Ambient aerosols increase stomatal transpiration and conductance of hydroponic sunflowers by extending the hydraulic system to the leaf surface.

Introduction: Many atmospheric aerosols are hygroscopic and play an important role in cloud formation. Similarly, aerosols become sites of micro-condensation when they deposit to the upper and lower surfaces of leaves. Deposited salts, in particular can trigger condensation at humidities considerably below atmospheric saturation, according to their hygroscopicity and the relative humidity within the leaf boundary layer. Salt induced water potential gradients and the resulting dynamics of concentrated salt solutions can be expected to affect plant water relations. Methods: Hydroponic sunflowers were grown in filtered (FA) and unfiltered, ambient air (AA). Sap flow was measured for 18 days and several indicators of incipient drought stress were studied. Results: At 2% difference in mean vapor pressure deficit (D), AA sunflowers had 49% higher mean transpiration rates, lower osmotic potential, higher proline concentrations, and different tracer transport patterns in the leaf compared to FA sunflowers. Aerosols increased plant conductance particularly at low D. Discussion: The proposed mechanism is that thin aqueous films of salt solutions from deliquescent deposited aerosols enter into stomata and cause an extension of the hydraulic system. This hydraulic connection leads - parallel to stomatal water vapor transpiration - to wick-like stomatal loss of liquid water and to a higher impact of D on plant water loss. Due to ample water supply by hydroponic cultivation, AA plants thrived as well as FA plants, but under more challenging conditions, aerosol deposits may make plants more susceptible to drought stress.

Nocturnal Transpiration May Be Associated with Foliar Nutrient Uptake.

Aerosols can contribute to plant nutrition via foliar uptake. The conditions for this are best at night because the humidity is high and hygroscopic, saline deposits can deliquesce as a result. Still, stomata tend to be closed at night to avoid unproductive water loss. However, if needed, nutrients are on the leaf surface, and plants could benefit from nocturnal stomatal opening because it further increases humidity in the leaf boundary layer and allows for stomatal nutrient uptake. We tested this hypothesis on P-deficient soil by comparing the influence of ambient aerosols and additional foliar P application on nocturnal transpiration. We measured various related leaf parameters, such as the foliar water loss, minimum leaf conductance (gmin), turgor loss point, carbon isotope ratio, contact angle, specific leaf area (SLA), tissue element concentration, and stomatal and cuticular characteristics. For untreated leaves grown in filtered, aerosol-free air (FA), nocturnal transpiration consistently decreased overnight, which was not observed for leaves grown in unfiltered ambient air (AA). Foliar application of a soluble P salt increased nocturnal transpiration for AA and FA leaves. Crusts on stomatal rims were shown by scanning electron microscopy, supporting the idea of stomatal uptake of deliquescent salts. Turgor loss point and leaf moisture content indicated a higher accumulation of solutes, due to foliar uptake by AA plants than FA plants. The hypothesis that deliquescent leaf surface salts may play a role in triggering nocturnal transpiration was supported by the results. Still, further experiments are required to characterize this phenomenon better.

Heterogeneity of Stomatal Pore Area Is Suppressed by Ambient Aerosol in the Homobaric Species, Vicia faba.

Stomatal pore area is heterogeneous across leaf surfaces. This has been considered as "patchy stomatal conductance," and may have substantial implications for photosynthetic efficiency. Aerosols have always been important elements of plant environments, but their effects on stomatal control of plant water relations, and stomatal heterogeneity specifically, have not been considered. Here we evaluate the spatial coordination of pore area in the glabrous and homobaric leaves of Vicia faba grown under two aerosol treatments and measured at four levels of VPD. We construct a large dataset (n > 88,000 discrete comparisons) of paired pore areas and distances between the pores. Plants were grown in ambient urban air and in filtered air (FA) to determine the effect of ambient aerosol on stomatal properties. Pore area exhibited spatial organization, as well as considerable variability among closely co-located pores. The difference between pore areas was positively correlated with the distance between the pores, in both aerosol treatments and at all VPDs. However, aerosol deposition reduced both the magnitude of variability between pores and the rate at which this variability increased with pore separation distance. These data support previous conclusions that deposition of hygroscopic aerosol may create a thin aqueous film across the leaf surface that connects neighboring stomata to each other and to the leaf interior. Aerosol impacts on stomatal heterogeneity and gas exchange are not adequately considered in current assessments of stomatal control.

Ambient aerosol increases minimum leaf conductance and alters the aperture-flux relationship as stomata respond to vapor pressure deficit (VPD).

Aerosols are important components of the global plant environment, with beneficial and deleterious impacts. The direct effects of aerosol deposition on plant-water relationships remain poorly characterized but potentially important. Vicia faba was grown in ambient urban air and in the same air with aerosol excluded, in a moderately polluted environment using two exposure protocols. Simultaneous measurement of gas exchange and stomatal pore aperture was combined with leaf dehydration kinetics and microscopic evaluation of leaf wetness formation and aerosol deposition patterns. The ambient aerosol was shown to be hygroscopic. Aerosol exposure increased minimum leaf conductance, shown by dehydration kinetics, and nocturnal water vapor flux, shown by dark-adapted gas exchange. Aerosol exposure decreased stomatal apertures at each level of vapor pressure deficit (VPD) and increased stomatal conductance at comparable levels of aperture. Overall, these effects were modest, and largest when stomata were wide open. The uncoupling of conductance (flux-based) from aperture (directly measured microscopically) implies that aerosol-induced water loss is not fully under stomatal control. This reduces drought tolerance and may provide a mechanism by which deposited aerosol plays a direct role in stomatal response to VPD.

Xeromorphic traits help to maintain photosynthesis in the perhumid climate of a Taiwanese cloud forest.

Previous flux measurements in the perhumid cloud forest of northeastern Taiwan have shown efficient photosynthesis of the endemic tree species Chamaecyparis obtusa var. formosana even under foggy conditions in which leaf surface moisture would be expected. We hypothesized this to be the result of 'xeromorphic' traits of the Chamaecyparis leaves (hydrophobicity, stomatal crypts, stomatal clustering), which could prevent coverage of stomata by precipitation, fog, and condensation, thereby maintaining CO2 uptake. Here we studied the amount, distribution, and composition of moisture accumulated on Chamaecyparis leaf surfaces in situ in the cloud forest. We studied the effect of surface tension on gas penetration to stomata using optical O2 microelectrodes in the laboratory. We captured the dynamics of condensation to the leaf surfaces with an environmental scanning electron microscope (ESEM). In spite of substantial surface hydrophobicity, the mean water film thickness on branchlets under foggy conditions was 80 µm (upper surface) and 40 µm (lower surface). This amount of water could cover stomata and prevent CO2 uptake. This is avoided by the clustered arrangement of stomata within narrow clefts and the presence of Florin rings. These features keep stomatal pores free from water due to surface tension and provide efficient separation of plant and atmosphere in this perhumid environment. Air pollutants, particularly hygroscopic aerosol, may disturb this functionality by enhancing condensation and reducing the surface tension of leaf surface water.

Particulate pollutants are capable to 'degrade' epicuticular waxes and to decrease the drought tolerance of Scots pine (Pinus sylvestris L.).

Air pollution causes the amorphous appearance of epicuticular waxes in conifers, usually called wax 'degradation' or 'erosion', which is often correlated with tree damage symptoms, e.g., winter desiccation. Previous investigations concentrated on wax chemistry, with little success. Here, we address the hypothesis that both 'wax degradation' and decreasing drought tolerance of trees may result from physical factors following the deposition of salt particles onto the needles. Pine seedlings were sprayed with dry aerosols or 50 mM solutions of different salts. The needles underwent humidity changes within an environmental scanning electron microscope, causing salt expansion on the surface and into the epistomatal chambers. The development of amorphous wax appearance by deliquescent salts covering tubular wax fibrils was demonstrated. The minimum epidermal conductance of the sprayed pine seedlings increased. Aerosol deposition potentially 'degrades' waxes and decreases tree drought tolerance. These effects have not been adequately considered thus far in air pollution research.

"Breath figures" on leaf surfaces-formation and effects of microscopic leaf wetness.

"Microscopic leaf wetness" means minute amounts of persistent liquid water on leaf surfaces which are invisible to the naked eye. The water is mainly maintained by transpired water vapor condensing onto the leaf surface and to attached leaf surface particles. With an estimated average thickness of less than 1 µm, microscopic leaf wetness is about two orders of magnitude thinner than morning dewfall. The most important physical processes which reduce the saturation vapor pressure and promote condensation are cuticular absorption and the deliquescence of hygroscopic leaf surface particles. Deliquescent salts form highly concentrated solutions. Depending on the type and concentration of the dissolved ions, the physicochemical properties of microscopic leaf wetness can be considerably different from those of pure water. Microscopic leaf wetness can form continuous thin layers on hydrophobic leaf surfaces and in specific cases can act similar to surfactants, enabling a strong potential influence on the foliar exchange of ions. Microscopic leaf wetness can also enhance the dissolution, the emission, and the reaction of specific atmospheric trace gases e.g., ammonia, SO2, or ozone, leading to a strong potential role for microscopic leaf wetness in plant/atmosphere interaction. Due to its difficult detection, there is little knowledge about the occurrence and the properties of microscopic leaf wetness. However, based on the existing evidence and on physicochemical reasoning it can be hypothesized that microscopic leaf wetness occurs on almost any plant worldwide and often permanently, and that it significantly influences the exchange processes of the leaf surface with its neighboring compartments, i.e., the plant interior and the atmosphere. The omission of microscopic water in general leaf wetness concepts has caused far-reaching, misleading conclusions in the past.

Stomatal penetration by aqueous solutions--an update involving leaf surface particles.

The recent visualization of stomatal nanoparticle uptake ended a 40-yr-old paradigm. Assuming clean, hydrophobic leaf surfaces, the paradigm considered stomatal liquid water transport to be impossible as a result of water surface tension. However, real leaves are not clean, and deposited aerosols may change hydrophobicity and water surface tension. Droplets containing NaCl, NaClO(3), (NH(4))(2) SO(4), glyphosate, an organosilicone surfactant or various combinations thereof were evaporated on stomatous abaxial and astomatous adaxial surfaces of apple (Malus domestica) leaves. The effects on photosynthesis, necrosis and biomass were determined. Observed using an environmental scanning electron microscope, NaCl and NaClO(3) crystals on hydrophobic tomato (Solanum lycopersicum) cuticles underwent several humidity cycles, causing repeated deliquescence and efflorescence of the salts. All physiological parameters were more strongly affected by abaxial than adaxial treatments. Spatial expansion and dendritic crystallization of the salts occurred and cuticular hydrophobicity was decreased more rapidly by NaClO(3) than NaCl. The results confirmed the stomatal uptake of aqueous solutions. Humidity fluctuations promote the spatial expansion of salts into the stomata. The ion-specific effects point to the Hofmeister series: chaotropic ions reduce surface tension, probably contributing to the defoliant action of NaClO(3), whereas the salt spray tolerance of coastal plants is probably linked to the kosmotropic nature of chloride ions.