Induction of stomatal opening following a night-chilling event alleviates physiological damage in mango trees.

Chilling events have become more frequent with climate change and are a significant abiotic factor causing physiological damage to plants and, consequently, reducing crop yield. Like other tropical and subtropical plants, mango (Mangifera indica L.) is particularly sensitive to chilling events, especially if they are followed by bright sunny days. It was previously shown that in mango leaves stomatal opening is restricted in the morning following a night-chilling event. This impairment results in restraint of carbon assimilation and subsequently, photoinhibition and reactive oxygen species production, which leads to chlorosis and in severe cases, cell death. Our detailed physiological analysis showed that foliar application of the guard cell H+-ATPase activator, fusicoccin, in the morning after a cold night, mitigates the physiological damage from 'cold night-bright day' abiotic stress. This application restored stomatal opening, thereby enabling gas exchange, releasing the photosynthetic machinery from harmful excess photon energy, and improving the plant's overall physiological state. The mechanisms by which plants react to this abiotic stress are examined in this work. The foliar application of compounds that cause stomatal opening as a potential method of minimizing physiological damage due to night chilling is discussed.

A role for ethylene signaling and biosynthesis in regulating and accelerating CO2 - and abscisic acid-mediated stomatal movements in Arabidopsis.

Little is known about long-distance mesophyll-driven signals that regulate stomatal conductance. Soluble and/or vapor-phase molecules have been proposed. In this study, the involvement of the gaseous signal ethylene in the modulation of stomatal conductance in Arabidopsis thaliana by CO2 /abscisic acid (ABA) was examined. We present a diffusion model which indicates that gaseous signaling molecule/s with a shorter/direct diffusion pathway to guard cells are more probable for rapid mesophyll-dependent stomatal conductance changes. We, therefore, analyzed different Arabidopsis ethylene-signaling and biosynthesis mutants for their ethylene production and kinetics of stomatal responses to ABA/[CO2 ]-shifts. According to our research, higher [CO2 ] causes Arabidopsis rosettes to produce more ethylene. An ACC-synthase octuple mutant with reduced ethylene biosynthesis exhibits dysfunctional CO2 -induced stomatal movements. Ethylene-insensitive receptor (gain-of-function), etr1-1 and etr2-1, and signaling, ein2-5 and ein2-1, mutants showed intact stomatal responses to [CO2 ]-shifts, whereas loss-of-function ethylene receptor mutants, including etr2-3;ein4-4;ers2-3, etr1-6;etr2-3 and etr1-6, showed markedly accelerated stomatal responses to [CO2 ]-shifts. Further investigation revealed a significantly impaired stomatal closure to ABA in the ACC-synthase octuple mutant and accelerated stomatal responses in the etr1-6;etr2-3, and etr1-6, but not in the etr2-3;ein4-4;ers2-3 mutants. These findings suggest essential functions of ethylene biosynthesis and signaling components in tuning/accelerating stomatal conductance responses to CO2 and ABA.

Physiological characterization of the wild almond Prunus arabica stem photosynthetic capability.

Leaves are the major plant tissue for transpiration and carbon fixation in deciduous trees. In harsh habitats, atmospheric CO2 assimilation via stem photosynthesis is common, providing extra carbon gain to cope with the detrimental conditions. We studied two almond species, the commercial Prunus dulcis cultivar "Um-el-Fahem" and the rare wild Prunus arabica. Our study revealed two distinctive strategies for carbon gain in these almond species. While, in P. dulcis, leaves possess the major photosynthetic surface area, in P. arabica, green stems perform this function, in particular during the winter after leaf drop. These two species' anatomical and physiological comparisons show that P. arabica carries unique features that support stem gas exchange and high-gross photosynthetic rates via stem photosynthetic capabilities (SPC). On the other hand, P. dulcis stems contribute low gross photosynthesis levels, as they are designed solely for reassimilation of CO2 from respiration, which is termed stem recycling photosynthesis (SRP). Results show that (a) P. arabica stems are covered with a high density of sunken stomata, in contrast to the stomata on P. dulcis stems, which disappear under a thick peridermal (bark) layer by their second year of development. (b) P. arabica stems contain significantly higher levels of chlorophyll compartmentalized to a mesophyll-like, chloroplast-rich, parenchyma layer, in contrast to rounded-shape cells of P. dulcis's stem parenchyma. (c) Pulse amplitude-modulated (PAM) fluorometry of P. arabica and P. dulcis stems revealed differences in the chlorophyll fluorescence and quenching parameters between the two species. (d) Gas exchange analysis showed that guard cells of P. arabica stems tightly regulate water loss under elevated temperatures while maintaining constant and high assimilation rates throughout the stem. Our data show that P. arabica uses a distinctive strategy for tree carbon gain via stem photosynthetic capability, which is regulated efficiently under harsh environmental conditions, such as elevated temperatures. These findings are highly important and can be used to develop new almond cultivars with agriculturally essential traits.

Physiological Characterization of Young 'Hass' Avocado Plant Leaves Following Exposure to High Temperatures and Low Light Intensity.

The worldwide demand for avocados has resulted in the planting of millions of young plants each year. However, global warming, resulting in high temperatures, sensed as heat stress, may severely damage these new plantings. The objective of this study was to assess the risks of heat stress on young avocado plants. We aimed to characterize different physiological parameters of young 'Hass' plant leaves following exposure to high temperatures under low light (LL) intensity and to pinpoint the temperature threshold for significant heat stress damage in these plants. To this end, young potted plants were subjected to different temperature gradients in a controlled-climate chamber. Minor and severe leaf damage was apparent in plants subjected to the 51 °C and 53 °C treatments, respectively. Minor and vast reductions in optimal quantum yield efficiency of photosystem II (Fv/Fm) values were observed in plants subjected to 51 °C and 53 °C, respectively. Heat stress treatments significantly reduced CO2 assimilation in plants subjected to 49 °C and higher temperatures. Stomatal conductance to water vapour and substomatal internal CO2 concentration were less sensitive to the heat treatments. These results imply that the heat damage threshold for young avocado plants under LL conditions is between 49 °C and 51 °C, whereas at 53 °C, severe and irreversible leaf damage occurs.

The effect of blue light on stomatal oscillations and leaf turgor pressure in banana leaves.

Stomatal oscillations are cyclic opening and closing of stomata, presumed to initiate from hydraulic mismatch between leaf water supply and transpiration rate. To test this assumption, mismatches between water supply and transpiration were induced using manipulations of vapour pressure deficit (VPD) and light spectrum in banana (Musa acuminata). Simultaneous measurements of gas exchange with changes in leaf turgor pressure were used to describe the hydraulic mismatches. An increase of VPD above a certain threshold caused stomatal oscillations with variable amplitudes. Oscillations in leaf turgor pressure were synchronized with stomatal oscillations and balanced only when transpiration equaled water supply. Surprisingly, changing the light spectrum from red and blue to red alone at constant VPD also induced stomatal oscillations - while the addition of blue (10%) to red light only ended oscillations. Blue light is known to induce stomatal opening and thus should increase the hydraulic mismatch, reduce the VPD threshold for oscillations and increase the oscillation amplitude. Unexpectedly, blue light reduced oscillation amplitude, increased VPD threshold and reduced turgor pressure loss. These results suggest that additionally, to the known effect of blue light on the hydroactive opening response of stomata, it can also effect stomatal movement by increased xylem-epidermis water supply.

Lower leaf gas-exchange and higher photorespiration of treated wastewater irrigated Citrus trees is modulated by soil type and climate.

Water quality, soil and climate can interact to limit photosynthesis and to increase photooxidative damage in sensitive plants. This research compared diffusive and non-diffusive limitations to photosynthesis as well as photorespiration of leaves of grapefruit trees in heavy clay and sandy soils having a previous history of treated wastewater (TWW) irrigation for >10 years, with different water qualities [fresh water (FW) vs TWW and sodium amended treated wastewater (TWW + Na)] in two arid climates (summer vs winter) and in orchard and lysimeter experiments. TWW irrigation increased salts (Na(+) and Cl(-) ), membrane leakage, proline and soluble sugar content, and decreased osmotic potentials in leaves of all experiments. Reduced leaf growth and higher stomatal and non-stomatal (i.e. mesophyll) limitations were found in summer and on clay soil for TWW and TWW + Na treatments in comparison to winter, sandy soil and FW irrigation, respectively. Stomatal closure, lower chlorophyll content and altered Rubisco activity are probable causes of higher limitations. On the other hand, non-photochemical quenching, an alternative energy dissipation pathway, was only influenced by water quality, independent of soil type and season. Furthermore, light and CO2 response curves were investigated for other possible causes of higher non-stomatal limitation. A higher proportion of non-cyclic electrons were directed to the O2 dependent pathway, and a higher proportion of electrons were diverted to photorespiration in summer than in winter. In conclusion, both diffusive and non-diffusive limitations contribute to the lower photosynthetic performance of leaves following TWW irrigation, and the response depends on soil type and environmental factors.

Modification of non-stomatal limitation and photoprotection due to K and Na nutrition of olive trees.

Potassium (K) is an essential macronutrient shown to play a fundamental role in photosynthetic processes and may facilitate photoinhibition resistance. In some plant species, sodium (Na) can partially substitute for K. Although photosynthetic enhancement has been well established, the mechanisms by which K or Na affects photosynthesis are not fully understood. Olive (Olea europaea L.) trees were previously shown to benefit from Na nutrition when K is limiting. In order to study the effect of K and Na on photosynthetic performance, we measured gas exchange and chlorophyll fluorescence in young olive trees supplied with either K, Na or no fertilizer, and subjected to manipulated levels of CO2, O2 and radiation. Light and CO2 response curves indicate substantially superior photosynthetic capacity of K-sufficient trees, while Na substitution generated intermediate results. The enhanced performance of K, and to a lesser extent, Na-supplied trees was found to be related mainly to modification of non-stomatal limitation. This indicates that K deficiency promotes inhibition of enzymatic-photochemical processes. Results indicate lower chlorophyll content and altered Rubisco activity as probable causes of photosynthetic impairment. Potassium deficiency was found to diminish photoprotection mechanisms due to reduced photosynthetic and photorespiratory capacity. The lower CO2 and O2 assimilation rate in K-deficient trees caused elevated levels of exited energy. Consequently, non-photochemical quenching, an alternative energy dispersion pathway, was increased. Nonetheless, K-deficient trees were shown to suffer from photodamage to photosystem-II. Sodium replacement considerably diminished the negative effect of K deficiency on photoprotection mechanisms. The overall impact of K and Na nutrition plays down any indirect effect on stomatal limitation and rather demonstrates the centrality of these elements in photochemical processes of photosynthesis and photoprotection.

Direct in vivo evidence of immense stem water exploitation in irrigated date palms.

During the summer, evaporative demand at midday often exceeds the transport capacity of most desert plants. However, date palms maintain their ecological dominance with sustained and uniquely high rates of transpiration. This high rate of flow cannot be attributed to soil water supply alone. In order to quantify intra-plant water allocation in irrigated date palms, three water-sensing techniques have been incorporated: heat dissipation, gravimetric sampling, and time domain reflectrometry. Each of these methods has known limitations but their integration resulted in a quantitative in vivo accounting of the date palm diurnal and seasonal water mass balance. By incorporating these methods it was possible to determine that date palms substantially rely on the exploitation and recharge of the stem reservoir in their water budget. The stem of mature date palms can hold up to 1 m(3) of water and supply 25% of daily transpiration (i.e. 5000 l of water in 100 d of summer). The internal stem water reservoir is consistently recharged by over 50 l per night which allows for successive daytime reuse throughout the entire growing season. More broadly, these findings suggest that internal water allocation and night-time soil-water availability could provide useful information for improving date palm irrigation practices.

Estimating sap flux densities in date palm trees using the heat dissipation method and weighing lysimeters.

In a world of diminishing water reservoirs and a rising demand for food, the practice and development of water stress indicators and sensors are in rapid progress. The heat dissipation method, originally established by Granier, is herein applied and modified to enable sap flow measurements in date palm trees in the southern Arava desert of Israel. A long and tough sensor was constructed to withstand insertion into the date palm's hard exterior stem. This stem is wide and fibrous, surrounded by an even tougher external non-conducting layer of dead leaf bases. Furthermore, being a monocot species, water flow does not necessarily occur through the outer part of the palm's stem, as in most trees. Therefore, it is highly important to investigate the variations of the sap flux densities and determine the preferable location for sap flow sensing within the stem. Once installed into fully grown date palm trees stationed on weighing lysimeters, sap flow as measured by the modified sensors was compared with the actual transpiration. Sap flow was found to be well correlated with transpiration, especially when using a recent calibration equation rather than the original Granier equation. Furthermore, inducing the axial variability of the sap flux densities was found to be highly important for accurate assessments of transpiration by sap flow measurements. The sensors indicated no transpiration at night, a high increase of transpiration from 06:00 to 09:00, maximum transpiration at 12:00, followed by a moderate reduction until 08:00; when transpiration ceased. These results were reinforced by the lysimeters' output. Reduced sap flux densities were detected at the stem's mantle when compared with its center. These results were reinforced by mechanistic measurements of the stem's specific hydraulic conductivity. Variance on the vertical axis was also observed, indicating an accelerated flow towards the upper parts of the tree and raising a hypothesis concerning dehydrating mechanisms of the date palm tree. Finally, the sensors indicated reduction in flow almost immediately after irrigation of field-grown trees was withheld, at a time when no climatic or phenological conditions could have led to reduction in transpiration.

Functional anatomy controls ion distribution in banana leaves: significance of Na+ seclusion at the leaf margins.

Typical salt stress symptoms appear in banana (Musa sp., cv. 'Grand Nain' AAA) only along the leaf margins. Mineral analysis of the dry matter of plants treated with increasing concentrations of KCl or NaCl revealed significant accumulation of Na+, but not of K+ or Cl(-), in the affected leaf margins. The differential distribution of the three ions suggests that water and ion movement out of the xylem is mostly symplastic and, in contrast to K+ and Cl(-), there exists considerable resistance to the flow of Na+ from the xylem to the adjacent mesophyll and epidermis. The parallel veins of the lamina are enclosed by several layers of bundle sheath parenchyma; in contrast, the large vascular bundle that encircles the entire lamina, and into which the parallel veins merge, lacks a complete bundle sheath. Xylem sap containing a high concentration of Na+ is 'pulled' by water tension from the marginal vein back into the adjacent mesophyll without having to cross a layer of parenchyma tissue. When the marginal vein was dissected from the lamina, the pattern of Na+ distribution in the margins changed markedly. The distinct anatomy of the marginal vein plays a major role in the accumulation of Na+ in the margins, with the latter serving as a 'dumping site' for toxic molecules.