Foliar water uptake: a common water acquisition strategy for plants of the redwood forest

Evaluations of plant water use in ecosystems around the world reveal a shared capacity by many different species to absorb rain, dew, or fog water directly into their leaves or plant crowns. This mode of water uptake provides an important water subsidy that relieves foliar water stress. Our study provides the first comparative evaluation of foliar uptake capacity among the dominant plant taxa from the coast redwood ecosystem of California where crown-wetting events by summertime fog frequently occur during an otherwise drought-prone season. Previous research demonstrated that the dominant overstory tree species, Sequoia sempervirens, takes up fog water by both its roots (via drip from the crown to the soil) and directly through its leaf surfaces. The present study adds to these early findings and shows that 80% of the dominant species from the redwood forest exhibit this foliar uptake water acquisition strategy. The plants studied include canopy trees, understory ferns, and shrubs. Our results also show that foliar uptake provides direct hydration to leaves, increasing leaf water content by 2–11%. In addition, 60% of redwood forest species investigated demonstrate nocturnal stomatal conductance to water vapor. Such findings indicate that even species unable to absorb water directly into their foliage may still receive indirect benefits from nocturnal leaf wetting through suppressed transpiration. For these species, leaf-wetting events enhance the efficacy of nighttime re-equilibration with available soil water and therefore also increase pre-dawn leaf water potentials.     

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 (R² = 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.     

Fog in the California redwood forest: ecosystem inputs and use by plants

Fog has been viewed as an important source of moisture in many coastal ecosystems, yet its importance for the plants which inhabit these ecosystems is virtually unknown. Here, I report the results of a 3-year investigation of fog inputs and the use of fog water by plants inhabiting the heavily fog inundated coastal redwood (Sequoia sempervirens) forests of northern California. During the study period, 34%, on average, of the annual hydrologic input was from fog drip off the redwood trees themselves (interception input). When trees were absent, the average annual input from fog was only 17%, demonstrating that the trees significantly influence the magnitude of fog water input to the ecosystem. Stable hydrogen and oxygen isotope analyses of water from fog, rain, soil water, and xylem water extracted from the dominant plant species were used to characterize the water sources used by the plants. An isotopic mixing model was employed to then quantify how much fog water each plant used each month during the 3-year study. In summer, when fog was most frequent, ～19% of the water within S.sempervirens, and ～66% of the water within the understory plants came from fog after it had dripped from tree foliage into the soil; for S.sempervirens, this fog water input comprised 13–45% of its annual transpiration. For all plants, there was a significant reliance on fog as a water source, especially in summer when rainfall was absent. Dependence on fog as a moisture source was highest in the year when rainfall was lowest but fog inputs normal. Interestingly, during the mild El Niño year of 1993, when the ratio of rainfall to fog water input was significantly higher and fog inputs were lower, both the proportion and coefficient of variation in how much fog water was used by plants increased. An explanation for this is that while fog inputs were lower than normal in this El Niño year, they came at a time when plant demand for water was highest (summer). Therefore, proportional use of fog water by plants increased. The results presented suggest that fog, as a meteorological factor, plays an important role in the water relations of the plants and in the hydrology of the forest. These results demonstrate the importance of understanding the impacts of climatic factors and their oscillations on the biota. The results have important implications for ecologists, hydrologists, and forest managers interested in fog-inundated ecosystems and the plants which inhabit them.     

Polystichum munitum (Dryopteridaceae) varies geographically in its capacity to absorb fog water by foliar uptake within the redwood forest ecosystem

? Premise of the study: Fog provides a critical water resource to plants around the world. In the redwood forest ecosystem of northern California, plants depend on fog absorbed through foliar uptake to stay hydrated during the rainless summer. In this study, we identified regions within the redwood ecosystem where the fern Polystichum munitum canopy most effectively absorbs fog drip that reaches the forest floor. ? Methods: We measured the foliar uptake capacity of P. munitum fronds at seven sites along 700 km of the redwood forest ecosystem. We quantified the canopy cover of P. munitum at each site and estimated how much water the fern canopy can acquire aboveground through fog interception and absorption. ? Key results: Throughout the ecosystem, nocturnal foliar uptake increased the leaf water content of P. munitum by 7.2%, and we estimated that the P. munitum canopy can absorb 5 ± 3% (mean ± SE) of intercepted fog precipitation. Strikingly, P. munitum had the highest foliar uptake capacity in the center of the ecosystem and may absorb 10% more of the fog its canopy intercepts in this region relative to other regions studied. Conversely, P. munitum had no foliar uptake capacity in the southern end of the ecosystem. ? Conclusions: This study shows the first evidence that foliar uptake varies within species at the landscape scale. Our findings suggest that the P. munitum at the southern tip of the redwood ecosystem may suffer most from low summertime water availability because it had no potential to acquire fog as an aboveground water subsidy.     

Landscape scale variation in the hydrologic niche of California coast redwood

Topoclimatic diversity within forest landscapes can underlie variation in water availability, which may correspond to patterns in habitat suitability of tree species with differing hydrologic niches. However, the trade-off between the collection of data at a fine grain size over large spatial extents has limited comprehensive analyses of landscape scale variation in habitat suitability. We present a fine scale analysis of the roles of topographic gradients in moisture availability, soil water storage and fog frequency in the spatial pattern of habitat suitability for coast redwood Sequoia sempervirens, at 10 m resolution across 34 800 ha and three landscapes spanning approximately one-third of redwood's latitudinal range. A new 10 m resolution presence-absence map derived from airborne imaging spectroscopy was used to characterize current redwood distributions in three landscapes. Access to belowground moisture was assessed through four measures calculated from a high-resolution LiDAR digital elevation model, soil water storage was mapped from the USDA Soil Survey Geographic Database, and fog frequency was calculated from MODIS cloud cover data. Access to belowground moisture and fog predicted variation in suitability as redwood habitat from 22–75% and predicted redwood presence and absence at 10 m resolution with 63–74% accuracy. Across all three landscapes, redwood density consistently decreased with increasing interpolated height above a stream (IHAS), but the role of other predictors varied among the sites. Our results challenge previous assumptions that redwood habitat suitability within much coarser resolutions (800 m × 800 m) can be described by a single value. The elevated habitat suitability of sites close to streams suggests that these sites should be evaluated for their potential to become microrefugia within landscapes that may become climatically-unsuitable for redwoods at a regional scale.     

Shade-avoidance responses in two common coastal redwood forest species,Sequoia sempervirens (Taxodiaceae) and Satureja douglasii (Lamiaceae), occurring in various light quality environments

Shade-avoidance responses were examined for two species common to the coastal redwood forest, Sequoia sempervirens and Satureja douglasii. Sequoia seedlings demonstrated a shade-avoidance response when given end-of-day far-red light by increased hypocotyl, epicotyl, and first-node extension, and greater total number of needles and reduced anthocyanin concentration. Thus, Sequoia seedlings respond as sun-adapted plants. Satureja has several leaf monoterpene chemotypes that occur in different light environments including the redwood forest, and the types responded differently to the light treatments. The pulegone type responded to end-of-day far-red light as a sun-adapted plant with significant extension growth, increased leaf area and chlorophyll, and reduced anthocyanin. The isomenthone type responded as a shade-tolerant plant and did not exhibit extension growth nor a change in other parameters with end-of-day far-red light. However, the carvone and bicyclic types had variable responses depending on the parameter studied, which indicated genetic variation for these traits.     

Water relations of coast redwood planted in the semi-arid climate of southern California

Trees planted in urban landscapes in southern California are often exposed to an unusual combination of high atmospheric evaporative demand and moist soil conditions caused by irrigation. The water relations of species transplanted into these conditions are uncertain. We investigated the water relations of coast redwood (Sequoia sempervirens) planted in the urbanized semi-arid Los Angeles Basin, where it often experiences leaf chlorosis and senescence. We measured the sap flux (J(O)) and hydraulic properties of irrigated trees at three sites in the Los Angeles region. We observed relatively strong stomatal regulation in response to atmospheric vapour pressure deficit (D; J(O) saturated at D < 1 kPa), and a linear response of J(O) to photosynthetically active radiation. Total tree water use by coast redwood was relatively low, with plot-level transpiration rates below 1 mm d(-1) . There was some evidence of xylem cavitation during the summer, which appeared to be reversed in fall and early winter. We conclude that water stress was not a direct factor in causing leaf chlorosis and senescence as has been proposed. Instead, the relatively strong stomatal control that is adaptive in the native habitat of coast redwood may lead to carbon limitation and other stresses in semi-arid, irrigated habitats.     

Seasonal and diurnal patterns of photosynthetic gas exchange for Lonicera sempervirens and L. Japonica (Caprifoliaceae)

Photosynthesis, stomatal conductance, and water use efficiency were compared between Lonicera japonica (Japanese honeysuckle), an invasive species in the southeastern United States, and its native congener, Lonicera sempervirens (coral honeysuckle), to determine the role of seasonal patterns of water loss and carbon gain in the invasive ability of the exotic. Diurnal measurements were taken monthly for 1 year under a closed forest canopy and in an open field. There were few significant differences in photosynthetic rates between the two species in either environment. However, at both sites, Lonicera japonica retained its old leaves over winter while old leaves of L. sempervirens senesced. Also, new leaves of L. japonica had significantly higher photosynthetic rates than the emerging leaves of L. sempervirens (6.2 vs. 4.4 μmol m ²sec^-1 under the canopy; 4.4 vs. 3.0 μmol m^-2 sec^-1 in the open). Although differences in conductance and water use efficiency between species were seldom significant, L. japonica tended to have higher maximum values than L. sempervirens. Retention of old leaves by L. japonica during new leaf formation (January–March) as well as higher photosynthetic rates in new leaves contribute to greater annual carbon gain and help explain the invasive ability of Japanese honeysuckle.     

Influence of leaf size,orientation, and arrangement on temperature and transpiration in three high-elevation,large-leafed herbs

Summary The temperature and water relations of the largleafed, high-elevation species Frasera speciosa, Balsamorhiza sagittata, and Rumex densiflorus were evaluated in the Medicine Bow Mountains of southeast Wyoming (USA) to determine the influence of leaf size, orientation, and arrangement on transpiration. These species characteristically have low minimum stomatal resistances (<60 s m^-1) and high maximum transpiration rates (>260 mg m^-2s^-1 for F. speciosa). Field measurements of leaf and microclimatic parameters were incorporated into a computer simulation using standard energy balance equations which predicted leaf temperature (T _leaf) and transpiration for various leaf sizes. Whole-plant transpiration during a day was simulated using field measurements for plants with natural leaf sizes and compared to transpiration rates simulated for plants having identical, but hypothetically smaller (0.5 cm) leaves during a clear day and a typically cloudy day. Although clear-day transpiration for F. speciosa plants with natural size leaves was only 2.0% less per unit leaf area than that predicted for plants with much smaller leaves, daily transpiration of B. sagittata and R. densiflorus plants with natural leaf sizes was 16.1% and 21.1% less, respectively. The predicted influence of a larger leaf size on transpiration for the cloudy day was similar to clear-day results except that F. speciosa had much greater decreases in transpiration (12.7%). The different influences of leaf size on transpiration between the three species was primarily due to major differences in leaf absorptance to solar radiation, orientation, and arrangement which caused large differences in T _leaf. Also, simulated increases in leaf size above natural sizes measured in the field resulted in only small additional decreases in predicted transpiration, indicating a leaf size that was nearly optimal for reducing transpiration. These results are discussed in terms of the possible evolution of a larger leaf size in combination with specific leaf absorptances, orientations and arrangements which could act to reduce transpiration for species growing in short-season habitats where the requirement for rapid carbon fixation might necessitate low stomatal resistances.     

MICROCLIMATIC EFFECTS ON WATER RELATIONS,LEAF TEMPERATURES,AND THE DISTRIBUTION OF HERACLEUM LANATUM AT HIGH ELEVATIONS

The small-scale distribution of an understory herb, Heracleum lanatum, was evaluated in terms of leaf temperature and water relations limitations due to a large leaf size (630 cm²). Diurnal variations in transpiration (4 to 60 mg m⁻² s⁻¹) were influenced by fluctuations in solar irradiance, wind speed, leaf temperature and stomatal conductance. Computer simulations indicated that leaf temperatures in a forest clearing would be > 12 C above air temperature, with maximum transpiration rates of 140 mg m⁻² s⁻¹, and daily water loss to be over 200% greater than values at natural understory locations. Simulations of nocturnal temperature relations indicated ~100 W m ⁻² less incident longwave irradiance in the forest clearing as compared to the understory (560 vs. 660 W m⁻² at 400 hr). This difference led to predicted leaf temperatures being as low as 6 C below air temperature in the forest clearing while measured leaf temperatures in the forest understory were within 1.5 C of air temperature throughout the night. Furthermore, minimum air temperatures were at or below 6 C on 36% of the nights during the summer growth period indicating that in open areas leaves of H. lanatum would frequently be below 0 C and subject to possible freeze damage. Heracleum lanatum may be more abundant in the shaded understory of the subalpine forest because exposure in open environments would result in high leaf temperatures and increased transpirational water loss during the day, as well as low leaf temperatures with the possibility of freeze damage at night.     

Relationships between vessel element anatomy and physiological as well as morphological traits of leaves in <Emphasis Type="Italic">Fagus crenata</Emphasis> seedlings originating from different provenances

We attempted to observe differences in vessel element anatomy, and physiological and morphological traits of leaves in Fagus crenata seedlings originated from seven different provenances grown under the uniform environmental conditions. We also investigated the relationships between the anatomical characteristics of the vessel elements and physiological plus morphological traits of leaves in Fagus crenata seedlings. To carry out the experiments, Fagus crenata samples were prepared from Chichibu Research Forest of Tokyo University. For anatomical studies of the vessel elements, vessel number per mm², average vessel area, and the percentage of vessel area from the pith to the bark side were measured. We also measured transpiration rate, stomatal conductance, leaf area, leaf thickness, leaf dry mass per unit leaf area and leaf density for foliar studies. The anatomical characteristics of the vessel elements as well as the physiological and morphological traits of leaves were noticeably different among provenances. In addition, we found significant correlations between the foliar characteristics not only with vessel number per mm², but also with vessel area percentage and sum of the (vessel area)², theoretical hydraulic conductivity, from the pith to the bark side. Therefore, we concluded that variations in physiological and morphological traits of leaves in response to provenance variation were in correspondence with vessel number per mm² as a parameter that determines the total vessel area and consequent water hydraulic conductance in Fagus crenata as a diffuse porous hardwood.     

The role of marginal hydathodes in foliar water absorption

Summary

Marginal epithem hydathodes may, in some conditions, absorb water. A toothed leaf margin can be regarded as a water-holding structure. It is suggested that the lack of hydathodes in north-temperate evergreens and in the leaves of rain forest trees is related to the need for such species to reduce foliar water absorption. The high proportion of deciduous species with hydathodes in woodland vegetation is thought to be related to the summer potential soil water deficit in Britain, leaf-absorbed water supplementing water from the soil. The term ‘potomorphic’ is suggested for leaves with water absorbing structures.     

Foliar water uptake: Processes,pathways, and integration into plant water budgets

Nearly all plant families, represented across most major biomes, absorb water directly through their leaves. This phenomenon is commonly referred to as foliar water uptake. Recent studies have suggested that foliar water uptake provides a significant water subsidy that can influence both plant water and carbon balance across multiple spatial and temporal scales. Despite this, our mechanistic understanding of when, where, how, and to what end water is absorbed through leaf surfaces remains limited. We first review the evidence for the biophysical conditions necessary for foliar water uptake to occur, focusing on the plant and atmospheric water potentials necessary to create a gradient for water flow. We then consider the different pathways for uptake, as well as the potential fates of the water once inside the leaf. Given that one fate of water from foliar uptake is to increase leaf water potentials and contribute to the demands of transpiration, we also provide a quantitative synthesis of observed rates of change in leaf water potential and total fluxes of water into the leaf. Finally, we identify critical research themes that should be addressed to effectively incorporate foliar water uptake into traditional frameworks of plant water movement.     

Photosynthesis and water-relation traits of the summer annual C4 grasses, Eleusine indica and Digitaria adscendens, with contrasting trampling tolerance

Two summer annual C₄ grasses with different trampling susceptibilities were grown as potted plants, and diurnal leaf gas exchange and leaf water potential in each grass were compared. The maximum net photosynthetic rate, leaf conductance and transpiration rate were higher in the trampling-tolerant Eleusine indica (L.) Gaertn. than in trampling sensitive Digitaria adscendens (H. B. K.) Henr. Leaf water potential was much lower in E. indica than in D. adscendens. There were no differences in soil-to-leaf hydraulic conductance and leaf osmotic potential at full turgor as obtained by pressure–volume analysis. However, the bulk modulus of elasticity in cell walls was higher in E. indica leaves than in D. adscendens leaves. This shows that the leaves of E. indica are less elastic. Therefore, the rigid cell walls of E. indica leaves reduced leaf water potential rapidly by decreasing the leaf water content, supporting a high transpiration rate with high leaf conductance. In trampled habitats, such lowering of leaf water potential in E. indica might play a role in water absorption from the compacted soil. In contrast, the ability of D. adscendens to colonize dry habitats such as coastal sand dunes appears to be due to its lower transpiration rate and its higher leaf water potential which is not strongly affected by decreasing leaf water content.     

Transpiration and Calcium Deposition by Unifoliate Leaves of Phaseolus vulgaris Differing in Maturity

Unifoliate leaves were individually enclosed in clear, plastic chambers for the 24 hour treatment periods and then sacrificed for Ca analysis. Two transpiration rates were obtained by passing dry air through the chambers tising flow rates of 160 and 260 cm³/min. A third rate was obtained by a combination of shade and the lower air flow rate. Neither the transpiration rate nor solution-Ca concentration (0.5mM and 2.5 mM of 0.1, and 0.5 strength Hoagland solution) altered the amount of Ca deposited in the unifoliate leaves of 22 day old bean plants (Phaseolus vulgaris). The transpiration rate per unit area of leaf remained constant for all ages studied (1l–20 days) and was 1.8, 2.7, 3.6 g H₂O per dm² day for the three different imposed conditions. A definite pattern of Ca deposition occurred. With all the transpiration rates there was a maximum rate of calcium deposition at 13 days of growth and a gradual decrease thereafter. When the Ca concentration of the nutrient solution was 20 μg/ml the daily Ca deposition in terms of water transpired by the unifoliate leaves exceeded this amount, except for the oldest leaf tested, and, the maximum Ca to water ratios were 250, 320, and 430 (μg Ca/g) in order of decreasing transpiration rates. The uptake of Ca against a concentration gradient and approximately the same total uptake regardless of transpiration rates and solution concentrations used, firmly suggest that Ca secretion into root-xylem elements from a surrounding low level Ca solution requires energy expenditure by the plant. A possible explanation was proposed for the decreased rate of Ca deposition by the unifoliate leaves subsequent to the 13th day.     

Two Steps of Gas Exchange in Leaf Photosynthesis

Photosynthesis and transpiration of excised leaves of Taraxacum officinale L. and a few other species of plants were measured, using an open gas analysis system. The rates of CO₂ uptake and transpiration increased in two steps upon illumination of stomata-bearing epidermis of these leaves at a light intensity of 50 mW × cm⁻². Abscisic acid inhibited only the second step of gas exchange. Illumination of the astomatous epidermis of hypostomatous leaves caused only the first step of gas exchange. These data indicate that the first and second steps arise from cuticular and stomatal gas exchange, respectively. The rate of the cuticular photosynthesis in a Taraxacum leaf reached saturation at a light intensity of 5 mW × cm⁻², and the rates of the stomatal photosynthesis and transpiration reached saturation at a higher intensity of 35 mW × cm⁻². The cuticular photosynthesis of a Taraxacum leaf was 18% of the stomatal photosynthesis at 50 mW × cm⁻² and 270% at 5 mW × cm⁻². The other species of leaves showed the same trend. The importance of cuticular CO₂ uptake in leaf photosynthesis, especially under low light intensity was stressed from these data.     

Taxodioxylon gypsaceum (Göppert) Kräusel,legno fossile rinvenuto nel miocene dell'Italia centrale

Abstract

Taxodioxylon gypsaceum (Göppert) Kräusel, wood fossil discovered in the central italian miocene.—A sample of silicified wood fossil has been discovered in the miocenic formations of the Castellaro mountain in the neighbourhood of Pesaro a city in Central Italy. An anatomical study has made it possible to accept that the species is Taxodioxylon gypsaceum, which resembles the actual Sequoia sempervirens for its woody structure, conservative endemism of a coastal region in California. The fossil species that is known for its numerous tertiary deposits in the Northern, Central and Eastern Europe, is now indicated for the first time in Italy; the mount Castellaro station results to be the more south among those in Europe.

Various paleobotanical data concerning traces of leaves, fruits and seeds gathered in localities near to the deposit and in the region complies with the discovery of Taxodioxylon gypsaceum because of the presence of varions species of Sequoia and Taxodium and in particular of the Sequoia langsdorfii leaves which is considered as the ancient form of the Sequoia sempervirens.     

Effect of Abscisic Acid on the Transpiration of Zea mays

Intact plants of Zea mays L. were treated with foliar sprays of cis-trans-abscisic acid (ABA) at concentrations from 10⁻⁹ to 10⁻⁴M. Even the lowest concentration caused a reduction of the transpiration rate as measured between 1 and 33 h after spraying. With increasing ABA concentrations, there was a nearly linear relationship between the logarithm of the ABA concentration and the (decreasing) transpiration rate within that period. Subsequently a partial recovery of the transpiration rate set in, beginning progressively later as the ABA concentration was increased. After 5 1/2 days the transpiration rate of plants treated with 10⁻⁹ and 10⁻⁸M was nearly back to normal, whereas plants treated with 10⁻⁴M transpiration at only about 2/3 their normal rate. In experiments with detached maize leaves supplied with water or ABA solutions (10⁻⁸ to 10⁻⁵M) through their cut bases, the transpiration of control leaves decreased gradually to a low level in 24 h. ABA caused a marked and rapid reduction of the transpiration rate compared to that of the controls. After a few hours, the transpiration of the treated leaves decreased at a slower rate than that of the controls, thus approaching the control values. After 35 h, the transpiration of leaves treated with 10⁻⁵M ABA was nearly the same as in untreated leaves. Exchanging the ABA solution for distilled water after 24 h had little effect on the subsequent course of the transpiration rate.     

Water balance in the arborescent palm, Sabal palmetto. I. Stem structure, tissue water release properties and leaf epidermal conductance

The functional importance of water storage in the arborescent palm, Sabal palmetto, was investigated by observing aboveground water content, pressure-volume curve parameters of leaf and stem tissue and leaf epidermal conductance rates. The ratio of the amount of water stored within the stem to the leaf area (kg m^?2) increased linearly with plant height. Pressure-volume curves for leaf and stem parenchyma differed markedly; leaves lost turgor at 0.90 relative water content and –3.81 MPa, while the turgor loss point for stem parenchyma occurred at 0–64 relative water content and ?0.96 MPa. Specific capacitance (change in relative water content per change in tissue water potential) of stem parenchyma tissue was 84 times higher than that of leaves, while the bulk modulus of elasticity was 346 times lower. Leaf epidermal conductance rates were extremely low (0.32–0.56 mmol m^?2 s^?1) suggesting that S. palmetto are able to strongly restrict foliar water loss rates. Structurally, stems of S. palmetto appear to be well suited to act as a water storage reservoir, and coupled with the ability to restrict water loss from leaf surfaces, may play an important role in tree survival during periods of low water availability.