Thomas Eichert

Uptake of Hydrophilic Solutes Through Plant Leaves: Current State of Knowledge and Perspectives of Foliar Fertilization

Foliar fertilization is an agricultural practice of increasing importance in practical terms. Since nutrient sprays are mostly applied as water solutions, the focus of the article was placed on the penetration of ionic, polar solutes through the leaf surface, although the mechanisms of cuticular penetration of lipophilics are also taken into consideration. In theory, application of foliar nutrient sprays may indeed be a more target-oriented and environmentally friendly fertilization method since nutrients are directly delivered to the target organism in limited amounts, thereby helping to reduce the environmental impact associated with soil fertilization. However, response to foliar sprays is often variable and not reproducible due to the existing lack of knowledge of many factors related to the penetration of the leaf-applied solution. To overcome the current “spray and pray” situation, there is a need to critically analyze the major factors involved and the existing experimental approaches to safely assess the penetration mechanisms, which is the final aim of this review. Beginning with the significance of foliar fertilization in agriculture, a historical overview regarding foliar uptake related studies is given, with especial emphasis on the penetration of solutes through the cuticle and stomata. The existing models of cuticular and stomatal uptake are analyzed separately considering among other factors the physico-chemistry of the solution including the role of adjuvants and the effect of the environment. Methods employed to estimate the process of cuticular and stomatal penetration of solutes are critically assessed. Finally, conclusions related to multidisciplinary research perspectives for improving the efficiency of foliar sprays are drawn.

Size exclusion limits and lateral heterogeneity of the stomatal foliar uptake pathway for aqueous solutes and water-suspended nanoparticles

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.

Equivalent pore radii of hydrophilic foliar uptake routes in stomatous and astomatous leaf surfaces--further evidence for a stomatal pathway.

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.

Leaf structural changes associated with iron deficiency chlorosis in field-grown pear and peach: physiological implications

Plants grown in calcareous, high pH soils develop Fe deficiency chlorosis. While the physiological parameters of Fe-deficient leaves have been often investigated, there is a lack of information regarding structural leaf changes associated with such abiotic stress. Iron-sufficient and Fe-deficient pear and peach leaves have been studied, and differences concerning leaf epidermal and internal structure were found. Iron deficiency caused differences in the aspect of the leaf surface, which appeared less smooth in Fe-deficient than in Fe-sufficient leaves. Iron deficiency reduced the amount of soluble cuticular lipids in peach leaves, whereas it reduced the weight of the abaxial cuticle in pear leaves. In both plant species, epidermal cells were enlarged as compared to healthy leaves, whereas the size of guard cells was reduced. In chlorotic leaves, bundle sheaths were enlarged and appeared disorganized, while the mesophyll was more compacted and less porous than in green leaves. In contrast to healthy leaves, chlorotic leaves of both species showed a significant transient opening of stomata after leaf abscission (Iwanoff effect), which can be ascribed to changes found in epidermal and guard cells. Results indicate that Fe-deficiency may alter the barrier properties of the leaf surface, which can significantly affect leaf water relations, solute permeability and pest and disease resistance.

Review: Mechanisms for boron deficiency-mediated changes in plant water relations

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.

Greenhouse evaluation and environmental impact assessment of different urine-derived struvite fertilizers as phosphorus sources for plants.

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.

Solar thermal evaporation of human urine for nitrogen and phosphorus recovery in Vietnam

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.

Uptake and Release of Elements by Leaves and Other Aerial Plant Parts

Publisher Summary This chapter focuses on the importance of uptake and release of elements by leaves and other aerial parts and discusses the practical aspects of foliar fertilization and the ecological consequences of nutrient uptake and release. The surface of plants is covered by the cuticle, which is a bio-polymer synthesized by epidermal cells and primarily consists of cutin, a polyester matrix of polymerized long-chain fatty acids in which waxes are embedded. It protects the leaf from excessive water loss by transpiration and against excessive leaching of inorganic and organic solutes by rain. The hydrophobic nature of the cuticle makes it an effective barrier against the penetration by hydrophilic polar solutes, whereas lipophilic molecules may penetrate cuticles at much higher rates. Stomata act as adjustable apertures in the leaf surface, which optimize the trade-off between CO − uptake and water loss of plants. Various environmental conditions during plant growth, such as shading, high temperatures, and humidity, as well as nutrient deficiency, affect the structure and anatomy of the leaf surface, thereby affecting the uptake and release of elements by the leaves. Foliar sprays are widely used in agricultural production as an alternative or complementary strategy to soil fertilization, and the plant responses to elements supplied via foliar sprays are normally more rapid than to soil treatment. Thus it can help in preserving crop yields and quality, with low environmental impact.

Effects of iron chlorosis and iron resupply on leaf xylem architecture, water relations, gas exchange and stomatal performance of field-grown peach (Prunus persica)

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.

Allelopathic Monoterpenes Interfere with Arabidopsis thaliana Cuticular Waxes and Enhance Transpiration

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.