water potential; stress
Using a stomatal optimization model, an abrupt change is foundnd in the relationship between carbon and water fluxes along the evolutionary gradient from C3 to C4 photosynthetic types.
C4 photosynthesis evolved independently numerous times, probably in response to declining atmospheric CO2 concentrations, but also to high temperatures and aridity, which enhance water losses through transpiration. Here, the environmental factors controlling stomatal behaviour of leaf-level carbon and water exchange were examined across the evolutionary continuum from C3 to C4 photosynthesis at current (400 μmol mol–1) and low (280 μmol mol–1) atmospheric CO2 conditions. To this aim, a stomatal optimization model was further developed to describe the evolutionary continuum from C3 to C4 species within a unified framework. Data on C3, three categories of C3–C4 intermediates, and C4
Flaveria species were used to parameterize the stomatal model, including parameters for the marginal water use efficiency and the efficiency of the CO2-concentrating mechanism (or C4 pump); these two parameters are interpreted as traits reflecting the stomatal and photosynthetic adjustments during the C3 to C4 transformation. Neither the marginal water use efficiency nor the C4 pump strength changed significantly from C3 to early C3–C4 intermediate stages, but both traits significantly increased between early C3–C4 intermediates and the C4-like intermediates with an operational C4 cycle. At low CO2, net photosynthetic rates showed continuous increases from a C3 state, across the intermediates and towards C4 photosynthesis, but only C4-like intermediates and C4 species (with an operational C4 cycle) had higher water use efficiencies than C3
Flaveria. The results demonstrate that both the marginal water use efficiency and the C4 pump strength increase in C4
Flaveria to improve their photosynthesis and water use efficiency compared with C3 species. These findings emphasize that the advantage of the early intermediate stages is predominantly carbon based, not water related.
C3–C4 intermediates; C3 photosynthesis; leaf gas exchange; photosynthetic model; stomatal conductance; water use efficiency.
The tolerance of lettuce plants (Lactuca sativa L. cv. Romana) to drought stress differed with the arbuscular-mycorrhizal fungal isolate with which the plants were associated. Seven fungal species belonging to the genus Glomus were studied for their ability to enhance the drought tolerance of lettuce plants. These fungi had different traits that affected the drought resistance of host plants. The ranking of arbuscular-mycorrhizal fungal effects on drought tolerance, based on the relative decreases in shoot dry weight, was as follows: Glomus deserticola > Glomus fasciculatum > Glomus mosseae > Glomus etunicatum > Glomus intraradices > Glomus caledonium > Glomus occultum. In this comparative study specific mycorrhizal fungi had consistent effects on plant growth, mineral uptake, the CO(inf2) exchange rate, water use efficiency, transpiration, stomatal conductance, photosynthetic phosphorus use efficiency, and proline accumulation under either well-watered or drought-stressed conditions. The ability of the isolates to maintain plant growth effectively under water stress conditions was related to higher transpiration rates, levels of leaf conductance, and proline, N, and P contents. Differences in proline accumulation in leaves among the fungal symbioses suggested that the fungi were able to induce different degrees of osmotic adjustment. The detrimental effects of drought were not related to decreases in photosynthesis or water use efficiency. Neither of these parameters was related to P nutrition. The differences in P and K acquisition, transpiration, and stomatal conductance were related to the mycorrhizal efficiencies of the different fungi. Our observations revealed the propensities of different Glomus species to assert their protective effects during plant water stress. The greater effectiveness of G. deserticola in improving water deficit tolerance was associated with the lowest level of growth reduction (9%) under stress conditions. The growth of plants colonized by G. occultum was reduced by 70% after a progressive drought stress period. In general, the different protective effects of the mycorrhizal isolates were not associated with colonizing ability. Nevertheless, G. deserticola was the most efficient fungus and exhibited the highest levels of mycorrhizal colonization, as well as the greatest stimulation of physiological parameters.
Seasonal and regional variations in kiwifruit storage quality imply a weather effect. This is perhaps mediated via fruit transpiration and fruit mineral nutrition. Concurrent measurements of fruit transpiration and weather are modelled to predict cumulative fruit transpiration throughout the season.
Background and aims
In most fruit crops, storage quality varies greatly between regions and seasons, causing significant commercial loss. Understanding the sources of this variability will contribute to the knowledge of fruit developmental physiology and may also benefit commercial fruit production via altered managements that reduce it or forecasts that predict it. A causal-chain relationship is proposed to help elucidate the sources of variability in fruit storage quality: the weather →(i)→ fruit transpiration →(ii)→ fruit calcium →(iii)→ fruit storage quality. This paper explores the first link of this hypothesis, →(i)→, for Hayward kiwifruit using field measurements of fruit transpiration rate and concurrent meteorological recordings. The aims are to identify the key environmental variables driving fruit transpiration and develop a predictive fruit transpiration model.
Fruit transpiration was determined hourly over several 24-h periods by recording weight loss of detached fruit, on Days 23, 35, 49, 65, 94 and 140 after full bloom. Meteorological records were made every 15 min throughout the season at an adjacent regional weather station. A model of fruit transpiration was developed in which the usual meteorological variables (radiation, temperature, windspeed and relative humidity) were incorporated in a Fick's Law transpiration flux equation.
Fruit transpiration rate (i.e. the molar flux density, mmol cm−2 h−1) varied diurnally and decreased during the season. The dominant fruit variable governing transpiration rate was skin conductance and the dominant environmental variables were relative humidity and temperature. Radiation and windspeed were not significantly influential.
The model provides a good fit to the fruit transpiration rate measurements regardless of the time of day/night or the stage of fruit development. The model allows reasonably accurate and continuous predictions of fruit transpiration rate throughout fruit development based on standard meteorological recordings. It also allows estimates of cumulative fruit transpiration throughout the season.
Effect of different photosynthetic photon flux densities (0, 500, 1000, 1500 and 2000 μmol m−2s−1), temperatures (20, 25, 30, 35 and 40 °C) and CO2 concentrations (250, 350, 450, 550, 650 and 750 μmol mol−1) on gas and water vapour exchange characteristics of Cannabis sativa L. were studied to determine the suitable and efficient environmental conditions for its indoor mass cultivation for pharmaceutical uses. The rate of photosynthesis (PN) and water use efficiency (WUE) of Cannabis sativa increased with photosynthetic photon flux densities (PPFD) at the lower temperatures (20–25 °C). At 30 °C, PN and WUE increased only up to 1500 μmol m−2s−1 PPFD and decreased at higher light levels. The maximum rate of photosynthesis (PN max) was observed at 30 °C and under 1500 μmol m−2s−1 PPFD. The rate of transpiration (E) responded positively to increased PPFD and temperature up to the highest levels tested (2000 μmol m−2s−1 and 40 °C). Similar to E, leaf stomatal conductance (gs) also increased with PPFD irrespective of temperature. However, gs increased with temperature up to 30 °C only. Temperature above 30 °C had an adverse effect on gs in this species. Overall, high temperature and high PPFD showed an adverse effect on PN and WUE. A continuous decrease in intercellular CO2 concentration (Ci) and therefore, in the ratio of intercellular CO2 to ambient CO2 concentration (Ci/Ca) was observed with the increase in temperature and PPFD. However, the decrease was less pronounced at light intensities above 1500 μmol m−2s−1. In view of these results, temperature and light optima for photosynthesis was concluded to be at 25–30 °C and ∼1500 μmol m−2s−1 respectively. Furthermore, plants were also exposed to different concentrations of CO2 (250, 350, 450, 550, 650 and 750 μmol mol−1) under optimum PPFD and temperature conditions to assess their photosynthetic response. Rate of photosynthesis, WUE and Ci decreased by 50 %, 53 % and 10 % respectively, and Ci/Ca, E and gs increased by 25 %, 7 % and 3 % respectively when measurements were made at 250 μmol mol-1 as compared to ambient CO2 (350 μmol mol−1) level. Elevated CO2 concentration (750 μmol mol−1) suppressed E and gs ∼ 29% and 42% respectively, and stimulated PN, WUE and Ci by 50 %, 111 % and 115 % respectively as compared to ambient CO2 concentration. The study reveals that this species can be efficiently cultivated in the range of 25 to 30 °C and ∼1500 μmol m−2s−1 PPFD. Furthermore, higher PN, WUE and nearly constant Ci/Ca ratio under elevated CO2 concentrations in C. sativa, reflects its potential for better survival, growth and productivity in drier and CO2 rich environment.
Cannabis sativa; Photosynthesis; Transpiration; Water use efficiency
In contrast to C3 photosynthesis, the response of C4 photosynthesis to water stress has been less-well studied in spite of the significant contribution of C4 plants to the global carbon budget and food security. The key feature of C4 photosynthesis is the operation of a CO2-concentrating mechanism in the leaves, which serves to saturate photosynthesis and suppress photorespiration in normal air. This article reviews the current state of understanding about the response of C4 photosynthesis to water stress, including the interaction with elevated CO2 concentration. Major gaps in our knowledge in this area are identified and further required research is suggested.
Evidence indicates that C4 photosynthesis is highly sensitive to water stress. With declining leaf water status, CO2 assimilation rate and stomatal conductance decrease rapidly and photosynthesis goes through three successive phases. The initial, mainly stomatal phase, may or may not be detected as a decline in assimilation rates depending on environmental conditions. This is because the CO2-concentrating mechanism is capable of saturating C4 photosynthesis under relatively low intercellular CO2 concentrations. In addition, photorespired CO2 is likely to be refixed before escaping the bundle sheath. This is followed by a mixed stomatal and non-stomatal phase and, finally, a mainly non-stomatal phase. The main non-stomatal factors include reduced activity of photosynthetic enzymes; inhibition of nitrate assimilation, induction of early senescence, and changes to the leaf anatomy and ultrastructure. Results from the literature about CO2 enrichment indicate that when C4 plants experience drought in their natural environment, elevated CO2 concentration alleviates the effect of water stress on plant productivity indirectly via improved soil moisture and plant water status as a result of decreased stomatal conductance and reduced leaf transpiration.
It is suggested that there is a limited capacity for photorespiration or the Mehler reaction to act as significant alternative electron sinks under water stress in C4 photosynthesis. This may explain why C4 photosynthesis is equally or even more sensitive to water stress than its C3 counterpart in spite of the greater capacity and water use efficiency of the C4 photosynthetic pathway.
C3 and C4 photosynthesis; stomatal and non-stomatal limitation; high CO2; water stress
Leaves within a canopy may experience rapid and extreme fluctuations in ambient conditions. A shaded leaf, for example, may become exposed to an order of magnitude increase in solar radiation within a few seconds, due to sunflecks or canopy motions. Considering typical time scales for stomatal adjustments, (2 to 60 minutes), the gap between these two time scales raised the question whether leaves rely on their hydraulic and thermal capacitances for passive protection from hydraulic failure or over-heating until stomata have adjusted. We employed a physically based model to systematically study effects of short-term fluctuations in irradiance on leaf temperatures and transpiration rates. Considering typical amplitudes and time scales of such fluctuations, the importance of leaf heat and water capacities for avoiding damaging leaf temperatures and hydraulic failure were investigated. The results suggest that common leaf heat capacities are not sufficient to protect a non-transpiring leaf from over-heating during sunflecks of several minutes duration whereas transpirative cooling provides effective protection. A comparison of the simulated time scales for heat damage in the absence of evaporative cooling with observed stomatal response times suggested that stomata must be already open before arrival of a sunfleck to avoid over-heating to critical leaf temperatures. This is consistent with measured stomatal conductances in shaded leaves and has implications for water use efficiency of deep canopy leaves and vulnerability to heat damage during drought. Our results also suggest that typical leaf water contents could sustain several minutes of evaporative cooling during a sunfleck without increasing the xylem water supply and thus risking embolism. We thus submit that shaded leaves rely on hydraulic capacitance and evaporative cooling to avoid over-heating and hydraulic failure during exposure to typical sunflecks, whereas thermal capacitance provides limited protection for very short sunflecks (tens of seconds).
Water is a key resource, and the plant water transport system sets limits on maximum growth and drought tolerance. When plants open their stomata to achieve a high stomatal conductance (gs) to capture CO2 for photosynthesis, water is lost by transpiration1,2. Water evaporating from the airspaces is replaced from cell walls, in turn drawing water from the xylem of leaf veins, in turn drawing from xylem in the stems and roots. As water is pulled through the system, it experiences hydraulic resistance, creating tension throughout the system and a low leaf water potential (Ψleaf). The leaf itself is a critical bottleneck in the whole plant system, accounting for on average 30% of the plant hydraulic resistance3. Leaf hydraulic conductance (Kleaf = 1/ leaf hydraulic resistance) is the ratio of the water flow rate to the water potential gradient across the leaf, and summarizes the behavior of a complex system: water moves through the petiole and through several orders of veins, exits into the bundle sheath and passes through or around mesophyll cells before evaporating into the airspace and being transpired from the stomata. Kleaf is of strong interest as an important physiological trait to compare species, quantifying the effectiveness of the leaf structure and physiology for water transport, and a key variable to investigate for its relationship to variation in structure (e.g., in leaf venation architecture) and its impacts on photosynthetic gas exchange. Further, Kleaf responds strongly to the internal and external leaf environment3. Kleaf can increase dramatically with irradiance apparently due to changes in the expression and activation of aquaporins, the proteins involved in water transport through membranes4, and Kleaf declines strongly during drought, due to cavitation and/or collapse of xylem conduits, and/or loss of permeability in the extra-xylem tissues due to mesophyll and bundle sheath cell shrinkage or aquaporin deactivation5-10. Because Kleaf can constrain gs and photosynthetic rate across species in well watered conditions and during drought, and thus limit whole-plant performance they may possibly determine species distributions especially as droughts increase in frequency and severity11-14.
We present a simple method for simultaneous determination of Kleaf and gs on excised leaves. A transpiring leaf is connected by its petiole to tubing running to a water source on a balance. The loss of water from the balance is recorded to calculate the flow rate through the leaf. When steady state transpiration (E, mmol • m-2 • s-1) is reached, gs is determined by dividing by vapor pressure deficit, and Kleaf by dividing by the water potential driving force determined using a pressure chamber (Kleaf= E /- Δψleaf, MPa)15.
This method can be used to assess Kleaf responses to different irradiances and the vulnerability of Kleaf to dehydration14,16,17.
Plant Biology; Issue 70; Molecular Biology; Physiology; Ecology; Biology; Botany; Leaf traits; hydraulics; stomata; transpiration; xylem; conductance; leaf hydraulic conductance; resistance; evaporative flux method; whole plant
Background and Aims
The influence of temperature on the timing of budbreak in woody perennials is well known, but its effect on subsequent shoot growth and architecture has received little attention because it is understood that growth is determined by current temperature. Seasonal shoot development of grapevines (Vitis vinifera) was evaluated following differences in temperature near budbreak while minimizing the effects of other microclimatic variables.
Dormant buds and emerging shoots of field-grown grapevines were heated above or cooled below the temperature of ambient buds from before budbreak until individual flowers were visible on inflorescences, at which stage the shoots had four to eight unfolded leaves. Multiple treatments were imposed randomly on individual plants and replicated across plants. Shoot growth and development were monitored during two growing seasons.
Higher bud temperatures advanced the date of budbreak and accelerated shoot growth and leaf area development. Differences were due to higher rates of shoot elongation, leaf appearance, leaf-area expansion and axillary-bud outgrowth. Although shoots arising from heated buds grew most vigorously, apical dominance in these shoots was reduced, as their axillary buds broke earlier and gave rise to more vigorous lateral shoots. In contrast, axillary-bud outgrowth was minimal on the slow-growing shoots emerging from buds cooled below ambient. Variation in shoot development persisted or increased during the growing season, well after temperature treatments were terminated and despite an imposed soil water deficit.
The data indicate that bud-level differences in budbreak temperature may lead to marked differences in shoot growth, shoot architecture and leaf-area development that are maintained or amplified during the growing season. Although growth rates commonly are understood to reflect current temperatures, these results demonstrate a persistent effect of early-season temperatures, which should be considered in future growth models.
Apical dominance; budbreak; grapevine; growth; leaf expansion; shoot elongation; temperature; Vitis vinifera
The common reed (Phragmites australis) is a clonal wetland grass with high genetic variability. Clone-specific differences are reflected in morphological and physiological traits, and hence in the ability to cope with environmental stress. The responses to progressively increasing salinity of fifteen distinct Phragmites australis clones reveal genotype-related strategies of salt avoidance and exclusion. The salinity-induced inhibition in shoot elongation rate and photosynthesis varies widely between clones. The differences can be partially attributed to their geographic range, but not correlated to ploidy level. Thus, the genetic background is a major factor influencing the salinity tolerance of distinct Phragmites australis clones.
Different clones of the wetland grass Phragmites australis differ in their morphology and physiology, and hence in their ability to cope with environmental stress. We analysed the responses of 15 P. australis clones with distinct ploidy levels (PLs) (4n, 6n, 8n, 10n, 12n) and geographic origins (Romania, Russia, Japan, Czech Republic, Australia) to step-wise increased salinity (8, 16, 24, 32, 40, 56 and 72 ppt). Shoot elongation rate, photosynthesis and plant part-specific ion accumulation were studied in order to assess if traits associated with salinity tolerance can be related to the genetic background and the geographic origin of the clones. Salt stress affected all clones, but at different rates. The maximum height was reduced from 1860 mm in control plants to 660 mm at 40 ppt salinity. The shoot elongation rate of salt-exposed plants varied significantly between clones until 40 ppt salinity. The light-saturated photosynthesis rate (Pmax) was stimulated by a salinity of 8 ppt, but decreased significantly at higher salinities. The stomatal conductance (gs) and the transpiration rate (E) decreased with increasing salinity. Only three clones survived at 72 ppt salinity, although their rates of photosynthesis were strongly inhibited. The roots and basal leaves of the salt-exposed plants accumulated high concentrations of water-extractable Na+ (1646 and 1004 µmol g−1 dry mass (DM), respectively) and Cl− (1876 and 1400 µmol g−1 DM, respectively). The concentrations of water-extractable Mg2+ and Ca2+ were reduced in salt-exposed plants compared with controls. The variation of all the measured parameters was higher among clones than among PLs. We conclude that the salinity tolerance of distinct P. australis clones varies widely and can be partially attributed to their longitudinal geographic origin, but not to PL. Further investigation will help in improving the understanding of this species' salt tolerance mechanisms and their connection to genetic factors.
Common reed; ecophysiology; geographic range; ion concentration; ploidy level; salt-stress tolerance
The theoretical basis for the link between the leaf exchange of carbonyl sulfide (COS), carbon dioxide (CO2) and water vapour (H2O) and the assumptions that need to be made in order to use COS as a tracer for canopy net photosynthesis, transpiration and stomatal conductance, are reviewed. The ratios of COS to CO2 and H2O deposition velocities used to this end are shown to vary with the ratio of the internal to ambient CO2 and H2O mole fractions and the relative limitations by boundary layer, stomatal and internal conductance for COS. It is suggested that these deposition velocity ratios exhibit considerable variability, a finding that challenges current parameterizations, which treat these as vegetation-specific constants. COS is shown to represent a better tracer for CO2 than H2O. Using COS as a tracer for stomatal conductance is hampered by our present poor understanding of the leaf internal conductance to COS. Estimating canopy level CO2 and H2O fluxes requires disentangling leaf COS exchange from other ecosystem sources/sinks of COS. We conclude that future priorities for COS research should be to improve the quantitative understanding of the variability in the ratios of COS to CO2 and H2O deposition velocities and the controlling factors, and to develop operational methods for disentangling ecosystem COS exchange into contributions by leaves and other sources/sinks. To this end, integrated studies, which concurrently quantify the ecosystem-scale CO2, H2O and COS exchange and the corresponding component fluxes, are urgently needed.
We investigate the potential of carbonyl sulfide (COS) for being used as a tracer for canopy net photosynthesis, transpiration and stomatal conductance by examining the theoretical basis of the link between leaf COS, carbon dioxide (CO2) and water vapour (H2O) exchange. Our analysis identifies several limitations that need to be overcome to this end, however at present we lack appropriate ecosystem-scale field measurements for assessing their practical significance. It however appears that COS represents a better tracer for CO2 than H2O. Concurrent measurements of ecosystem scale COS, CO2 and H2O exchange are advocated.
carbon dioxide; gross photosynthesis; internal conductance; net photosynthesis; water vapour
Effects of water deficit on plant water status, gas exchange and hydraulic conductance were investigated in Betula pendula under artificially manipulated air humidity in Eastern Estonia. The study was aimed to broaden an understanding of the ability of trees to acclimate with the increasing atmospheric humidity predicted for northern Europe. Rapidly-induced water deficit was imposed by dehydrating cut branches in open-air conditions; long-term water deficit was generated by seasonal drought.
The rapid water deficit quantified by leaf (ΨL) and branch water potentials (ΨB) had a significant (P < 0.001) effect on gas exchange parameters, while inclusion of ΨB in models resulted in a considerably better fit than those including ΨL, which supports the idea that stomatal openness is regulated to prevent stem rather than leaf xylem dysfunction. Under moderate water deficit (ΨL≥-1.55 MPa), leaf conductance to water vapour (gL), transpiration rate and leaf hydraulic conductance (KL) were higher (P < 0.05) and leaf temperature lower in trees grown in elevated air humidity (H treatment) than in control trees (C treatment). Under severe water deficit (ΨL<-1.55 MPa), the treatments showed no difference. The humidification manipulation influenced most of the studied characteristics, while the effect was to a great extent realized through changes in soil water availability, i.e. due to higher soil water potential in H treatment. Two functional characteristics (gL, KL) exhibited higher (P < 0.05) sensitivity to water deficit in trees grown under increased air humidity.
The experiment supported the hypothesis that physiological traits in trees acclimated to higher air humidity exhibit higher sensitivity to rapid water deficit with respect to two characteristics - leaf conductance to water vapour and leaf hydraulic conductance. Disproportionate changes in sensitivity of stomatal versus leaf hydraulic conductance to water deficit will impose greater risk of desiccation-induced hydraulic dysfunction on the plants, grown under high atmospheric humidity, in case of sudden weather fluctuations, and might represent a potential threat in hemiboreal forest ecosystems. There is no trade-off between plant hydraulic capacity and photosynthetic water-use efficiency on short time scale.
Betula pendula; Branch water potential; Climate change; Hydraulic conductance; Leaf water potential; Net photosynthesis; Silver birch; Stomatal conductance; Water-use efficiency
Photosynthesis is fundamental to biomass production, but sensitive to drought. To understand the genetics of leaf photosynthesis, especially under drought, upland rice cv. Haogelao, lowland rice cv. Shennong265, and 94 of their introgression lines (ILs) were studied at flowering and grain filling under drought and well-watered field conditions. Gas exchange and chlorophyll fluorescence measurements were conducted to evaluate eight photosynthetic traits. Since these traits are very sensitive to fluctuations in microclimate during measurements under field conditions, observations were adjusted for microclimatic differences through both a statistical covariant model and a physiological approach. Both approaches identified leaf-to-air vapour pressure difference as the variable influencing the traits most. Using the simple sequence repeat (SSR) linkage map for the IL population, 1–3 quantitative trait loci (QTLs) were detected per trait–stage–treatment combination, which explained between 7.0% and 30.4% of the phenotypic variance of each trait. The clustered QTLs near marker RM410 (the interval from 57.3 cM to 68.4 cM on chromosome 9) were consistent over both development stages and both drought and well-watered conditions. This QTL consistency was verified by a greenhouse experiment under a controlled environment. The alleles from the upland rice at this interval had positive effects on net photosynthetic rate, stomatal conductance, transpiration rate, quantum yield of photosystem II (PSII), and the maximum efficiency of light-adapted open PSII. However, the allele of another main QTL from upland rice was associated with increased drought sensitivity of photosynthesis. These results could potentially be used in breeding programmes through marker-assisted selection to improve drought tolerance and photosynthesis simultaneously.
Chlorophyll fluorescence; drought; gas exchange; photosynthesis; physiological model; quantitative trait locus (QTL)
The dual-source Shuttleworth-Wallace model has been widely used to estimate and partition crop evapotranspiration (λET). Canopy stomatal conductance (Gsc), an essential parameter of the model, is often calculated by scaling up leaf stomatal conductance, considering the canopy as one single leaf in a so-called “big-leaf” model. However, Gsc can be overestimated or underestimated depending on leaf area index level in the big-leaf model, due to a non-linear stomatal response to light. A dual-leaf model, scaling up Gsc from leaf to canopy, was developed in this study. The non-linear stomata-light relationship was incorporated by dividing the canopy into sunlit and shaded fractions and calculating each fraction separately according to absorbed irradiances. The model includes: (1) the absorbed irradiance, determined by separately integrating the sunlit and shaded leaves with consideration of both beam and diffuse radiation; (2) leaf area for the sunlit and shaded fractions; and (3) a leaf conductance model that accounts for the response of stomata to PAR, vapor pressure deficit and available soil water. In contrast to the significant errors of Gsc in the big-leaf model, the predicted Gsc using the dual-leaf model had a high degree of data-model agreement; the slope of the linear regression between daytime predictions and measurements was 1.01 (R2 = 0.98), with RMSE of 0.6120 mm s−1 for four clear-sky days in different growth stages. The estimates of half-hourly λET using the dual-source dual-leaf model (DSDL) agreed well with measurements and the error was within 5% during two growing seasons of maize with differing hydrometeorological and management strategies. Moreover, the estimates of soil evaporation using the DSDL model closely matched actual measurements. Our results indicate that the DSDL model can produce more accurate estimation of Gsc and λET, compared to the big-leaf model, and thus is an effective alternative approach for estimating and partitioning λET.
Background and Aims
Rice (Oryza sativa) plants lose significant amounts of volatile NH3 from their leaves, but it has not been shown that this is a consequence of photorespiration. Involvement of photorespiration in NH3 emission and the role of glutamine synthetase (GS) on NH3 recycling were investigated using two rice cultivars with different GS activities.
NH3 emission (AER), and gross photosynthesis (PG), transpiration (Tr) and stomatal conductance (gS) were measured on leaves of ‘Akenohoshi’, a cultivar with high GS activity, and ‘Kasalath’, a cultivar with low GS activity, under different light intensities (200, 500 and 1000 µmol m−2 s−1), leaf temperatures (27·5, 32·5 and 37·5 °C) and atmospheric O2 concentrations ([O2]: 2, 21 and 40 %, corresponding to 20, 210 and 400 mmol mol−1).
An increase in [O2] increased AER in the two cultivars, accompanied by a decrease in PG due to enhanced photorespiration, but did not greatly influence Tr and gS. There were significant positive correlations between AER and photorespiration in both cultivars. Increasing light intensity increased AER, PG, Tr and gS in both cultivars, whereas increasing leaf temperature increased AER and Tr but slightly decreased PG and gS. ‘Kasalath’ (low GS activity) showed higher AER than ‘Akenohoshi’ (high GS activity) at high light intensity, leaf temperature and [O2].
Our results demonstrate that photorespiration is strongly involved in NH3 emission by rice leaves and suggest that differences in AER between cultivars result from their different GS activities, which would result in different capacities for reassimilation of photorespiratory NH3. The results also suggest that NH3 emission in rice leaves is not directly controlled by transpiration and stomatal conductance.
Ammonia assimilation; ammonia emission; glutamine synthetase; nitrogen; Oryza sativa; photorespiration; rice cultivars