Trichomes are common in plants from dry environments, and despite their recognized role in protection and defense, little is known about their role as absorptive structures and in other aspects of leaf ecophysiology. We combine anatomical and ecophysiological data to evaluate how trichomes affect leaf gas exchange and water balance during drought. We studied two congeneric species with pubescent leaves which co-occur in Brazilian Caatinga: Croton blanchetianus (dense trichomes) and Croton adenocalyx (sparse trichomes). We found a novel foliar water uptake (FWU) pathway in C. blanchetianus composed of stellate trichomes and underlying epidermal cells and sclereids that interconnect the trichomes from both leaf surfaces. The water absorbed by these trichomes is redistributed laterally by pectin protuberances on mesophyll cell walls. This mechanism enables C. blanchetianus leaves to absorb water more efficiently than C. adenocalyx. Consequently, the exposure of C. blanchetianus to dew during drought improved its leaf gas exchange and water status more than C. adenocalyx. C. blanchetianus trichomes also increase their leaf capacity to reflect light and maintain lower temperatures during drought. Our results emphasize the multiple roles that trichomes might have on plant functioning and the importance of FWU for the ecophysiology of Caatinga plants during drought.

Hydathodes are usually associated with water exudation in plants. However, foliar water uptake (FWU) through the hydathodes has long been suspected in the leaf-succulent genus Crassula (Crassulaceae), a highly diverse group in southern Africa, and, to our knowledge, no empirical observations exist in the literature that unequivocally link FWU to hydathodes in this genus. FWU is expected to be particularly beneficial on the arid western side of southern Africa, where up to 50% of Crassula species occur and where periodically high air humidity leads to fog and/or dew formation. To investigate if hydathode-mediated FWU is operational in different Crassula species, we used the apoplastic fluorescent tracer Lucifer Yellow in combination with different imaging techniques. Our images of dye-treated leaves confirm that hydathode-mediated FWU does indeed occur in Crassula and that it might be widespread across the genus. Hydathodes in Crassula serve as moisture-harvesting structures, besides their more common purpose of guttation, an adaptation that has likely played an important role in the evolutionary history of the genus. Our observations suggest that ability for FWU is independent of geographical distribution and not restricted to arid environments under fog influence, as FWU is also operational in Crassula species from the rather humid eastern side of southern Africa. Our observations point towards no apparent link between FWU ability and overall leaf surface wettability in Crassula. Instead, the hierarchically sculptured leaf surfaces of several Crassula species may facilitate FWU due to hydrophilic leaf surface microdomains, even in seemingly hydrophobic species. Overall, these results confirm the ecophysiological relevance of hydathode-mediated FWU in Crassula and reassert the importance of atmospheric humidity for some arid-adapted plant groups.

Analysis of functional traits is a cornerstone of ecology, yet individual traits seldom explain useful amounts of variation in species distribution or climatic tolerance, and their functional significance is rarely validated experimentally. Multivariate suites of interacting traits could build an understanding of ecological processes and improve our ability to make sound predictions of species success in our rapidly changing world. We use foliar water uptake capacity as a case study because it is increasingly considered to be a key functional trait in plant ecology due to its importance for stress-tolerance physiology. However, the traits behind the trait, that is, the features of leaves that determine variation in foliar water uptake rates, have not been assembled into a widely applicable framework for uptake prediction. Focusing on trees, we investigated relationships among 25 structural traits, leaf osmotic potential (a source of free energy to draw water into leaves), and foliar water uptake in 10 diverse angiosperm and conifer species. We identified consistent, multitrait \"uptake syndromes\" for both angiosperm and conifer trees, with differences in key traits revealing suspected differences in the water entry route between these two clades and an evolutionarily significant divergence in the function of homologous structures. A literature review of uptake-associated functional traits, which largely documents similar univariate relationships, provides additional support for our proposed \"uptake syndrome.\" Importantly, more than half of shared traits had opposite-direction influences on the capacity of leaves to absorb water in angiosperms and conifers. Taxonomically targeted multivariate trait syndromes provide a useful tool for trait selection in ecological research, while highlighting the importance of micro-traits and the physiological verification of their function for advancing trait-based ecology.

In arid and semi-arid areas, plants can directly absorb and use dew through their leaves, and some plants have the ability for hydraulic redistribution of their roots. Therefore, in arid areas, plants may redistribute dew to the soil, using the soil as a reservoir for short-term dry seasons, i.e. dew may participate in the hydraulic redistribution process of plants. This process plays an important role in plant survival and community stability.
To verify this hypothesis, we investigated the water use mechanism of Populus euphratica through a comprehensive observation of sap flow, water potential and soil water content using a heavy water tracer experiment under in situ field conditions.
Dewdrops contributed 28.3 % of soil moisture near the roots, and applying dew on leaves for several days significantly improved soil moisture status. Hydraulic redistribution in the roots mainly occurred from 2200 h at night to 800 h the following day and mainly occurred in the 20- to 80-cm soil layer. Water storage in the trunk is the intermediate link in the coupling process of foliar water uptake and hydraulic redistribution; water storage in the trunk is mainly replenished from May to July and consumed throughout the rest of the year. In conclusion, dew redistributes water into soil through the coupling process of foliar water uptake and hydraulic redistribution. Populus euphratica uses the trunk and soil for water storage to cope with water stress during short-term drought periods. Our findings provide a scientific basis for the restoration of different species in water-deficient areas, which is conducive to maintaining vegetation ecosystem stability in areas of desertification and improving the soil water balance.

Trichomes are epidermal outgrowths on plant shoots. Their roles in protecting plants against herbivores and in the biosynthesis of specialized metabolites have long been recognized. Recently, studies are increasingly showing that trichomes also play important roles in water absorption and metal detoxication, with these roles having important implications for ecology, the environment, and agriculture. However, these two functions of trichomes have been largely overlooked and much remains unknown. In this review, we show that the trichomes of 37 plant species belonging to 14 plant families are involved in water absorption, while the trichomes of 33 species from 13 families are capable of sequestering metals within their trichomes. The ability of trichomes to absorb water results from their decreased hydrophobicity compared to the remainder of the leaf surface as well as the presence of special structures for collecting and absorbing water. In contrast, the metal detoxication function of trichomes results not only from the good connection of their basal cells to the underlying vascular tissues, but also from the presence of metal-chelating ligands and transporters within the trichomes themselves. Knowledge gaps and critical future research questions regarding these two trichome functions are highlighted. This review improves our understanding on trichomes.

Foliar water uptake (FWU) has been documented in many species and is increasingly recognized as a non-trivial factor in plant-water relationships. However, it remains unknown whether FWU is a widespread phenomenon in Pinus species, and how it may relate to needle traits such as the form and structure of stomatal wax plugs. In this contribution, these questions were addressed by studying FWU in current-year and 1-year-old needles of seven Pinus species.
We monitored FWU gravimetrically and analysed the needle surface via cryo-scanning electron microscopy. Additionally, we considered the effect of artificial wax erosion by application of the surfactant Triton X-100, which is able to alter wax crystals.
The results show for all species that (1) FWU occurred, (2) FWU is higher in old needles compared to young needles and (3) there is substantial erosion of stomatal wax plugs in old needles. FWU was highest in Pinus canariensis, which has a thin stomatal wax plug. Surfactant treatment enhanced FWU.
The results of this study provide evidence for (1) widespread FWU in Pinus, (2) the influence of stomatal wax plugs on FWU and (3) age-related needle surface erosion.

Embolism refilling is thought to require relaxation of xylem tension, and it is unclear whether and how tall trees or plants growing in arid or saline soils recover from embolism. We tested whether foliar water uptake could enable embolism refilling in dehydrated twigs of the grey mangrove (Avicennia marina). Four dehydrated twigs were imaged by laboratory-based micro-computed tomography before and after wetting leaves. Emboli were observed in dehydrated stems and leaves. Embolism decreased with increasing distance from the cut end of stems, suggesting that stem emboli were caused by cutting. A significant (P = 0.026) c. 80% reduction in the embolised area was observed in leaves between the start and the end of the experiment (29 ± 10 h after wetting). Embolus diameter was unaffected by wetting. Embolism refilling occurred slowly, in stems embolised by cutting and leaves embolised by cutting and/or dehydration. The lack of response of embolus diameter to wetting suggests that capillarity was not the main mechanism for refilling. Results show that excised twigs of A. marina are able to recover from embolism by absorption of atmospheric water and call for studies under natural conditions.

Absorption of water across the surfaces of leaves is an ecologically important aspect of tree physiology. Variation in foliar water uptake capacity depends on environmental conditions when traits associated with the uptake pathway respond to climatic signals. Using a series of experiments, we verify that water enters Sequoia sempervirens (D. Don) Endl. leaves by crossing the cuticle, and show that surface-trait acclimation alters the kinetic parameters of foliar water uptake. Under our experimental conditions, the cuticle was the primary pathway for water entry into the leaf. Exposure to climatic variation may induce surface acclimations, such as increased waxiness, that reduce water-film formation over stomata at the expense of dry-season foliar uptake rates. We found that water uptake is negatively associated with the interaction of leaf-surface wax coverage and stomatal density, and provide an accessible protocol to measure these key traits in Sequoia. Linking absorptive pathways and trait acclimation to physiological performance can provide a foundation for range-wide or genomic investigations of forest interactions with water and a mechanism-centered means to monitor canopy hydraulic parameters over time.

Tracheid buckling may protect leaves in the dynamic environments of forest canopies, where rapid intensifications of evaporative demand, such as those brought on by changes in light availability, can result in sudden increases in transpiration rate. While treetop leaves function in reliably direct light, leaves below the upper crown must tolerate rapid, thermally driven increases in evaporative demand. Using synchrotron-based X-ray microtomography, we visualized impacts of experimentally induced water stress and subsequent fogging on living cells in redwood leaves, adding ecological and functional context through crown-wide explorations of variation in leaf physiology and microclimate. Under drought, leaf transfusion tracheids buckle, releasing water that supplies sufficient temporal reserves for leaves to reduce stomatal conductance safely while stopping the further rise of tension. Tracheid buckling fraction decreases with height and is closely coordinated with transfusion tissue capacity and stomatal conductance to provide temporal reserves optimized for local variation in microclimate. Foliar water uptake fully restores collapsed and air-filled transfusion tracheids in leaves on excised shoots, suggesting that trees may use aerial water sources for recovery. In the intensely variable deep-crown environment, foliar water uptake can allow for repetitive cycles of tracheid buckling and unbuckling, protecting the tree from damaging levels of hydraulic tension and supporting leaf survival.

The ecohydrological significance of leaf wetting due to atmospheric water in arid and semiarid ecosystems is not well understood. In these environments, the inputs of precipitation or dew formation resulting in leaf wetting have positive effects on plant functioning. However, its impact on plant water relations may depend on the degree of leaf surface wettability. In this study we evaluated leaf wettability and other leaf traits and its effects on foliar water uptake and canopy interception in plant species of a Patagonian steppe. We also studied how leaf traits affecting wettability vary seasonally from growing to dry season.
Contact angle of a water droplet with the leaf surface, water adhesion, droplet retention angle, stomatal density, cuticular conductance, canopy interception and maximum foliar water uptake were determined in six dominant shrub species.
All species increased leaf wettability during the dry season and most species were considered highly wettable. The leaf surface had very high capacity to store and retain water. We found a negative correlation between foliar water uptake and leaf hydrophilia.
Despite the diversity of life forms, including cushion shrubs and tall shrubs, as well as phenological variability, all species converged in similar seasonal changes in leaf traits that favour wettability. Intercepted water by crowns and the extremely high capacity of retention of droplets on leaf surfaces can have a significant impact on eco-hydrological process in water limited ecosystems where most of water sources during the growing and the dry season may be small rainfall events or dew, which do not always increase soil water availability.