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The absorption of water and solutes by plant leaves has been recognised since more than two centuries. Given the polar nature of water and solutes, the mechanisms of foliar uptake have been proposed to be similar for water and electrolytes, including nutrient solutions. Research efforts since the 19th century focussed on characterising the properties of cuticles and applying foliar sprays to crop plants as a tool for improving crop nutrition. This was accompanied by the development of hundreds of studies aimed at characterising the chemical and structural nature of plant cuticles from different species and the mechanisms of cuticular and, to a lower extent, stomatal penetration of water and solutes. The processes involved are complex and will be affected by multiple environmental, physico-chemical and physiological factors which are only partially clear to date. During the last decades, the body of evidence that water transport across leaf surfaces of native species may contribute to water balances (absorption and loss) at an ecosystem level has grown. Given the potential importance of foliar water absorption for many plant species and ecosystems as shown in recent studies, the aim of this review is to first integrate current knowledge on plant surface composition, structure, wettability and physico-chemical interactions with surface-deposited matter. The different mechanisms of foliar absorption of water and electrolytes and experimental procedures for tracing the uptake process are discussed before posing several outstanding questions which should be tackled in future studies.

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RATIONALE: Boron (B) is an essential micronutrient in plants and its isotope variations are used to gain insights into plant metabolism, which is important for crop plant cultivation. B isotope variations were used to trace intra-plant fractionation mechanisms in response to the B concentration in the irrigation water spanning the range from B depletion to toxic levels. METHODS: A fully validated analytical procedure based on multi-collector inductively coupled plasma mass spectrometry (MC-ICP-MS), sample decomposition and B matrix separation was applied to study B isotope fractionation. The validation was accomplished by establishing a complete uncertainty budget and by applying reference materials, yielding expanded measurement uncertainties of 0.8‰ for pure boric acid solutions and ≤1.5‰ for processed samples. With this validated procedure SI traceable B isotope amount ratios were determined in plant reference materials for the first time. RESULTS: The B isotope compositions of irrigation water and bell pepper samples suggest passive diffusion of the heavy 11 B isotope into the roots during low to high B concentrations while uptake of the light 10 B isotope was promoted during B depletion, probably by active processes. A systematic enrichment of the heavy 11 B isotope in higher located plant parts was observed (average Δ11 Bleaf-roots = 20.3 ± 2.8‰ (1 SD)), possibly by a facilitated transport of the heavy 11 B isotope to growing meristems by B transporters. CONCLUSIONS: The B isotopes can be used to identify plant metabolism in response to the B concentration in the irrigation water and during intra-plant B transfer. The large B isotope fractionation within the plants demonstrates the importance of biological B cycling for the global B cycle.

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Most aerial plant surfaces are covered with a lipid-rich cuticle, which is a barrier for the bidirectional transport of substances between the plant and the surrounding environment. This review article provides an overview of the significance of the leaf cuticle as a barrier for the deposition and absorption of water and electrolytes. After providing insights into the physico-chemical properties of plant surfaces, the mechanisms of foliar absorption are revised with special emphasis on solutes. Due to the limited information and relative importance of the leaf cuticle of herbaceous and deciduous cultivated plants, an overview of the studies developed with Alpine conifers and treeline species is provided. The significance of foliar water uptake as a phenomenon of ecophysiological relevance in many areas of the world is also highlighted. Given the observed variability in structure and composition among, for example, plant species and organs, it is concluded that it is currently not possible to establish general permeability and wettability models that are valid for predicting liquid-surface interactions and the subsequent transport of water and electrolytes across plant surfaces.

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Foliar application of micronutrients (e.g. Fe3+ ) onto plants over an extended time is challenging and often not possible due to insufficient rainfastness. Smart delivery systems which enable micronutrient release over several weeks would offer innovative and sustainable options to improve plant health and food production. Herein, we report a novel foliar fertilizer delivery system based on functional pH-responsive biohybrid microgels that have orthogonal functionality as carriers of micronutrients and employ peptides (termed anchor peptides) as foliar adhesion promoters. The anchor peptides bind to hydrophobic surfaces and the waxy "islands" of plant leaves. Our system requires no auxiliaries and is loadable, storable, and applicable from aqueous dispersion. We report the synthesis and functionalization of microgels, their loading with Fe3+ ions, and a proof of concept for the biofunctional microgel-based fertilizer system is demonstrated for iron-deficient cucumber plants.

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Laser scanning is a non-invasive method for collecting and parameterizing 3D data of well reflecting objects. These systems have been used for 3D imaging of plant growth and structure analysis. A prerequisite is that the recorded signals originate from the true plant surface. In this paper we studied the effects of species, leaf chlorophyll content and sensor settings on the suitability and accuracy of a commercial 660 nm active laser triangulation scanning device. We found that surface images of Ficus benjamina leaves were inaccurate at low chlorophyll concentrations and a long sensor exposure time. Imaging of the rough waxy leaf surface of leek (Allium porrum) was possible using very low exposure times, whereas at higher exposure times penetration and multiple refraction prevented the correct imaging of the surface. A comparison of scans with varying exposure time enabled the target-oriented analysis to identify chlorotic, necrotic and healthy leaf areas or mildew infestations. We found plant properties and sensor settings to have a strong influence on the accuracy of measurements. These interactions have to be further elucidated before laser imaging of plants is possible with the high accuracy required for e.g., the observation of plant growth or reactions to water stress.

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Boron (B) is an essential microelement for plants and is constantly needed throughout the plant life due to its function as a structural element of the plant cell wall. B deficiency is a wide-spread problem in agricultural areas world-wide, and management of B nutrition is challenged by sudden occurrences of B deficiency or inconsistent effects of foliar B application. The effects of insufficient B supply on different structures relevant for the plant water status have been heavily researched, but the resulting conclusions are contradictory and no clear picture has so far emerged that fully explains the inconsistencies. B deficiency can affect water uptake by inhibition of root and shoot growth and by upregulation of water channels. Structural damage to xylem vessels can limit water transport to arial plant parts, while water loss can be altered by impaired barrier functions of leaf surfaces and reduced photosynthesis. In consequence of all these effects, transpiration is reduced in B-deficient plants under well-watered conditions. Under drought conditions, the responsiveness of stomata is impaired. Possible consequences of damaged vasculature for plant B nutrition include the reduced effectiveness of foliar B fertilization, especially in species with high B phloem mobility. Changes in leaf surface properties can further reduce B uptake after foliar application. In species with low B phloem mobility, weakened xylem vessels may not be able to supply sufficient B to arial parts under conditions of increased B demand, such as during bud development of trees. Since structural damage to vessels is hardly reversible, these effects could be permanent, even if B deficiency was only transient. Another consequence of reduced water status is the higher susceptibility of B-deficient plants to other abiotic stresses, which also impair water relations, especially drought. Since damage to vasculature can occur before visible symptoms of B deficiency appear in shoots, the importance to develop reliable diagnostic tools for detection of sub-acute B deficiency is highlighted.

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A selection of six urine-derived struvite fertilizers generated by innovative precipitation technologies was assessed for their quality and their effectiveness as phosphorus sources for crops. Struvite purity was influenced by drying techniques and magnesium dosage. In a greenhouse experiment, the urine fertilizers led to biomass yields and phosphorus uptakes comparable to or higher than those induced by a commercial mineral fertilizer. Heavy metal concentrations of the different struvite fertilizers were below the threshold limits specified by the German Fertilizer and Sewage Sludge Regulations. The computed loading rates of heavy metals to agricultural land were also below the threshold limits decreed by the Federal Soil Protection Act. Urine-derived struvite contributed less to heavy metal inputs to farmland than other recycling products or commercial mineral and organic fertilizers. When combined with other soil conditioners, urine-derived struvite is an efficient fertilizer which covers the magnesium and more than half of the phosphorus demand of crops.

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A No Mix sanitation system was installed in a dormitory at the University of Can Tho in Vietnam, with the objective of recycling nutrients from source separated urine. This paper presents a pilot scale evaporation technology, and investigates the feasibility of recovering nitrogen and phosphorus from human urine by solar still for use as fertilizer. After 26 days of sun exposure, 360 g of solid fertilizer material was recovered from 50 L undiluted urine. This urine-derived fertilizer was mainly composed of sodium chloride, and had phosphorus and nitrogen contents of almost 2%. When tested with maize and ryegrass, the urine fertilizer led to biomass yields and phosphorus and nitrogen uptakes comparable to those induced by a commercial mineral fertilizer. Urine acidification with sulfuric or phosphoric acid prior treatment reduced nitrogen losses, improved the nutrient content of the generated fertilizers, and induced higher biomass yields and nitrogen and phosphorus uptakes than the commercial mineral fertilizer. However, acidification is not recommended in developing countries due to additional costs and handling risks. The fate of micropollutants and the possibility of separating sodium chloride from other beneficial nutrients require further investigation.

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There is increasing evidence suggesting that iron (Fe) deficiency induces not only leaf chlorosis and a decline of photosynthesis, but also structural changes in leaf morphology, which might affect the functionality of leaves. In this study, we investigated the effects of Fe deficiency on the water relations of peach (Prunus persica (L.) Batsch.) leaves and the responses of previously chlorotic leaves to Fe resupply via the root or the leaf. Iron deficiency induced a decline of maximum potential photosystem II (PSII) efficiency (F(V)/F(M)), of rates of net photosynthesis and transpiration and of water use efficiency. Iron chlorosis was associated with a reduction of leaf xylem vessel size and of leaf hydraulic conductance. In the course of the day, water potentials in chlorotic leaves remained higher (less negative) than in green leaves. In chlorotic leaves, normal stomatal functioning was disturbed, as evidenced by the lack of opening upon withdrawal of external CO(2) and stomatal closure after sudden illumination of previously darkened leaves. We conclude that the Fe deficiency induced limitations of xylem conductivity elicited a water saving strategy, which poses an additional challenge to plant growth on high pH, calcareous soils. Fertilisation with Fe improved photosynthetic performance but the proper xylem structure and water relations of leaves were not fully restored, indicating that Fe must be available at the first stages of leaf growth and development.

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Penetration rates of foliar-applied polar solutes are highly variable and the underlying mechanisms are not yet fully understood. The contribution of stomata especially, is still a matter of debate. Thus, the size exclusion limits of the stomatal foliar uptake pathway, its variability and its transport capacity have been investigated. The size exclusion limits were analyzed by studying the penetration of water-suspended hydrophilic particles of two different sizes (43 nm or 1.1 microm diameter) into leaves of Vicia faba (L.). To avoid agglutination of the particles, plants were kept in water-saturated atmosphere. Penetration of the larger particles was never detected, whereas after 2 to 9 days, the smaller particles occasionally penetrated the leaf interior through stomatal pores. Permeability of stomata to Na(2)-fluorescein along the leaf blade of Allium porrum (L.) was highly variable and not correlated with the position on the leaf. When evaporated residues of the foliar-applied solutions were rewetted repeatedly, approximately 60% of the previously penetrated stomata were penetrated again. The average rate constant of penetration of an individual stoma was in the same order of magnitude as typical rate constants reported for the cuticular pathway. The observed sparseness of stomatal penetration together with its high lateral variability but local and temporal persistency was taken as evidence that stomata contributing to uptake differ from non-penetrated ones in the wettability of their guard cell cuticle. These results show that the stomatal pathway is highly capacitive because of its large size exclusion limit above 10 nm and its high transport velocity, but at the same time the high variability renders this pathway largely unpredictable.

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Foliar uptake pathways for hydrophilic solutes were studied by the analysis of co-uptake of 15N-labelled urea, NH4+ or NO3- and 13C-labelled sucrose across leaf surfaces of various plant species. Uptake of N (y) and sucrose (x) were strongly correlated. Curvilinear regression revealed significantly positive intercepts with the y-axis indicating the involvement of a sucrose-excluding pathway consisting of small pores with radii <0.5 nm. Depending on plant species, N source, leaf side and aperture of stomata, these small pores accounted for 6-62% of total N uptake. Regression analysis revealed that in stomatous leaf surfaces of Vicia faba L., Coffea arabica L. and Prunus cerasus L., the remaining N uptake occurred via another pathway with an estimated average pore radius (r(P)) greater than 20 nm. This is two orders of magnitude greater than previous estimations of cuticular r(P), indicating that this pathway, which was only found in stomatous leaf surfaces, was probably not located in the cuticle but at the surfaces of the stomatal pores. In astomatous leaf surfaces of C. arabica and Populus x canadensis Moench, average r(P) was 2.0 and 2.4 nm, respectively, which is four to eight times larger than previous estimations of cuticular r(P). These results indicate that for polar solutes, the size exclusion limits of plant surfaces can be considerably larger than previously estimated. The far-reaching implications of these findings are discussed.

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Exposure to the allelopathic monoterpenes camphor (100 mg/10 L) and menthol (50 mg/10 L) for 24 h enhanced transpiration of Arabidopsis thaliana fully developed rosette leaves similar to de-waxing. As ascertained by ESEM analyses the leaf surfaces were spotted with platelet like structures which seem to be partly mixed with the lipophilic epicuticular layers. The structures are supposed to contain the condensed monoterpenes, which could be identified by GC. Long term exposure (more than 48 h) to 100 mg/50 mg killed the plants by desiccation, a 24 h exposure caused necrotic spots that became visible one to two days after the treatment. Examinations of the stomatal apertures indicated that monoterpenes induced stomatal opening followed by extreme swelling and a final break down of the protoplasts. Exposure of Arabidopsis thaliana to volatiles of Mentha piperita, Lavandula latifolia and Artemisia camphorata resulted in a dramatic increase of the stomata aperture but swelling of the protoplasts was less exhibited.In contrast to de-waxing, expression of the fatty acid condensing enzyme encoding CER6 gene and de novo synthesis of CER6 protein was not induced after 24 h of exposure to the monoterpenes.The aim of the study was to demonstrate that the lipophilic layers of the leaf surface and the stomata are primary targets of monoterpene allelopathic attack. Enhanced transpiration results from a combination of affected lipophilic wax layers and a disturbed stomata function.