Research progress on water uptake through foliage

Leaf is one of the most sensitive organs that response to atmospheric environment changes. In arid region, especially under water stress conditions, water from foliar uptake is one of the main water subsidy sources, which plays an important role in plant survival, growth and ecosystem sustainable development. Foliar water uptake attracts more attention in physiological and ecological research all over the world. This essay summarized the main measurement methods of foliar water uptake at home and abroad in detail: vessel control method, stable isotope technology and heat ratio method. And analyzed the significance of foliar water uptake from physiological and ecological aspects; and discussed the influence factors of foliar water uptake from water condition perspective, water absorption and leaf structure aspects. The following aspects should be paid more attention in future researches: (1) measure the amount of foliar water uptake by using stable isotope technology and heat ratio method; (2) the relationship among foliar uptake of fog, dew, rain and water vapor in arid region; (3) the foliar uptake amount and rate should be simulated by models; (4) the mechanisms of foliar water uptake should be researched; (5) study foliar water uptake of different plants at ecosystem scale.     

Research progress on water uptake through foliage

Leaf is one of the most sensitive organs that response to atmospheric environment changes. In arid region, especially under water stress conditions, water from foliar uptake is one of the main water subsidy sources, which plays an important role in plant survival, growth and ecosystem sustainable development. Foliar water uptake attracts more attention in physiological and ecological research all over the world. This essay summarized the main measurement methods of foliar water uptake at home and abroad in detail: vessel control method, stable isotope technology and heat ratio method. And analyzed the significance of foliar water uptake from physiological and ecological aspects; and discussed the influence factors of foliar water uptake from water condition perspective, water absorption and leaf structure aspects. The following aspects should be paid more attention in future researches: (1) measure the amount of foliar water uptake by using stable isotope technology and heat ratio method; (2) the relationship among foliar uptake of fog, dew, rain and water vapor in arid region; (3) the foliar uptake amount and rate should be simulated by models; (4) the mechanisms of foliar water uptake should be researched; (5) study foliar water uptake of different plants at ecosystem scale.     

中国东部南北样带森林优势植物叶片的水分利用效率和氮素利用效率

通过测定中国东部南北样带主要森林生态系统中10种优势植物(兴安落叶松、蒙古栎、水曲柳、紫椴、色木槭、红松、杉木、木荷、马尾松、锥栗)叶片的碳氮含量(Cmass、Nmass)、同位素丰度(δ13C、δ15N)以及光合响应曲线,分析了不同优势植物叶片的水分利用效率和氮素利用效率之间的差异及其相互关系.结果表明： 不同生活型植物叶片的Nmass和δ15N差异显著,表现为阔叶植物>针叶植物,落叶植物>常绿植物;最大光合速率(Pn max)表现为针叶植物>阔叶植物,落叶植物>常绿植物;植物叶片的瞬时水分利用效率(WUEi)和长期水分利用效率(WUE)均表现为阔叶植物>针叶植物,常绿植物>落叶植物;植物叶片的瞬时氮素利用效率(NUEi)和长期氮素利用效率(NUE)则表现出相反的规律,且常绿植物和落叶植物叶片的NUE差异显著;WUEi和WUE之间相关性不显著,而NUEi和NUE之间呈显著正相关.植物叶片的水分利用效率与氮素利用效率显著负相关.两种资源利用效率均受植物生活型的影响,并且存在一定的制约关系.     

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.     

雪被覆盖对高山森林凋落物分解过程中水溶性和有机溶性组分含量的影响

季节性雪被可能通过冻结、淋溶以及冻融循环等对高山森林凋落物水溶性和有机溶性组分含量产生影响.本文采用凋落物分解袋法,以川西高山森林典型乔木（四川红杉、岷江冷杉、红桦、方枝柏）和灌木（高山杜鹃、康定柳）凋落物为研究对象,研究了雪被覆盖不同时期（雪被形成期、雪被覆盖期和雪被融化期）和雪被厚度（厚型雪被、中型雪被、薄型雪被和无雪被）下凋落物水溶性和有机溶性组分含量的动态变化特征.结果表明： 在一个冬季的分解过程中,6种凋落物水溶性组分含量在雪被形成期和融化期降低而雪被覆盖期增加,但除高山杜鹃凋落物有机溶性组分含量在雪被覆盖期增加外,其他5种凋落物有机溶性组分含量在整个冬季呈降低趋势.相对于凋落物有机溶性组分含量,不同厚度雪被斑块对凋落物水溶性组分含量变化的影响更大,且主要表现在雪被形成期和雪被覆盖期.相对于其他雪被斑块,薄型雪被斑块更加显著地促进了高山柳和高山杜鹃凋落物水溶性组分含量降低,但显著抑制了方枝柏凋落物水溶性组分含量降低,而其他凋落物水溶性组分含量变化在不同斑块间无显著差异.冬季高山森林雪被对凋落物水溶性和有机溶性组分含量的影响主要受控于凋落物质量.     

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.     

北京西山侧柏林冠层不同高度处叶片水分利用效率

以北京西山广泛分布的侧柏林为研究对象,综合考虑冠层不同高度处气象因子、大气CO₂浓度以及大气CO₂中碳同位素组成的差异,对其冠层不同高度处叶片的瞬时水分利用效率和短期水分利用效率分别进行了测定,以期为区域森林生态系统固碳与耗水研究提供理论依据,为区域森林生态系统经营与维护提供技术支撑.结果表明: 侧柏林冠层不同高度处叶片的瞬时水分利用效率随冠层高度的变化规律表现为上层>中层>下层,多种气象因子协同影响气孔运动,使瞬时水分利用效率受气孔限制;侧柏林冠层不同高度处的环境因子、大气CO₂浓度以及大气CO₂的δ13C均存在一定差异,导致了林冠各层叶片的短期水分利用效率的变化.林冠上层叶片通过提高水分利用效率适应环境.     

沙漠腹地天然绿洲胡杨和柽柳叶片δ13C值对不同地下水埋深的响应

干旱区植物的水分利用效率对植物的分布及水分利用状况具有重要意义。基于不同地下水埋深条件下沙漠腹地绿洲优势种胡杨和柽柳叶片δ¹³C值,分析了胡杨和柽柳的水分利用效率对不同地下水埋深的响应。结果表明: 随着地下水埋深由2.1 m增加到4.3 m,柽柳叶片的δ¹³C值先略有增加后处于较为稳定状态,柽柳采取较为稳定的水分利用效率适应干旱环境;胡杨叶片的δ¹³C值呈现先略有减小后增加的趋势,胡杨通过提高水分利用效率的策略适应干旱胁迫。同一地下水埋深条件下柽柳叶片的δ¹³C值高于胡杨叶片,表明柽柳的水分利用效率高于胡杨。     

Sapling growth as a function of light and landscape-level variation in soil water and foliar nitrogen in northern Michigan

Interspecific differences in sapling growth responses to soil resources could influence species distributions across soil resource gradients. I calibrated models of radial growth as a function of light intensity and landscape-level variation in soil water and foliar N for saplings of four canopy tree species, which differ in adult distributions across soil resource gradients. Model formulations, characterizing different resource effects and modes of influencing growth, were compared based on relative empirical support using Akaike’s Information Criterion. Contrary to expectation, the radial growth of species associated with lower fertility (Acer rubrum and Quercus rubra) was more sensitive to variation in soil resources than the high fertility species Acer saccharum. Moreover, there was no species tradeoff between growth under high foliar N versus growth under low foliar N, which would be expected if growth responses to foliar N mediated distributions. In general, there was functional consistency among species in growth responses to light, foliar N, and soil water availability, respectively. Foliar N influenced primarily high-light growth in F. grandifolia, A. rubrum, and Q. rubra (but was not significant for A. saccharum). In A. saccharum and A. rubrum, for which soil water availability was a significant predictor, soil water and light availability simultaneously limited growth (i.e., either higher light or water increased growth). Simple resource-based models explained 0.74–0.90 of growth variance, indicating a high degree of determinism. Results suggest that nitrogen effects on forest dynamics would be strongest in high-light early successional communities but that water availability influences growth in both early successional and understory environments.     

Boron foliar supplementation as a strategy to attenuate drought stress in soybean

Drought is the major abiotic stress that limits growth, development and yield of crops worldwide. In this scenario, mineral nutrients, such as boron (B), have been promising for increasing the tolerance of plants to abiotic stresses because of their physiological roles in plants. We aimed to evaluate the benefits of foliar supplementation of B in either relieving or reducing the physiological damages caused by water stress in soybean (Glycine max L.). A greenhouse trial was carried out in a 2 × 3 factorial scheme, with two water conditions (well-watered and drought stress) and three doses of B (0, 150 and 300 mg B L⁻¹). Foliar application of B was before inducing the water stress, which remained for 20 days during vegetative stage. Plants were evaluated at three moments, according to the water conditions: maximum stress, rehydration and just before harvesting. The levels of hydrogen peroxide and lipid peroxidation increased in soybean leaves and roots under water stress, resulting in impaired plant growth. However, the foliar supplementation with B before the stress increased activities of the antioxidant enzymes and reduced the levels of stress markers. Furthermore, B applied foliar increased the nutrient concentration in the leaves of plants and stimulated root growth, which resulted in higher harvest index related to yield. The foliar application of B has shown as an alternative management to mitigate the damages caused by drought stress in soybean.     

Experimental cloud immersion and foliar water uptake in saplings of Abies fraseri and Picea rubens

Key message

Frequent cloud immersion events result in direct uptake of cloud water and improve plant water potentials during daylight hours in saplings of two dominant cloud forest species.

Abstract

In ecosystems with frequent cloud immersion, the influence on plant water balance can be important. While cloud immersion can reduce plant water loss via transpiration, recent advances in methodology have suggested that many species also absorb water directly into leaves (foliar water uptake). The current study examines foliar water uptake and its influence on daily plant water balance in tree species of the endangered spruce–fir forest of the southern Appalachian Mountains, USA. These mountain-top communities are considered relic, boreal forests that may have persisted because of the benefits of frequent cloud immersion. We examined changes in needle water content, xylem water potentials, and stable isotope values in saplings of the two dominant tree species, Abies fraseri and Picea rubens before and after a 24 h period of experimental cloud immersion. Both species exhibited foliar water uptake following immersion, evidenced by substantial changes in stable isotope values of extracted needle water that reflected the composition of the fog water. In addition, total needle water content improved 3.7–6.4 % following experimental submersion and xylem water potentials were significantly greater (up to 0.33 MPa) in cloud-immersed plants over control plants. These results indicate that foliar water uptake may be an adaptive strategy for utilizing cloud water and improving overall tree vigor in these most southerly distributed boreal species.     

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.     

Inferring foliar water uptake using stable isotopes of water

A growing number of studies have described the direct absorption of water into leaves, a phenomenon known as foliar water uptake. The resultant increase in the amount of water in the leaf can be important for plant function. Exposing leaves to isotopically enriched or depleted water sources has become a common method for establishing whether or not a plant is capable of carrying out foliar water uptake. However, a careful inspection of our understanding of the fluxes of water isotopes between leaves and the atmosphere under high humidity conditions shows that there can clearly be isotopic exchange between the two pools even in the absence of a change in the mass of water in the leaf. We provide experimental evidence that while leaf water isotope ratios may change following exposure to a fog event using water with a depleted oxygen isotope ratio, leaf mass only changes when leaves are experiencing a water deficit that creates a driving gradient for the uptake of water by the leaf. Studies that rely on stable isotopes of water as a means of studying plant water use, particularly with respect to foliar water uptake, must consider the effects of these isotopic exchange processes.     

不同林龄杉木人工林的水分利用效率与叶片养分浓度

以亚热带不同林龄(3、8、14、21、46年生)杉木人工林为研究对象,探索不同叶龄(当年生、1年生、2年生和3年生)叶片氮、磷养分状况和水分利用效率的差异及其相互关系.结果表明: 不同叶龄水分利用效率差异显著,总体趋势为当年生>1年生>2年生>3年生,而林龄对水分利用效率影响不显著.叶片N/P为11.4～19.6,其中,幼龄林和老龄林叶片N/P高于速生期林分叶片N/P,氮、磷浓度随叶龄的变化趋势一致,为当年生>1年生>2年生>3年生.水分利用效率随林龄变化不显著,可能是因为叶片光合速率和气孔导度同时随林龄下降.水分利用效率与叶片氮浓度相关不显著,而与叶片磷浓度呈显著正相关,与N/P呈显著负相关,表明在氮沉降增加的背景下,亚热带森林中植物磷含量逐渐成为影响水分利用效率的重要因子.     

Foliar water and solute absorption: an update

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.     

武夷山不同海拔植物水分利用效率的变化及其与养分变化的关系

以武夷山不同海拔(600、900、1300、1500、1800、2000和2100 m)的乔木、灌木和草本3种生活型植物为研究对象,观测其水分利用效率和叶片氮、磷养分浓度随海拔的变化趋势,旨在探索海拔梯度间水分、养分变化与植物水分利用效率变化的关系.结果表明: 植物水分利用效率随海拔的增加呈显著上升趋势,乔木叶片δ¹⁸O随海拔变化不显著.总体上看,叶片氮浓度未观测到规律性的变化,高海拔地区的叶片磷浓度显著高于低海拔地区.水分利用效率与叶片磷浓度呈显著正相关,而与叶片氮浓度相关性不显著.不同海拔植物水分利用效率变化主要由光合速率的变化引起,水分状况对植物水分利用效率的影响不显著.海拔梯度间植物叶片磷浓度的差异对光合速率影响显著,进而影响水分利用效率.     

Divergent patterns of foliar δ13C and δ15N in Quercus aquifolioides with an altitudinal transect on the Tibetan Plateau: an integrated study based on multiple key leaf functional traits

Aims With a close association with plant water availability, foliar δ 13 C had been investigated extensively in alpine regions; however, foliar δ 15 N has rarely been concurrently used as an indicator of plant nitrogen availability. Due to the positive correlations between leaf nitrogen content and foliar δ 13 C and δ 15 N found in previous studies, we expected that they should show consistent patterns along an altitudinal gradient.Methods To test our hypothesis, we measured foliar δ 13 C and δ 15 N in conjunction with multiple key leaf functional traits of Quercus aquifolioides, a dominant species of alpine forest on the eastern slopes of the Sygera Mountains, southeastern Tibetan Plateau from 2500 to 3800 m.Important findings (i) Contrary to our hypothesis, foliar δ 13 C exhibited a significant positive linear relationship with altitude; in contrast, foliar δ 15 N initially increased and subsequently decreased with altitude, the change in trend occurring around 3300 m. (ii) Our analyses indicated that leaf internal resistance and stomatal conductance, rather than photosynthetic capacity indicated by leaf N concentration, apparently explained the altitudinal variation in foliar δ 13 C, while differences in foliar δ 15 N were likely the result of soil N availability. (iii) Principal component analysis revealed a clear association between δ 13 C and a tradeoff between water loss and carbon gain, indicated by traits related to gas exchange such as leaf thickness, density, stomatal properties. In contrast, the second axis was associated with δ 15 N and nitrogen acquisition strategy in Q. aquifolioides across its altitudinal distribution, represented by traits related to nitrogen concentration and stomata per gram of leaf nitrogen.     

Why might roadside mulgas be better mistletoe hosts?

We tested the hypothesis that roads in arid central Australia modify roadside hydrology and nutrient patterns creating better habitat for mistletoes (Loranthaceae). In two areas of mulga woodland near Alice Springs where mulgas (Acacia aneura) are the principal mistletoe host, we compared mistletoe abundance, and host foliar water and nutrient content between roadside and interior (100 m from the road) sites. Mistletoes were significantly more abundant close to roadsides than at interior sites and mulgas had significantly higher foliar water contents at roadside compared to interior sites. Mulga foliar P and N levels were not, however, significantly different between roadside and interior sites. We conclude that roads exert a strong influence on the adjacent biota in arid central Australia primarily through facilitating water infiltration at roadside sites.     

The incidence and implications of clouds for cloud forest plant water relations

Although clouds are the most recognisable and defining feature of tropical montane cloud forests, little research has focussed on how clouds affect plant functioning. We used satellite and ground‐based observations to study cloud and leaf wetting patterns in contrasting tropical montane and pre‐montane cloud forests. We then studied the consequences of leaf wetting for the direct uptake of water accumulated on leaf surfaces into the leaves themselves. During the dry season, the montane forest experienced higher precipitation, cloud cover and leaf wetting events of longer duration than the pre‐montane forest. Leaf wetting events resulted in foliar water uptake in all species studied. The capacity for foliar water uptake differed significantly between the montane and pre‐montane forest plant communities, as well as among species within a forest. Our results indicate that foliar water uptake is common in these forest plants and improves plant water status during the dry season.     

毛乌素沙地两种典型灌木叶片凝结水吸收能力及吸水途径

凝结水是半干旱地区生态系统重要的水源, 已有研究表明, 一些沙生植物可以通过叶片吸收凝结水以改善其水分状况。该研究以毛乌素沙地典型沙生灌木黑沙蒿(又称油蒿)(Artemisia ordosica)和北沙柳(Salix psammophila)为研究对象, 研究这两种植物的叶片是否具有吸收凝结水的能力, 并探究叶片吸水的途径及运移的通道。分别将黑沙蒿与北沙柳失水和未失水离体枝条置入人工模拟加湿室中, 使用高丰度氘水标记的凝结水进行浸润实验, 比对浸润前后枝条质量、叶片水及茎水氢同位素丰度变化, 确定黑沙蒿和北沙柳的叶片吸水能力; 并将盆栽黑沙蒿和北沙柳整株置入人工模拟加湿室, 使用荧光标记的凝结水进行浸润实验, 比对浸润前后叶片、小枝荧光显像, 确定黑沙蒿和北沙柳叶片吸收和运移凝结水的途径。结果显示: (1)黑沙蒿和北沙柳未失水枝条在浸润前后质量无显著差异, 黑沙蒿和北沙柳失水离体枝条在凝结水浸润后质量显著提高了2.04%和6.74%, 叶片水氘丰度提高了170.10‰和104.09‰, 茎水氘丰度提高了10.52‰和12.72‰; (2)荧光标记凝结水浸润后, 荧光示踪剂分布在黑沙蒿和北沙柳叶片的角质层、气孔、海绵组织、栅栏组织和维管束中, 黑沙蒿叶片的厚角组织中也发现了荧光示踪剂, 两种灌木小枝的表皮、韧皮部、木质部和髓中均观察到荧光。以上结果表明, 毛乌素沙地两种典型灌木叶片均具有吸收凝结水的能力, 水分亏缺植株的吸水能力更强; 两种灌木叶片通过气孔或角质层吸收凝结水, 并通过叶肉运移至维管束乃至小枝。黑沙蒿与北沙柳叶片具有的吸水功能可能是其适应干旱期水分亏缺的重要水分利用策略。