New insights into the role of the root system of epiphytic bromeliads: comparison of root and leaf trichome functions in acquisition of water and nutrients.

BACKGROUND: In epiphytic bromeliads, the roots used to be considered poorly functional organs in the processes of absorption and metabolization of water and nutrients, while the leaves always acted as protagonists in both functions. More recent discoveries have been changing this old view of the root system. SCOPE: In this review, we will address the old thoughts of the scientific community regarding the function performed by the roots of epiphytic bromeliads (mere holdfast structures with low physiological activity) and the importance of a reduced or lack of root system for the emergence of epiphytism. We will present indirect and direct evidence that contradicts this older hypothesis. Furthermore, the importance of the root absorptive function mainly for juvenile tankless epiphytic bromeliads and the characteristics of the root absorption process of adult epiphytic tank bromeliads will be thoroughly discussed in physiological aspects. Finally, some factors (species, substrate, environmental conditions) that influence the absorptive capability of the roots of epiphytic tank bromeliads will also be considered in this review, highlighting the importance that the absorptive role of the roots have for the plasticity of bromeliads that live on trees, which is an environment characterized by the intermittent availability of water and nutrients. CONCLUSIONS: The roots of tank-forming epiphytic bromeliads play important roles in the absorption and metabolization of nutrients and water. The importance of roots stands out mainly for juvenile tankless bromeliads since the root is the main absorptive organ. In larger plants with tank, although the leaves become the protagonists in the resource acquisition process, the roots complement the absorptive function of the leaf trichomes, resulting in a better growth of the bromeliad. The physiological and biochemical properties of the processes of absorption and distribution of resources in the tissues seem to differ between absorption by trichomes and roots.

Isotope Evidence for Rice Accumulation of Newly Deposited and Soil Legacy Cadmium: A Three-Year Field Study.

Biogeochemical processes of atmospherically deposited cadmium (Cd) in soils and accumulation in rice were investigated through a three-year fully factorial atmospheric exposure experiment using Cd stable isotopes and diffusive gradients in thin films (DGT). Our results showed that approximately 37-79% of Cd in rice grains was contributed by atmospheric deposition through root and foliar uptake during the rice growing season, while the deposited Cd accounted for a small proportion of the soil pools. The highly bioavailable metals in atmospheric deposition significantly increased the soil DGT-measured bioavailable fraction; yet, this fraction rapidly aged following a first-order exponential decay model, leading to similar percentages of the bioavailable fraction in soils exposed for 1-3 years. The enrichment of light Cd isotopes in the atmospheric deposition resulted in a significant shift toward lighter Cd isotopes in rice plants. Using a modified isotopic mass balance model, foliar and root uptake of deposited Cd accounted for 47-51% and 28-36% in leaves, 41-45% and 22-30% in stems, and 45-49% and 26-30% in grains, respectively. The implications of this study are that new atmospheric deposition disproportionately contributes to the uptake of Cd in rice, and managing emissions thus becomes very important versus remediation of impacted soils.

Acid waters in tank bromeliads: Causes and potential consequences.

PREMISE: The consequences of acidity for plant performance are profound, yet the prevalence and causes of low pH in bromeliad tank water are unknown despite its functional relevance to key members of many neotropical plant communities. METHODS: We investigated tank water pH for eight bromeliad species in the field and for the widely occurring Guzmania monostachia in varying light. We compared pH changes over time between plant and artificial tanks containing a solution combined from several plants. Aquaporin transcripts were measured for field plants at two levels of pH. We investigated relationships between pH, leaf hydraulic conductance, and CO2 concentration in greenhouse plants and tested proton pump activity using a stimulator and inhibitor. RESULTS: Mean tank water pH for the eight species was 4.7 ± 0.06 and was lower for G. monostachia in higher light. The pH of the solution in artificial tanks, unlike in plants, did not decrease over time. Aquaporin transcription was higher for plants with lower pH, but leaf hydraulic conductance did not differ, suggesting that the pH did not influence water uptake. Tank pH and CO2 concentration were inversely related. Fusicoccin enhanced a decrease in tank pH, whereas orthovanadate did not. CONCLUSIONS: Guzmania monostachia acidified its tank water via leaf proton pumps, which appeared responsive to light. Low pH increased aquaporin transcripts but did not influence leaf hydraulic conductance, hence may be more relevant to nutrient uptake.

Stable Isotope Ratios Trace the Rice Uptake of Cadmium from Atmospheric Deposition via Leaves and Roots.

Cadmium (Cd) stable isotopes provide a novel technique to investigate the fate of Cd in the environment, but challenges exist for tracing the sources in the plants. We performed individual rice leaf and root exposures to dry and wet deposition using customized open-top chambers (OTCs) in the greenhouse and in the field next to a smelter, respectively. The field experiment also included a control without Cd deposition and a "full" treatment. The exposure experiments and isotope signatures showed that leaves can directly take up atmospheric Cd and then translocate within rice plants to other tissues, contributing 52-70% of Cd in grains, which exceeded the contribution (30-48%) by root exposure. The Cd isotopes in leaves, nodes, internodes, and grains demonstrate that roots preferentially take up Cd from wet deposition, but leaves favor uptake of Cd from dry deposition. The Cd uptake by leaves is redistributed via nodes, allowing for upward transport to the grains but preventing downward transport to the roots. Leaves favor uptake of heavy isotopes from atmospheric deposition (ΔCd114/110Leaf-Dust: 0.10 ± 0.02‰) but retain light isotopes and transport heavy isotopes to the nodes and further to grains. These findings highlight the contribution of atmospheric deposition to rice and Cd isotopes as a useful tracer for quantifying sources in plants when different isotopic compositions are in sources.

Dynamic Behaviors of Newly Deposited Atmospheric Heavy Metals in the Soil-Pak Choi System.

Dynamic behaviors of the newly deposited atmospheric heavy metals in the soil-pak choi (Brassica chinensis L.) system are investigated by a fully factorial atmospheric exposure experiment using soils exposed to 0.5-year and 1.5-year atmospheric depositions. The results showed approximately 17-87%, 19-64%, and 43-84% of the Cu, Cd, and Pb in pak choi edible parts were contributed from the new depositions, respectively. For the newly deposited metals, foliar uptake was the key pathway of shoot bioaccumulation rather than from root uptake of the deposited metals in soils, resulting in no significant soil contribution differences between pak chois growing in 0.5-year and 1.5-year exposed soils. Indeed, highly bioavailable metals in atmospheric deposition significantly increased the soil plant-bioavailable Cu, Cd, and Pb fractions; however, soil aging resulted in similar percentages of the plant-bioavailable fractions in 0.5-year and 1.5-year exposed soils, which indicated the bioavailability of metals deposited into soils rapidly decreased with aging. The soil aging process of the deposited metals was well fitted with the first-order exponential decay model, and soil organic matter and clay were the major driving factors. Our findings highlight high plant bioaccumulation rates and the rapid soil aging process of newly deposited metals during the plant growth period.

Direct foliar uptake of phosphorus from desert dust.

Phosphorus (P) scarcity constrains plant growth in many ecosystems worldwide. In P-poor ecosystems, the biogeochemical paradigm links plant productivity with the deposition of P-rich dust originating from desert storms. However, dust P usually has low bioavailability and is thought to be utilized solely via roots. We applied desert dust on the leaf surface of P-deficient and P-sufficient wheat, chickpea and maize to test the feasibility of direct foliar uptake of dust-P and investigate its related acquisition mechanisms. Foliar dust doubled the growth of P-deficient chickpea and wheat, crops originating near the Syrian Desert. P deficiency stimulated several leaf modifications that enabled acquisition of up to 30% of the sparingly soluble dust-P that is conventionally perceived as unavailable. These modifications increased foliar dust capture, acidified the leaf surface and, in chickpea, enhanced exudation of P-solubilizing organic acids. Maize (originating far from deserts) displayed only a marginal response to dust. The dramatic response of chickpea and wheat in comparison to maize suggests that plants that evolved in dust-rich ecosystems adopted specialized utilization strategies. Interestingly, the abovementioned foliar responses are comparable to known P uptake root responses. Given that P limitation is almost universal, a foliar P uptake pathway will have significant ecological and agricultural implications.

Ecotoxicology of Heavy Metal(loid)-Enriched Particulate Matter: Foliar Accumulation by Plants and Health Impacts.

Atmospheric contamination by heavy metal-enriched particulate matter (metal-PM) is highly topical nowadays because of its high persistence and toxic nature. Metal-PMs are emitted to the atmosphere by various natural and anthropogenic activities, the latter being the major source. After being released into the atmosphere, metal-PM can travel over a long distance and can deposit on the buildings, water, soil, and plant canopy. In this way, these metal-PMs can contaminate different parts of the ecosystem. In addition, metal-PMs can be directly inhaled by humans and induce several health effects. Therefore, it is of great importance to understand the fate and behavior of these metal-PMs in the environment. In this review, we highlighted the atmospheric contamination by metal-PMs, possible sources, speciation, transport over a long distance, and deposition on soil, plants, and buildings. This review also describes the foliar deposition and uptake of metal-PMs by plants. Moreover, the inhalation of these metal-PMs by humans and the associated health risks have been critically discussed. Finally, the article proposed some key management strategies and future perspectives along with the summary of the entire review. The abovementioned facts about the biogeochemical behavior of metal-PMs in the ecosystem have been supported with well-summarized tables (total 14) and figures (4), which make this review article highly informative and useful for researchers, scientists, students, policymakers, and the organizations involved in development and management. It is proposed that management strategies should be developed and adapted to cope with atmospheric release and contamination of metal-PM.

Foliar uptake of atmospheric nitrate by two dominant subalpine plants: insights from in situ triple-isotope analysis.

The significance of foliar uptake of nitrogen (N) compounds in natural conditions is not well understood, despite growing evidence of its importance to plant nutrition. In subalpine meadows, N-limitation fosters the dominance of specific subalpine plant species, which in turn ensures the provision of essential ecosystems services. Understanding how these plants absorb N and from which sources is important in predicting ecological consequences of increasing N deposition. Here, we investigate the sources of N to plants from subalpine meadows with distinct land-use history in the French Alps, using the triple isotopes (Δ17 O, δ18 O, and δ15 N) of plant tissue nitrate (NO3- ). We use this approach to evaluate the significance of foliar uptake of atmospheric NO3- (NO3-atm ). The foliar uptake of NO3-atm accounted for 4-16% of the leaf NO3- content, and contributed more to the leaf NO3- pool after peak biomass. Additionally, the gradual 15 N enrichment of NO3- from the soil to the leaves reflected the contribution of NO3-atm assimilation to plants' metabolism. The present study confirms that foliar uptake is a potentially important pathway for NO3-atm into subalpine plants. This is of major significance as N emissions (and deposition) are predicted to increase globally in the future.

Dew-induced transpiration suppression impacts the water and isotope balances of Colocasia leaves.

Foliar uptake of water from the surface of leaves is common when rainfall is scarce and non-meteoric water such as dew or fog is more abundant. However, many species in more mesic environments have hydrophobic leaves that do not allow the plant to uptake water. Unlike foliar uptake, all species can benefit from dew- or fog-induced transpiration suppression, but despite its ubiquity, transpiration suppression has so far never been quantified. Here, we investigate the effect of dew-induced transpiration suppression on the water balance and the isotope composition of leaves via a series of experiments. Characteristically, hydrophobic leaves of a tropical plant, Colocasia esculenta, are misted with isotopically enriched water to reproduce dew deposition. This species does not uptake water from the surface of its leaves. We measure leaf water isotopes and water potential and find that misted leaves exhibit a higher water potential and a more depleted water isotope composition than dry leaves, suggesting a ∼ 30% decrease in transpiration rate compared to control leaves. We propose three possible mechanisms governing the interaction of water droplets with leaf energy balance: increase in albedo from the presence of dew droplets, decrease in leaf temperature from the evaporation of dew, and local decrease in vapor pressure deficit. Comparing previous studies on foliar uptake to our results, we conclude that transpiration suppression has an effect of similar amplitude, yet opposite sign to foliar uptake on leaf water isotopes.

Foliar uptake of radiocaesium from irrigation water by paddy rice (Oryza sativa): an overlooked pathway in contaminated environments.

Flooded (paddy) rice (Oryza sativa) can take up ions from the irrigation water by foliar uptake via the exposed stem base. We hypothesised that the stem base uptake of radiocaesium (RCs) is a pathway for rice grown in RCs-contaminated environments. We developed a bi-compartmental device which discriminates the stem base from root RCs uptake from solutions, thereby using RCs isotopes (137 Cs and 134 Cs) with < 2% solution leak between the compartments. Radiocaesium uptake was linear over time (0-24 h). Radiocaesium uptake to the entire plant, expressed per dry weight of the exposed parts, was sixfold higher for the roots than for the exposed stem base. At equal RCs concentrations in both compartments, the exposed stem base and root uptake contributed almost equally to the total shoot RCs concentrations. Reducing potassium supply to the roots not only increased the root RCs uptake but also increased RCs uptake by the stem base. This study was the first to experimentally demonstrate active and internally regulated RCs uptake by the stem base of rice. Scenario calculations for the Fukushima-affected area predict that RCs in irrigation water could be an important source of RCs in rice as indirectly suggested from field data.

Physico-chemical properties of plant cuticles and their functional and ecological significance.

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.

Bark water uptake promotes localized hydraulic recovery in coastal redwood crown.

Coastal redwood (Sequoia sempervirens), the world's tallest tree species, rehydrates leaves via foliar water uptake during fog/rain events. Here we examine if bark also permits water uptake in redwood branches, exploring potential flow mechanisms and biological significance. Using isotopic labelling and microCT imaging, we observed that water entered the xylem via bark and reduced tracheid embolization. Moreover, prolonged bark wetting (16 h) partially restored xylem hydraulic conductivity in isolated branch segments and whole branches. Partial hydraulic recovery coincided with an increase in branch water potential from about -5.5 ± 0.4 to -4.2 ± 0.3 MPa, suggesting localized recovery and possibly hydraulic isolation. As bark water uptake rate correlated with xylem osmotic potential (R(2) = 0.88), we suspect a symplastic role in transferring water from bark to xylem. Using historical weather data from typical redwood habitat, we estimated that bark and leaves are wet more than 1000 h per year on average, with over 30 events being sufficiently long (>24 h) to allow for bark-assisted hydraulic recovery. The capacity to uptake biologically meaningful volumes of water via bark and leaves for localized hydraulic recovery throughout the crown during rain/fog events might be physiologically advantageous, allowing for relatively constant transpiration.

Kinetic study of phytotoxicity induced by foliar lead uptake for vegetables exposed to fine particles and implications for sustainable urban agriculture.

At the global scale, foliar metal transfer occurs for consumed vegetables cultivated in numerous urban or industrial areas with a polluted atmosphere. However, the kinetics of metal uptake, translocation and involved phytotoxicity was never jointly studied with vegetables exposed to micronic and sub-micronic particles (PM). Different leafy vegetables (lettuces and cabbages) cultivated in RHIZOtest® devices were, therefore, exposed in a greenhouse for 5, 10 and 15days to various PbO PM doses. The kinetics of transfer and phytotoxicity was assessed in relation to lead concentration and exposure duration. A significant Pb accumulation in leaves (up to 7392mg/kg dry weight (DW) in lettuce) with translocation to roots was observed. Lead foliar exposure resulted in significant phytotoxicity, lipid composition change, a decrease of plant shoot growth (up to 68.2% in lettuce) and net photosynthesis (up to 58% in lettuce). The phytotoxicity results indicated plant adaptation to Pb and a higher sensitivity of lettuce in comparison with cabbage. Air quality needs, therefore, to be considered for the health and quality of vegetables grown in polluted areas, such as certain megacities (in China, Pakistan, Europe, etc.) and furthermore, to assess the health risks associated with their consumption.

Field study on the uptake pathways and their contributions to the accumulation of organophosphate esters, phthalates, and polycyclic aromatic hydrocarbons in upland rice.

Plant uptake of organic contaminants generally occurs through either root, gas-phase foliar, or particle-phase foliar uptake. Understanding these pathways is essential for food-system practitioners to reduce human exposures, and to clean contaminated-sites with phytoremediation. Herein, we conducted a field-based experiment using an improved specific exposure chamber to elucidate the uptake pathways of organophosphate esters, phthalates, and polycyclic aromatic compounds, and quantitatively assessed their contributions to organic contaminant accumulations in field-grown rice. For most target compounds, all three uptake pathways (root, foliar gas, and foliar particle uptakes) contributed substantially to the overall contaminant burden in rice. Compounds with lower octanol-water partition coefficients (Kow) were more readily translocated from roots to leaves, and compounds with higher octanol-air partition coefficients (Koa) tended to enter rice leaves mostly through particle deposition. Most compounds were mostly stored in the inner leaves (55.3-98.2 %), whereas the relatively volatile compounds were more readily absorbed by the waxy layer and then transferred to the inner leaves. Air particle desorption was a key process regulating foliar uptake of low-volatility compounds. The results can help us to better understand and predict the environmental fate of those contaminants, and develop more effective management strategies for reducing their human exposure through food ingestion.

Effects of uptake pathways on the accumulation, translocation, and metabolism of OPEs in rice: An emphasis on foliar uptake.

The often-overlooked importance of foliar absorption on the plant uptake of organic pollutants was investigated by an exposure chamber test. Rice seedlings were exposed to organophosphate esters (OPEs) through 8 scenarios arranged from 3 major uptake pathways: root uptake via solution, foliar uptake via gas, and foliar uptake via particles, to identify the contributions of these 3 uptake pathways and their influences on the translocation and metabolism of OPEs in rice. The concentration of OPEs in rice tissues showed an "additive effect" with the increase of exposure pathways. OPEs in rice shoots mainly originated from foliar uptake through particle (29.6 %-63.5 %) and gaseous (28.5 %-49.4 %) absorptions rather than root uptake (7.86 %-24.2 %) under the exposure condition. In comparison with stomal absorption, wax layer penetration was the main pathway for most OPEs to enter into leaves, especially for those compounds with high octanol-air partition coefficients. Although the subcellular distributions of OPEs in the rice tissues of the foliar exposure were slightly different from those of the root exposure, hydrophobic OPEs were mainly stored in the cell wall with hydrophilic OPEs mainly in the cytosol. The translocation of OPEs from the exposed tissue to the unexposed tissue were significantly negatively correlated with their octanol-water partition coefficients, but their basipetal translocation were limited. The result suggested that the translocation of OPEs within rice is prioritized over their degradation. This study deepens our understanding of the processes behind OPE uptake by rice and highlights the importance of foliar uptake, especially for those via particle absorption.

Effects of atmospheric deposition on heavy metal contamination in paddy field systems under different functional areas in ChangZhuTan, Hunan Province, China.

In recent decades, the problem of heavy metal contamination in rice paddies has attracted widespread attention. However, most studies on heavy metal contamination in paddy fields are biased towards soil and/or rice plants, without taking atmospheric deposition into account. In this study, atmospheric deposition, paddy soil, and rice samples were collected from three functional areas (area proximity to factories, along the roadside, and suburban) in ChangZhuTan, Hunan Province. The pollution characterization, translocation, and health risk of heavy metals were reassessed. The findings revealed that Cd and As contamination in the study area's soils was more severe, with point exceedance rates reaching 70 % and 35.9 %, respectively. The highest concentrations of As, Ni, Cd, and Pb in atmospheric deposition were found along the roadside, with 1.42 µg/m2/day, 3.21 µg/m2/day, 0.34 µg/m2/day, and 8.28 µg/m2/day, respectively. In area proximity to factories, As and Ni in atmospheric deposition showed to be lowest, whereas Cd and Pb concentrations showed lowest in suburban areas. Furthermore, the accumulation of Cd and Pb in rice grains in regions proximity to factories was significantly higher than in other regions. The human health risk assessment indicated the health risk caused by rice intake in areas proximity to factories was the highest and requires attention, which was mainly due to Cd accumulation, with HQ value reached 3.19. Correlation tests indicate that atmospheric deposition has a positive effect on heavy metal enrichment in rice grains. Further Random Forest analysis revealed that the transport of heavy metals from atmospheric deposition to leaves and shells were important influencing factors for As, Cd, Ni and Mg accumulation in rice grain. Therefore, more attention should be paid to the effects of atmospheric deposition on the accumulation of heavy metals in paddy fields in order to maintain the production safety of crops.

Foliar uptake screening: A promising strategy for identifying wheat varieties with low lead accumulation.

Lead (Pb) contamination in wheat grain is of great concern, especially in North China. Atmospheric deposition is a major contributor to Pb accumulation in wheat grain. Screening low Pb accumulating wheat varieties has been an effective method for addressing Pb contamination in wheat grain. However, identifying wheat varieties with low Pb accumulation based on foliar uptake of atmospheric Pb has been neglected. Therefore, two field trials with distinct atmospheric Pb deposition were conducted to screen for stable varieties with low Pb accumulation. It was verified that YB700 and CH58, which have high thousand-grain weights and stable low Pb accumulation in field 1 (0.19 and 0.13 mg kg-1) and field 2 (0.17 and 0.20 mg kg-1), respectively, were recommended for cultivation in atmospheric Pb contaminated farmlands in North China. Furthermore, indoor experiments were conducted to investigate Pb uptake by the roots and leaves of different wheat varieties. Our findings indicate that Pb accumulation in different wheat varieties is primarily influenced by foliar Pb uptake rather than root Pb uptake. Interestingly, there was a positive correlation (p < 0.05) between the Pb concentrations in leaves and the stomatal width and trichome length of the adaxial epidermal surface. Additionally, there is a positive correlation (p < 0.01) between the Pb concentration in the wheat grain and trichome length. In conclusion, the screening of wheat varieties with narrower stomatal widths or shorter trichomes based on foliar uptake pathways is an effective strategy for ensuring food safety in areas contaminated by atmospheric Pb.

Uptake, accumulation and translocation mechanisms of organophosphate esters in cucumber (Cucumis sativus) following foliar exposure.

Organophosphate esters (OPEs) have been frequently detected in crops. However, few studies have focused on the uptake and translocation of OPEs in plants following foliar exposure. Herein, to investigate the foliar uptake, accumulation and translocation mechanisms of OPEs in plant, the cucumber (Cucumis sativus) was selected as a model plant for OPEs exposure via foliar application under control conditions. The results showed that the content of OPEs in the leaf cuticle was higher than that in the mesophyll on exposed leaf. Significant positive correlations were observed between the content of OPEs in the leaf cuticle and their log Kow and log Kcw values (P < 0.01), suggesting that OPEs with high hydrophobicity could not easily move from the cuticle to the mesophyll. The moderately hydrophobic OPEs, such as tris (2-chloroisopropyl) phosphate (TCPP, log Kow = 2.59), were more likely to move not only from the cuticle to the mesophyll but also from the mesophyll to the phloem. The majority of the transported OPEs accumulated in younger leaves (32-45 %), indicating that younger tissue was the primary target organ for OPEs accumulation after foliar exposure. Compared to chlorinated OPEs (except TCPP) and aryl OPEs, alkyl OPEs exhibited the strongest transport capacity in cucumber seedling due to their high hydrophilicity. Interestingly, tri-p-cresyl phosphate was found to be more prone to translocation compared to tri-m-cresyl phosphate and tri-o-cresyl phosphate, despite having same molecular weight and similar log Kow value. These results can contribute to our understanding of foliar uptake and translocation mechanism of OPEs by plant.