Tight Stomatal Control Research Articles

Long‐term transpiration change with rainfall decline in a Mediterranean Quercus ilex forest

AbstractIn the Mediterranean basin, precipitation is expected to decline as a consequence of climate change. The response of a Quercus ilex forest in southern France to such a decline in water availability was studied using a 4‐year throughfall exclusion experiment. Seasonal courses of sap flow and leaf water potential were obtained from 2004 to 2007 and used to characterize tree water relations in a control and a dry treatment. The experiment reduced the average precipitation input to the soil by 29%, and resulted in a 23% reduction in annual transpiration. Soil water potential was significantly lower in the dry treatment only during summer drought, but transpiration was reduced all year round even during well‐watered periods. Despite a tight stomatal control over transpiration, whole‐tree hydraulic conductance was found to be lower in the trees growing in the driest conditions. This reduction in water transport capacity was observed jointly with a reduction in leaf transpiring area. Canopy leaf area decreased by 18% in the dry treatment as a consequence of the throughfall exclusion, which was found to validate the ecohydrological equilibrium theory.

Assessing environmental controls over conductances through the soil–plant–atmosphere continuum in an experimental olive tree plantation of southern Italy

Constraints on plant water status and water use indicators were studied in orchard grown Olea europaea L. trees, cv. Nocellara del Belice, in order to assess productivity and drought adaptation. The study was conducted on 16-year-old grafted plants, grown under different water treatments for 14 years in an experimental site plantation in southern Italy. Water treatments were: a non-irrigated, rainfed, control (T0) and three treatments that received seasonal water amount equivalent to 33 and 66% of crop evapotranspiration ( ET C) from the beginning of pit hardening to early fruit veraison (respectively T33 and T66), and 100% of ET C throughout the irrigation season (T100). During 2006 and 2007 growing seasons, plants were continuously monitored by automatic point dendrometers measuring stem radius variation and whole-plant water use was determined using a xylem sap flow method (compensation heat-pulse technique). Additional ecophysiological parameters, such as stomatal conductance and water potentials were periodically measured, as well as vegetative development. Predawn leaf water potential decreased in rainfed trees and this was associated with increasing soil moisture deficit, while the difference between predawn and midday leaf water potentials increased, suggesting anisohydric regulation of plant water potential. Olive trees exhibited a tight stomatal control over transpiration, but insufficient to prevent loss of hydraulic conductance under severe drought stress. The stem radial increment and sap flux did not differ consistently between water treatments in the mild year (2006), while irrigated trees had higher water use than rainfed trees in the dry year (2007). However, plants growing under rainfed conditions showed a small increase in maximum daily shrinkage during drought periods compared to irrigated ones, although no marked differences were recorded between irrigated treatments. The mean daily transpiration rate, canopy conductance and decoupling coefficient were higher in 2006 than in 2007, the driest year, only, and in fully irrigated rather than rainfed plants. These results support the idea that rainfed trees had more conservative water use strategies than irrigated trees, and that deficit-irrigated trees acclimated somewhat functionally and structurally to long-term partial watering. Based on these observations, we can argue that loss of hydraulic conductance is an important mechanism for increasing stomatal control of transpiration under progressive soil drying. Another view implies that the increasing difference between soil and leaf water potential during summer induced stomatal closure and minimized the risk of a collapse of the conductive system, decreasing transpiration and reducing hydraulic conductance.

Physiological performance of three pine species provides evidence for gap partitioning

Gradients of light and moisture availability peak at different positions within canopy gaps in northern latitudes providing the opportunity for niche partitioning in and around gaps based on differences in individual species’ life history attributes. This gap partitioning offers potential for increasing diversity in forests impacted by gap-creating disturbances. We examined resource availability and the physiological performance of three Pinus species with varying tolerances for shade and moisture stress across large (0.3 ha) canopy gaps to investigate relationships between gap position and species performance. Light availability was lowest in southern gap edges, while water availability was lowest in northern edges, and higher at gap interior positions than edges. Pinus banksiana seedlings had higher light-saturated CO 2 assimilation rates than P. resinosa or P. strobus seedlings at interior gap positions, and outperformed P. strobus at northern gap edges, but there were no differences between species at southern edges. Both transpiration and stomatal conductance were greatest for P. banksiana in gap centers, but showed few differences between species at edges. Foliar nitrogen concentrations were highest for P. banksiana, suggesting the dominance of this species in central gap locations may be due to a combination of high photosynthetic capacity and tight stomatal control to regulate moisture stress at drier gap positions. Our results suggest P. banksiana seedlings may be competitively superior in gap positions with high light and moisture availability, but P. resinosa and P. strobus become competitive under the drier conditions and moderate shade near gap edges. These findings support the concept of gap partitioning, and suggest silvicultural systems that incorporate patch cuttings could be used to promote diverse regeneration in northern pine forests.

Physiological characteristics of cassava tolerance to prolonged drought in the tropics: implications for breeding cultivars adapted to seasonally dry and semiarid environments

The paper summarizes research conducted at International Center for Tropical Agriculture (CIAT) on responses of cassava to extended water shortages in the field aided by modern gas-exchange and water-relation techniques as well as biochemical assays. The aim of the research was to coordinate basic and applied aspects of crop physiology into a breeding strategy with a multidisciplinary approach. Several physiological characteristics/traits and mechanisms underpinning tolerance of cassava to drought were elucidated using a large number of genotypes from the CIAT core germplasm collection grown in various locations representing ecozones where cassava is cultivated. Most notable among these characteristics are the high photosynthetic capacity of cassava leaves in favorable environments and the maintenance of reasonable rates throughout prolonged water deficits, a crucial characteristic for high and sustainable productivity. Cassava possess a tight stomatal control over leaf gas exchange that reduces water losses when plants are subjected to soil water deficits as well as to high atmospheric evaporative demands, thus protecting leaves from severe dehydration. During prolonged water deficits, cassava reduces its canopy by shedding older leaves and forming smaller new leaves leading to less light interception, another adaptive trait to drought. Though root yield is reduced (but much less than the reduction in top growth) under water stress, the crop can recover when water becomes available by rapidly forming new canopy leaves with much higher photosynthetic rates compared to unstressed crops, thus compensating for yield losses with final yields approaching those in well-watered crops. Cassava can extract slowly water from deep soils, a characteristic of paramount importance in seasonally dry and semiarid environments where deeply stored water needs to be tapped. Screening large accessions under seasonally dry and semiarid environments showed that yield is significantly correlated with upper canopy leaf photosynthetic rates, and the association was attributed mainly to nonstomatal (anatomical/biochemical) factors. Parental materials with both high yields and photosynthetic rates were identified for incorporation into breeding and selection programs for cultivars adapted to prolonged drought coupled with high temperatures and dry air, conditions that might be further aggravated by global climate changes in tropical regions.

Water relations of baobab trees (Adansonia spp. L.) during the rainy season: does stem water buffer daily water deficits?

Baobab trees are often cited in the literature as water-storing trees, yet few studies have examined this assumption. We assessed the role of stored water in buffering daily water deficits in two species of baobabs (Adansonia rubrostipa Jum. and H. Perrier and Adansonia za Baill.) in a tropical dry forest in Madagascar. We found no lag in the daily onset of sap flow between the base and the crown of the tree. Some night-time sap flow occurred, but this was more consistent with a pattern of seasonal stem water replenishment than with diurnal usage. Intrinsic capacitance of both leaf and stem tissue (0.07-0.08 and 1.1-1.43 MPa(-1), respectively) was high, yet the amount of water that could be withdrawn before turgor loss was small because midday leaf and stem water potentials (WPs) were near the turgor-loss points. Stomatal conductance was high in the daytime but then declined rapidly, suggesting an embolism-avoidance strategy. Although the xylem of distal branches was relatively vulnerable to cavitation (P50: 1.1-1.7 MPa), tight stomatal control and minimum WPs near--1.0 MPa maintained native embolism levels at 30-65%. Stem morphology and anatomy restrict water movement between storage tissues and the conductive pathway, making stored-water usage more appropriate to longer-term water deficits than as a buffer against daily water deficits.

Ozone uptake, water loss and carbon exchange dynamics in annually drought-stressed Pinus ponderosa forests: measured trends and parameters for uptake modeling.

This paper describes 3 years of physiological measurements on ponderosa pine (Pinus ponderosa Dougl. ex Laws.) growing along an ozone concentration gradient in the Sierra Nevada, California, including variables necessary to parameterize, validate and modify photosynthesis and stomatal conductance algorithms used to estimate ozone uptake. At all sites, gas exchange was under tight stomatal control during the growing season. Stomatal conductance was strongly correlated with leaf water potential (R2=0.82), which decreased over the growing season with decreasing soil water content (R2=0.60). Ozone uptake, carbon uptake, and transpirational water loss closely followed the dynamics of stomatal conductance. Peak ozone and CO2 uptake occurred in early summer and declined progressively thereafter. As a result, periods of maximum ozone uptake did not correspond to periods of peak ozone concentration, underscoring the inappropriateness of using current metrics based on concentration (e.g., SUM0, W126 and AOT40) for assessing ozone exposure risk to plants in this climate region. Both Jmax (maximum CO2-saturated photosynthetic rate, limited by electron transport) and Vcmax (maximum rate of Rubisco-limited carboxylation) increased toward the middle of the growing season, then decreased in September. Intrinsic water-use efficiency rose with increasing drought stress, as expected. The ratio of Jmax to Vcmax was similar to literature values of 2.0. Nighttime respiration followed a Q10 of 2.0, but was significantly higher at the high-ozone site. Respiration rates decreased by the end of the summer as a result of decreased metabolic activity and carbon stores.

Changes in whole-tree water relations during ontogeny of Pinus flexilis and Pinus ponderosa in a high-elevation meadow

We measured sap flux in Pinus ponderosa Laws. and Pinus flexilis James trees in a high-elevation meadow in northern Arizona that has been invaded by conifers over the last 150 years. Sap flux and environmental data were collected from July 1 to September 1, 2000, and used to estimate leaf specific transpiration rate (El), canopy conductance (Gc) and whole-plant hydraulic conductance (Kh). Leaf area to sapwood area ratio (LA/SA) increased with increasing tree size in P. flexilis, but decreased with increasing tree size in P. ponderosa. Both Gc and Kh decreased with increasing tree size in P. flexilis, and showed no clear trends with tree size in P. ponderosa. For both species, Gc was lower in the summer dry season than in the summer rainy season, but El did not change between wet and dry summer seasons. Midday water potential (Psi(mid)) did not change across seasons for either species, whereas predawn water potential (Psi(pre)) tracked variation in soil water content across seasons. Pinus flexilis showed greater stomatal response to vapor pressure deficit (VPD) and maintained higher Psi(mid) than P. ponderosa. Both species showed greater sensitivity to VPD at high photosynthetically active radiation (PAR; > 2500 micromol m-2 s-1) than at low PAR (< 2500 micromol m-2 s-1). We conclude that the direction of change in Gc and Kh with increasing tree size differed between co-occurring Pinus species, and was influenced by changes in LA/SA. Whole-tree water use and El were similar between wet and dry summer seasons, possibly because of tight stomatal control over water loss.

Evaluation of land‐surface interaction in ECMWF and NCEP/NCAR reanalysis models over grassland (FIFE) and boreal forest (BOREAS)

The National Center for Environmental Prediction and National Center for Atmospheric Research (NCEP/NCAR) and European Centre for Medium‐Range Weather Forecasts (ECMWF) reanalysis models are compared with First ISLSCP (International Satellite Land Surface Climatology Project) Field Experiment (FIFE) grassland data from Kansas in 1987 and Boreal Ecosystem‐Atmosphere Study (BOREAS) data from an old black spruce site in 1996 near Thompson, Manitoba. Some aspects of the comparison are similar for the two ecosystems. Over grassland and after snowmelt in the boreal forest, both models represent the seasonal cycle of near‐surface temperature well. The two models have quite different soil hydrology components. The ECMWF model includes soil water nudging based on low level humidity errors. While this works quite well for the FIFE grassland, it appears to give too high evaporation over the boreal forest. The NCEP/NCAR model constrains long‐term drifts by nudging deep soil water toward climatology. Over the FIFE site, this seems to give too low evaporation in midsummer, while at the BOREAS site, evaporation in this model is high. Both models have some difficulty representing the surface diurnal cycle of humidity. In the NCEP/NCAR reanalysis this leads to errors primarily in June, when the surface boundary layer stays saturated and too much precipitation occurs. In the ECMWF reanalysis there is a morning peak of mixing ratio, which an earlier work showed resulted from too shallow a boundary layer in the morning. Over the northern boreal forest there are important physical processes, which are not represented in either reanalysis model. In particular very high model albedos in spring, when there is snow under the forest canopy, lead to a very low daytime net radiation. This in turn leads to a large underestimate of the daytime surface fluxes, particularly the sensible heat flux, and to daytime model surface temperatures that are as much as 15 K low. In addition, the models do not account for the reduction in evaporation associated with frozen soil, and they generally have too large evapotranspiration in June and July, probably because they do not model the tight stomatal control of the coniferous forest.

Photosynthetic gas exchange and water use in tropical and subtropical populations of the mangrove Aegiceras corniculatum

Photosynthetic gas exchange and stomatal behaviour in the tropical (Darwin: 12°25′S) and subtropical (Ballina: 28°50′S) populations of the mangrove Aegiceras corniculatum (L.) Blanco were compared at elevated air temperatures (&gt;32°C) and leaf-to-air vapour pressure deficit (vpd &gt;25 mbar) with a quantum flux at, or above, their light saturation capacity (&gt;600 µmol m-2 s-1). At the lower end of the tested aridity range, the tropical population showed a less conservative water use than the subtropical population. As aridity increased, both populations showed a reduction in stomatal conductance. However, transpiration rates remained higher in the subtropical population at all times, reducing its water-use efficiency from that under less arid conditions. At extreme aridity (temperature &gt;37°C and vpd &gt;35 mbar), the efficiency of the evaporative cooling of fully exposed leaves was evident in the leaf-to-air temperature differential which remained minimal in the tropical population while it increased significantly in the subtropical population. Aridity tolerance was more pronounced in individuals from the tropical site than from the subtropical site, as evidenced by a tighter stomatal control on water use in the tropical population. These data suggest that the tropical and subtropical populations of A. corniculatum have different physiological responses to aridity.

Photosynthesis and Transpiration as a Function of Gaseous Diffusive Resistances in Orange Leaves

AbstractThe CO2 and H2O vapour exchange of single attached orange, Citrus sinensis (L.), leaves was measured under laboratory conditions using infrared gas analysis. Gaseous diffusive resistances were derived from measurements at a saturating irradiance and at a leaf temperature optimum for photosynthesis. Variation in leaf resistance (within the range 1.6 to 60 s cm‐1) induced by moisture status, or by cyclic oscillations in stomatal aperture, was associated with changes in both photosynthesis and transpiration.At low leaf resistance (ri less than 10 s cm‐1) the ratio of transpiration to photosynthesis declined with reduced stomatal aperture, indicating a tighter stomatal control over H2O vapour loss than over CO2 assimilation. At higher leaf resistance (ri greater than 10 s cm‐1) changes in transpiration and photosynthesis were linearly related, but leaf resistance and mesophyll resistance were also positively correlated, so that strictly stomatal control of photosynthesis became more apparent than real.This evidence, combined with direct measurements of CO2 diffusive resistances (in a ‐O2 gas stream) emphasised the presence of a significant mesophyll resistance; i.e., an additional and rate limiting resistance to CO2 assimilation over and above that encountered by H2O vapour escaping from the leaf.