Decoding Leaf Wettability: The Role of Physicochemical Leaf Traits in Hydrophytic Plants

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T. Bole, Ratan. V. More, Mansingraj. S. Nimbalkar, Tukaram. D. Dongale, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6731857/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 19 Feb, 2026 Read the published version in Wetlands Ecology and Management → Version 1 posted 10 You are reading this latest preprint version Abstract Leaf wettability refers to the extent to which water adheres on leaf surfaces., which can have positive or negative effect on leaf, plant and ecosystem functioning. Hydrophytic plants have developed morphological adaptations to cope up with the excess presence of water. In this study, we analyzed contact angles and key physicochemical leaf traits including length, width, area, thickness, water adhesion, stomatal density, trichome density, surface structure, and surface free energy of nine wetland plant species belonging to three different habitats i.e. floating, submerged and emergent, on the basis of leaf position, from the Western Ghats of Maharashtra. Measurements were taken for both the adaxial and abaxial surfaces to determine the correlations between these traits and leaf wettability. Among the examined traits, surface free energy emerged as the primary determinant of hydrophobicity, exhibiting a strong negative correlation with contact angle. Contact angles ranged from 61° to 143° on the adaxial surface and from 47° to 124° on the abaxial surface. While most other physical traits did not show significant correlation, the presence of trichomes in Pistia stratiotes a floating hydrophyte enabled the formation of water beads on its surface. Additionally, Scanning Electron Microscopy (SEM) was employed to investigate the microstructural and nano-structural features of both sides of the leaf, providing further insights into their role in wettability. Studying the adaptations of the hydrophytes to their environment reveals that incorporating wettability and nano structural surface traits, along with other plant characteristics, enhances our understanding of plant–water interactions in wetland ecosystems. contact angle hydrophobicity scanning electron microscopy surface free energy wettability Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction Leaf wettability refers to the interaction of water with respect to the leaf surface, either by spreading or forming droplet, primarily measured by by the contact angle (CA) formed between a water droplet and the leaf surface. A high contact angle (greater than 90°) indicates hydrophobicity, leading to the formation of water beads that quickly shed from the surface, whereas a low contact angle (less than 90°) signifies hydrophilicity, allowing water to spread across the surface (Koch and Barthlott 2009 ). According to Aryal and Neuner ( 2010 ) and Wang et al. ( 2014 ) the wettability of leaves can be classified as follows: Leaves with contact angles less than 40° are categorized as super hydrophilic, reflecting an exceptionally strong affinity for water. Surfaces with contact angles between 40° and 90° are regarded as highly wettable. When the angle lies between 90° and 110°, the surface is considered wettable, whereas angles from 110° to 130° suggest moderate hydrophobicity. Leaf surfaces exhibiting contact angles in the range of 130° to 150° are identified as hydrophobic, while those exceeding 150° are termed superhydrophobic, indicating pronounced water repellency. This classification system provides a useful framework for comparing the wettability of different plant species under varying environmental conditions. This property is influenced by both the chemical composition and the micro- to nanostructure of the leaf surface, including the presence of epicuticular waxes and specialized structures like trichomes (Bhushan and Jung 2011 ). With the rise in extreme weather events like heavy rainfall and droughts (Allan 2011 ), the interaction between leaves and atmospheric water has become increasingly relevant. Plant leaves not only interact directly with rainfall but also serve as condensation surfaces for fog and dew, thereby influencing the manner in which water reaches the ground (Dunkerley 2020 ). The retention of surface water on leaves not only alters the hydrological cycle but also has a diverse impact on the plant itself (Dawson and Goldsmith 2018 ). The effects on plants are both positive and negative. Depending on the conditions surface water retention can block transpiration and photosynthesis (Aparecido et al. 2016 , 2017 ; Berry and Goldsmith 2020 ) whereas in some cases foliar water uptake can boost photosynthesis and growth (Eller et al. 2013 ; Carmichael et al. 2020 ). The formation of water films on leaf surfaces can obstruct stomatal openings, thereby impeding CO₂ diffusion and reducing photosynthetic efficiency (Smith and McClean 1989 ). Given that CO₂ diffuses approximately 10,000 times slower in water than in air, even thin water layers can significantly limit gas exchange (Brewer 1991 ). Water films can alter the leaf's boundary layer, affecting transpiration rates. While a continuous water film may reduce transpiration by increasing resistance to water vapor diffusion, it can also lead to uncontrolled water loss if the cuticle is compromised (Brewer and Smith 1997 ). ​ In epiphytic Bromeliads, wettable leaves play a specialized role in absorption of water and nutrient (Zambrano et al. 2019 ). Hydrophobic leaf surfaces can deter pathogen establishment by minimizing water retention, which is essential for the germination of many fungal spores. Conversely, hydrophilic surfaces that retain water may create favorable conditions for pathogen proliferation (Holder 2007 ).​ The "lotus effect," characterized by superhydrophobic surfaces with self-cleaning properties, allows water droplets to remove contaminants and pathogens as they roll off the leaf, thereby maintaining optimal photosynthetic surfaces and reducing disease incidence (Barthlott and Neinhuis 1997 ).​ Now in general, it appears that water presence for brief periods on the leaf surfaces can be beneficial but prolonged wetness often proves disadvantageous. Therefore, the balance of leaf wettability is crucial, with adaptations varying depending on environmental conditions and ecological niches. Wetland habitats are characterized by unique environmental conditions such as constant or periodic waterlogging and high humidity. These conditions pose significant challenges to plants, including reduced gas exchange due to water films on leaf surfaces and increased susceptibility to pathogens that thrive in moist environments (Reynolds et al. 1989 ; Evans et al. 1992 ; Huang et al. 2006 ). To cope with excessive water, wetland plants have evolved various morphological and physicochemical adaptations. One crucial adaptation is the development of leaf surface traits that influence wettability, thereby affecting water retention and promoting efficient water shedding (Sikorska et al. 2017 ).​ The leaf wettability depends upon various factors like the epicuticular wax layer thickness and composition which play a major role in minimizing leaf wetting (Holloway 1969 ; Koch et al. 2006 ). Trichomes with higher densities often enhance hydrophobicity (Brewer et al. 1991 ; Pan et al. 2021 ). Stomatal density and spatial arrangement affect surface roughness and influence contact angle measurements (Brewer and Nuñez 2007 ; Wang et al. 2010 ). Surface free energy (SFE) affects the interaction between water and the leaf, with lower SFE correlating with higher hydrophobicity (Wang et al. 2014 ). Additionally, nano- and microstructural features like wax crystals and surface ridges further enhance water repellency by increasing roughness and modifying surface chemistry (Sikorska et al. 2017 ). So, in the previously done research on the wettability of plant leaves from various habitats the focus has been given to individual trait or general principle of wettability rather than systematically correlating multiple physicochemical and structural characteristics with wettability across diverse wetland taxa. In this study, we investigate the relationship between leaf wettability and key physicochemical traits in nine wetland (aquatic) plant species native to the Western Ghats of Maharashtra, India. Both adaxial and abaxial surfaces were assessed for contact angles and also analyzed a set of traits including leaf length, width, length to width ratio, area, thickness, stomatal and trichome densities, water adhesion, surface structure, and surface free energy. Scanning electron microscopy (SEM) was employed to examine micro- and nano-scale surface features. This study aims to identify the key leaf traits influencing hydrophobicity and to investigate their role in facilitating adaptive strategies in wetland plant species. Materials and Methods Using random sampling method for the selection, nine pond dwelling aquatic plants were selected for analyzing leaf wettability and the factors influencing it. The young and fully matured leaves after collecting were transported to the laboratory in the air tight packets so that they remain as fresh as possible. A total of 5 leaves were analyzed per plant species to determine the selected traits used in this study. Leaf wettability was determined measuring the contact angles of both adaxial and abaxial surfaces by the sessile drop method with the help of goniometer (rame hart goniometer). A 10 µl droplet of distilled water was deposited on the leaf surface employing a syringe and contact angle was determined by analyzing the image captured using CA meter software. For calculating surface free energy, we used 1-bromonapthalene as the nonpolar solvent. Using the Owens and Wendt ( 1969 ) method where the total surface free energy is divided into two components: a dispersive (non-polar) component and a polar component, surface free energy was estimated by measuring the contact angles of at least two different liquids typically one polar (distilled water) and one non-polar (1-bromonapthalene) whose surface free energy is known on the solid surface. The formula to calculate SFE is as following: $$\:{\gamma\:}_{s}=\:{\gamma\:}_{s}^{d}+\:{\gamma\:}_{s}^{p}$$ $$\:{\gamma\:}_{l}\left(1+cos\:\theta\:\right)=\:2\:\left.\left(\:\sqrt{\left\{{\gamma\:}_{s}^{d}\cdot\:{\gamma\:}_{l}^{d}\right\}}+\:\sqrt{\left\{{\gamma\:}_{s}^{p}\cdot\:{\gamma\:}_{l}^{p}\right\}}\right.\right)$$ \(\:{\gamma\:}_{s}\:\) : Total surface free energy of the solid, ​ \(\:{\gamma\:}_{s}^{d}\) : Dispersive component of the solid's surface free energy, \(\:{\gamma\:}_{s}^{p}:\) Polar component of the solid's surface free energy, \(\:{\gamma\:}_{l}\) \(\::\:\text{T}\text{o}\text{t}\text{a}\text{l}\:\text{s}\text{u}\text{r}\text{f}\text{a}\text{c}\text{e}\:\text{f}\text{r}\text{e}\text{e}\:\text{e}\text{n}\text{e}\text{r}\text{g}\text{y}\:\text{o}\text{f}\:\text{t}\text{h}\text{e}\:\text{l}\text{i}\text{q}\text{u}\text{i}\text{d}\) , \(\:{\gamma\:}_{l}^{d}\) : Dispersive component of the liquid's surface free, energy, \(\:{\gamma\:}_{l}^{p}\) : Polar component of the liquid's surface free energy, \(\:\theta\:\) : Contact angle of the liquid on the solid For surface structure analysis, air-dried leaf samples were placed on mounting stubs and gold sputtering is done, then imaged using a JEOL JSM-IT200 scanning electron microscope to assess structural differences in relation to contact angle (Pathan et.al. 2008 ). Leaf length and width were measured using a ruler, and fresh weight was recorded using a standard weight box. The length-to-width ratio was subsequently calculated. Leaf area was derived from length and width measurements, adjusted by a shape correction factor, following the method described by Schrader et al. ( 2021 ). Leaf thickness was measured using a micrometer gauge. Leaf adhesion on both the adaxial and abaxial surfaces was determined using the formula W = (W₁ – W₀)/S , where W₁ represents leaf weight after wetting, W₀ is the weight before wetting, and S denotes leaf’s surface area (Wang et al. 2014 ). Trichome density (trichomes/mm²) on both adaxial and abaxial surfaces was manually evaluated using a dissecting light microscope with observations taken from ten sites per leaf (Viz and Pacada 2022 ). Clear nail polish was thinly applied to both upper and lower leaf surfaces and left to dry. The dried film was peeled off with adhesive tape, mounted on slides, and examined under a light microscope to count stomata per mm²(Zhu et al. 2018 ). The parameters were analyzed for their correlation with the adaxial and abaxial contact angles to determine their effect on wettability. Pearson correlation coefficient was computed for each parameter against the contact angles of both surfaces, with statistical significance at p < 0.05, with the help of Statistica 10 software (StatSoft Inc., 2011 ). Results A total of nine wetland plant species belonging to three habitats i.e. submerged, floating, and emergent based on leaf’s position, were assessed for various morphological and surface traits in relation to leaf wettability, as quantified by contact angle (CA) measurements on both the leaf surfaces. On the adaxial surface, contact angles ranged between 61° and 143°, while on abaxial surface they varied from 47° to 124°, indicating a broad spectrum of wettability among the sampled species. Out of the nine species examined, only three i.e. Colocasia esculenta , Nelumbo nucifera from emergent habitat and Pistia stratiotes from floating habitat exhibited hydrophobic properties (CA > 90°), whereas the remaining six species displayed hydrophilic surfaces (CA < 90°). Additionally, in all but one species assessed, the adaxial surface showed contact angle more than the abaxial surface. All other physicochemical parameters were measured whose values are mentioned in the Table 1 . Pearson’s correlation analysis was employed to evaluate the strength, direction and significance of the associations between leaf traits (e.g., width, trichome density, surface free energy) and contact angle values. Table 1 shows the data of all the parameters studied. This data was then used to find out Pearson’s correlation coefficient values between the traits of leaf studied and the adaxial and abaxial contact angles measured at p < 0.05, using Statistica 10 software. Table 1 Leaf trait measurements of wetland plants used to assess correlations with contact angle and surface wettability: Plant name Habitat Length ± SD (cm) Width ± SD (cm) L/W ratio Weight ± SD (g) Thickness ± SD(mm) Area (mm²) Adaxial CA ± SD (°) Abaxial CA ± SD (°) Water adhesion ± SD (adaxial) Water adhesion ± SD (abaxial) Adaxial CA ± SD (°) nonpolar Abaxial CA ± SD (°) nonpolar SFE adaxial (mj/m2) SFE abaxial (mj/m2) Stomatal density adaxial (per mm² ) Stomatal density abaxial ( per mm² ) Trichome density adaxial (per mm²) Trichome density abaxial (per mm²) Colocasia esculenta emergent 18.68 ± 0.67 12.52 ± 0.13 1.49 3.768 ± 0.19 0.272 ± 0.007 16371 121.25 ± 2.1 95.55 ± 1.52 0.61 ± 0.23 g/m² 1.22 ± 0.45 g/m² 76 ± 3.42 45.3 ± 2.90 17.18 33 22.5 ± 1.1 148.75 ± 1.66 0 0 Nelumbo nucifera emergent 15.4 ± 0.64 18.46 ± 0.88 0.83 6.84 ± 0.27 0.318 ± 0.007 22458 143.8 ± 2.2 124.2 ± 1.8 0.97 ± 0.09 g/m² 1.33 ± 0.31 g/m² 62.2 ± 3.06 46.6 ± 2.75 28.75 29.5 498.75 ± 5.1 0 0 0 Typha angustifolia emergent 144.02 ± 2.98 2.8 ± 0.30 51.43 7.36 ± 0.17 1.296 ± 0.018 25808 79.5 ± 2.4 76.45 ± 1.9 6.58 ± 0.39 g/m² 7.05 ± 0.44 g/m² 30.7 ± 2.01 21.35 ± 2.04 42.29 45.7 417.5 ± 4.12 393.75 ± 3.65 0 0 Ipomoea aquatica floating 9.02 ± 0.68 5.62 ± 0.07 1.6 0.94 ± 0.05 0.31 ± 0.007 3549 75.15 ± 0.58 47.3 ± 0.8 14.09 ± 0.42 g/m² 19.72 ± 0.73 g/m² 16.9 ± 1.25 27.2 ± 1.63 47.09 59.3 156.25 ± 2.21 333.75 ± 3.75 0 0 Nymphaea nouchali floating 21.96 ± 1.59 18.6 ± 0.316 1.18 11.49 ± 0.54 0.42 ± 0.006 32268 63.1 ± 0.8 52 ± 1.5 13.94 ± 0.83 g/m² 16.42 ± 0.19 g/m² 44.05 ± 2.79 32.35 ± 2.94 46.18 55.6 397.5 ± 2.36 0 0 0 Pistia stratiotes floating 11.12 ± 0.23 8.36 ± 0.13 1.33 7.86 ± 0.13 0.718 ± 0.017 6322 95.64 ± 0.58 113.4 ± 0.7 36.38 ± 0.72 g/m² 104.4 ± 0.87 g/m² 73.1 ± 3.28 51.2 ± 2.56 21.68 29.1 65.2 ± 2.52 0 25.8 ± 1.6 29 ± 1.78 Cryptocoryne retrospiralis submerged 23.12 ± 0.28 1.48 ± 0.074 15.62 1.63 ± 0.028 0.292 ± 0.011 2258 62.85 ± 1.75 54.6 ± 1.77 10.62 ± 0.76 g/m² 15.5 ± 0.38 g/m² 45.6 ± 2.58 16.2 ± 2.71 46.5 56.9 98.75 ± 2.52 91.25 ± 1.46 0 0 Eriocaulon cuspidatum submerged 4.1 ± 0.109 0.92 ± 0.09 4.45 0.208 ± 0.007 1.14 ± 0.048 268 84.2 ± 1.7 58.5 ± 2.2 52.24 ± 0.33 g/m² 55.97 ± 0.67 g/m² 11.1 ± 1.04 9.7 ± 3.22 45.25 55.6 0 8.75 ± 0.89 0 0 Hygrophila auriculata submerged 6.24 ± 0.35 1.24 ± 0.048 5.03 0.22 ± 0.015 0.31 ± 0.011 511 61.8 ± 1.3 60.05 ± 1.8 29.41 ± 0.51 g/m² 35.29 ± 0.29 g/m² 34.2 ± 1.76 39.2 ± 2.90 49.49 49.2 226.25 ± 4.01 298.75 ± 4.21 7.6 ± 0.8 4 ± 0.63 The data in Table 1 had been used to find out the correlation between the various leaf parameters and the measure of wettability i.e. contact angle. The correlation coefficient ( r ) values show how strong is the relationship between the two parameters taken into consideration. of every parameter against adaxial contact angles. of every parameter against abaxial contact angle. Figures 1 and 2 show Pearson’s correlation matrix heatmaps representing the linear relationships between key leaf parameters and contact angles on the adaxial (Fig. 1 ) and abaxial (Fig. 2 ) surfaces. The color gradients reflect the strength (r-values) and direction of the correlations, with values ranging from − 1 (strong negative) to + 1 (strong positive). Using these Pearson’s correlation coefficient values we found out the significance of all the parameters affecting the contact angles i.e. wettability of the leaves. The values for Pearson correlation coefficient and p- values which shows significance of the correlation are mentioned in the Table 2 . Table 2 Pearson Correlation Coefficients (r) and p-values Between Leaf Parameters and Contact Angles. Parameter r-value (Adaxial) p-value (Adaxial) r-value (Abaxial) p-value (Abaxial) SFE abaxial -0.975 0.000 SFE adaxial -0.80 0.010 Width 0.561 0.116 0.480 0.191 Water adhesion adaxial -0.35 0.365 Stomatal density abaxial -0.36 0.345 Area 0.277 0.471 0.264 0.493 L/W ratio -0.235 0.543 -0.11 0.782 Weight 0.197 0.611 0.393 0.295 Water adhesion abaxial 0.160 0.681 Stomatal density adaxial 0.15 0.703 Thickness -0.111 0.776 -0.010 0.979 Length -0.094 0.810 0.02 0.961 Trichome density abaxial 0.47 0.203 Trichome density adaxial 0.017 0.966 Note: Significant results (p < 0.05) are displayed in bold. Among all measured parameters, surface free energy (SFE) showed the strongest correlations with contact angle. SFE on both adaxial and abaxial surfaces was negatively correlated with contact angle values, indicating that leaves with lower surface energy exhibited higher water repellency. For example, Colocasia esculenta , with low abaxial SFE (33 mJ/m²), showed high contact angles (adaxial: 121.25°, abaxial: 95.55°), suggesting strong hydrophobicity. Conversely, Ipomoea aquatica , which had the highest abaxial SFE (59.3) exhibited the lowest abaxial contact angle (47.3°), reflecting greater wettability. No consistent or significant correlations were found between contact angle and leaf length, width, thickness, area, or weight. In this study, trichome densities did not show significant associations with contact angle, likely due to their low or absent values across most species. But in case of Pistia stratiotes , a floating hydrophyte, the hydrophobic behavior is the result of presence of trichomes ( density more than 25 mm⁻²) on both of its surfaces. Regarding stomatal density, no significant association between stomatal density and contact angle on either surface was found as per the Pearson correlation analysis. Scanning electron microscopy (SEM) was used to examine the microstructural features of both adaxial and abaxial leaf surfaces for all nine wetland plant species. The SEM images revealed distinct variations in surface texture, presence/absence of trichomes, stomatal exposure, and cuticular structures, which were analyzed in relation to contact angle measurements to infer their influence on leaf wettability.The two hydrophobic species, Colocasia esculenta and Nelumbo nucifera , showed specialized surface features that are consistent with high contact angle values. The surfaces of these leaves displayed a hierarchical surface architecture, including micro- and nanoscale roughness combined with dense epicuticular wax crystals. In Nelumbo nucifera ( Fig. 3 ; cand d) , papillate epidermal cells and dense wax tubules were observed—hallmarks of the "lotus effect," which is known to promote superhydrophobicity by minimizing contact area between water and the surface (Barthlott and Neinhuis 1997 ; Koch et al. 2009). Similarly, the morphology of Colocasia esculenta ( Fig. 3 ; a and b) , leaf exhibits a two-tier surface structure characterized by hexagonal microcavities resembling a honeycomb-like pattern (Kumar and Bhardwaj 2020 ) with uniform wax coverage and elevated surface ridges that contribute to its water-repellent properties. The surface structures of Pistia stratiotes ( Fig. 3 ; e and f) show the waxy trichomes that help bead up the water drops. The other six plants’ contact angle in this study is below 90 degrees showing their affinity towards water. Their SEM analysis revealed surface structures consistent with those hydrophilic surfaces reported in previous studies. The scanning electron micrographs (Fig. 4 a–l) reveal consistent structural adaptations among the studied wetland species, supporting their hydrophilic surface behavior. Across all samples, the absence of micro- and nanoscale hierarchical features, including wax crystal deposition, correlates with low contact angles and increased wettability. In Nymphaea nouchali (Fig. 4 a, b), both leaf surfaces show an undulating and irregular epidermis with no epicuticular projections. This simple surface structure promotes close contact with water, enhancing hydrophilicity. Similarly, Cryptocoryne retrospiralis (Fig. 4 c, d) has an uneven surface with sunken stomata but lacks wax crystals, which likely aids in water absorption and gas exchange in moist or submerged environments. Ipomoea aquatica (Fig. 4 e, f) exhibits a smooth surface with fine cuticular striations, which may help water spread evenly while reducing air trapping. The absence of wax further supports its wettable nature. In Hygrophila auriculata (Fig. 4 g, h), the stomata are located in shallow depressions surrounded by smooth cuticular ridges, likely guiding water movement without blocking gas exchange. Eriocaulon cuspidatum (Fig. 4 i, j) shows a mostly uniform surface with slight striations near the stomata, consistent with water-retaining surfaces adapted for diffusion-based gas exchange. Lastly, Typha angustifolia (Fig. 4 k, l) displays uneven, micropatterned surfaces without waxes, suggesting a structural adaptation that maintains hydrophilicity while providing surface stability in wet habitats. Discussion This study investigated the relationship between leaf surface traits and wettability in nine wetland plant species by analyzing contact angles on both adaxial and abaxial surfaces. The results revealed a broad spectrum of wettability, with only three species exhibiting hydrophobic characteristics and the remainder showing hydrophilic behavior. Surface free energy (SFE) was found to be the most consistent predictor of contact angle, while traits like stomatal and trichome density showed inconsistent or weak correlations. These findings provide insight into how structural and physicochemical leaf traits contribute to ecological adaptations in wetland environments. Hydrophobicity and leaf positioning Interestingly, except Pistia stratiotes , the floating hydrophyte, whose hydrophobic behavior is attributed to the trichomes present on it, the other hydrophobic species are characterized by leaves that are held above the water surface rather than being submerged or floating. This leaf position suggests an ecological adaptation that favors water repellency, likely to facilitate self-cleaning (lotus effect) and prevent prolonged water retention, which can block gas exchange or promote microbial colonization (Barthlott and Neinhuis 1997 ; Koch et al. 2009). The elevated and hydrophobic leaves of Nelumbo nucifera are particularly well-known for their superhydrophobic nature, attributed to hierarchical surface structuring and wax crystals, which significantly reduce surface free energy and promote water rolling (Bhushan and Jung 2006 ). Similarly, Colocasia esculenta has been reported to exhibit a highly water-repellent surface, potentially serving to maintain gas exchange and light capture in wetland conditions (Ensikat et al. 2011 ). These surface adaptations can be viewed as a response to the plants’ aerial leaf posture, which exposes them to rain and surface splash, increasing the need for self-cleaning mechanisms (Barthlott et al. 2007 ). Adaptation of hydrophilic surfaces in aquatic plants In contrast, the remaining hydrophilic species, often with submerged or floating leaves, do not exhibit such repellency. This observation supports earlier studies showing that there is reduced selective pressure for hydrophobic surfaces in aquatic or semi-aquatic environments where leaves remain consistently wet or underwater (Vogel 1988 ; Riedl and Riedl 2004 ). In such species, hydrophilic surfaces may even aid in water absorption or nutrient exchange, depending on environmental conditions. This surface polarity in wettability is well-documented in foliage, often reflecting functional specialization. The adaxial surface, being more exposed to environmental factors like rainfall and sunlight, may evolve more hydrophobic traits to facilitate rapid droplet shedding and self-cleaning. The abaxial surface, typically more shaded and involved in gas exchange through stomata, may maintain a more hydrophilic profile to support physiological functions (Neinhuis and Barthlott 1997 ; Brewer 1991 ). Contribution of surface free energy to wettability The inverse relationship between SFE and contact angle aligns with established principles of surface chemistry, where lower SFE facilitates water repellency (Bhushan and Jung 2011 ). This relationship emphasizes the significance of surface physicochemical properties in determining leaf wettability across species with different ecological strategies. Trichomes as determinants of water repellency Hairy leaf surfaces, such as those of Alchemilla vulgaris , are highly water-repellent because their stiff hairs keep water droplets elevated, preventing contact with the leaf surface and creating a fakir-like effect (Otten and Herminghaus 2004 ). Brewer et al. ( 1991 ) found that plant species with dense trichomes (more than 25 mm⁻²) tended to be more water-repellent, though some still retained droplets. Similarly, Neinhuis and Barthlott ( 1997 ) observed that leaves with wax-covered trichomes were extremely water-repellent, while those with non-waxy trichomes only resisted water briefly after droplet contact. Inconsistent influence of stomatal density This observation aligns with the findings of Lott ( 2021 ), who reported that stomatal density in five plant species did not significantly influence leaf wettability, possibly due to limited sample size or species-specific traits. All plants in Lott’s study had stomata only on the abaxial surface, as is typical in many plant species. However, this finding contrasts with studies by Brewer and Nuñez ( 2007 ), who observed that surfaces with higher stomatal densities tended to be less wettable, possibly due to the microstructural complexity that encourages droplet retention. Unlike our findings, Wang et al. ( 2021 ) found a positive correlation between stomatal density and contact angle in species studied in Xi’an, China—suggesting that in some cases, higher stomatal density may increase hydrophobicity by contributing to surface microtexture or by trapping air, similar to the Cassie–Baxter wetting regime. Taken together, these contrasting reports suggest that the influence of stomatal density on wettability is context-dependent and may be modulated by other surface features such as cuticular wax, trichomes, and surface roughness. In this study, the lack of a clear trend indicates that stomatal density alone may not be a reliable predictor of hydrophobicity across diverse wetland species. Surface structure of hydrophilic leaves Hydrophilic leaf surfaces typically lack the complex micro- and nano-scale structures found on superhydrophobic leaves. They often appear smooth or slightly rough under SEM and show little to no wax crystalloids, which contributes to lower contact angles and increased water spreading (Neinhuis and Barthlott 1997 ). The cuticle on these leaves may be thin, cracked, or composed of more polar compounds, allowing greater wettability (Riederer and Schreiber 2001 ). In some cases, visible features such as stomata or trichomes can retain water, further promoting hydrophilic behavior (Brewer and Smith 1997 ). These surfaces also tend to have higher surface free energy due to the absence of hydrophobic waxes (Koch et al. 2008 ). As a result, contact angles are typically below 90°, indicating that water readily spreads rather than beads up (Mittal and Davis 2007 ).The collective absence of epicuticular wax crystals—a key determinant of hydrophobicity—across all species is a defining trait of these hydrophytes. These traits are not coincidental; rather, they represent evolutionary responses to persistently wet environments where maintaining a thin water layer on the leaf surface is advantageous. Such features may enhance nutrient absorption, support foliar respiration during submersion, and minimize damage from waterlogging (Rascio 2002 ; Sakamoto and Yamaguchi 2017 ). These findings reinforce the view that hydrophilic leaf surface structures are ecologically advantageous in aquatic and semi-aquatic environments. They contrast starkly with the superhydrophobic adaptations found in xerophytic and canopy-dwelling species, where water repellence is vital for self-cleaning and pathogen resistance (Barthlott and Neinhuis 1997 ; Koch et al. 2009). Conclusion This research undertaken examined the relationship between leaf’s physicochemical traits and wettability in hydrophytes living in three different ecological conditions i.e. emergent, floating, and submerged. Three plants from each were studied. Contact angle measurements revealed that only three species— Nelumbo nucifera , Colocasia esculenta (emergent), and Pistia stratiotes (floating) exhibited hydrophobic behavior, while the remaining species displayed hydrophilic surfaces. Notably, the two emergent species had aerial leaves that avoid direct water contact, and the hydrophobicity of Pistia stratiotes was attributed to the presence of dense, waxy trichomes. All other plant species showed wetting behavior. Among the various morphological and anatomical leaf traits analyzed, surface free energy (SFE) emerged as the strongest predictor of contact angle. The inverse correlation between SFE and contact angle reinforces the importance of physicochemical properties over purely structural features in determining leaf wettability. All other leaf parameters studied i.e. leaf length, width, area, water adhesion etc. did not have stark impact in determining wettability. SEM analysis further showed that most hydrophilic species lacked micro- and nanoscale hierarchical surface features and epicuticular wax crystalloids—structures typically associated with water repellency in terrestrial species. These findings suggest that leaf wettability in wetland plants is a complex trait shaped by both ecological context and adaptive function. While hydrophobic surfaces in emergent and floating species may aid in self-cleaning and gas exchange, hydrophilic surfaces in submerged or consistently wet leaves may facilitate foliar water uptake, nutrient absorption, or sustained physiological activity under saturated conditions. Thus, wettability patterns reflect functional adaptations aligned with each species’ microhabitat and life form. This indicates that leaf wettability is closely linked to habitat-specific ecological pressures and contributes to the functional diversity of plant responses in wet environments. Further research is needed to explore how these traits interact with environmental variables across diverse wetland habitats. Declarations Competing interests : The authors have no financial or nonfinancial interests to disclose. Funding: no funding Author Contribution All authors contributed significantly to the research and manuscript preparation. STB was responsible for conceptualization, methodology, and experimentation, literature review, drafting the original manuscript. MSN designed the experiments, collected data, and performed formal analysis. TDD conducted the curated data, and contributed to reviewing and editing the manuscript. RVM plant material collection, data validation, critical manuscript review. NVP handled final manuscript editing, proofreading, supervision, and correspondence. All authors reviewed and approved the final version of the manuscript. Acknowledgement The authors are thankful to the Principal, The New College, Kolhapur, for providing the laboratory facilities. STB is also thankful to University Grants Commission, New Delhi, [ UGC Ref. No.: 835/ (CSIR-UGC NET JUNE 2019] for providing the financial assistance through UGC-JRF award for research work. Data availability: The datasets generated during and/or analyzed during the current study are available from the corresponding author on reasonable request. References Allan, R. P. (2011). Climate change: Human influence on rainfall. Nature , 470 (7334), 344–345. https://doi.org/10.1038/470344a Aparecido, L. M. T., Woo, S., Suazo, A. A., and Goldsmith, G. R. (2016). Foliar water uptake: A common water acquisition strategy for plants of the redwood forest. Oecologia , 182 (3), 731–744. Aparecido, L. M. T., et al. (2017). Seasonal variation in leaf wetness and its role in foliar water uptake in a tropical montane cloud forest. Plant Ecology , 218 , 1001–1013. 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Pan, X., Zhang, H., Liu, Z., Wu, H., and Wang, C. (2021). The relationship between trichome density and wettability in desert plants. Plants , 10 (6), 1233. Pathan, A. K., Bond, J., and Gaskin, R. E. (2008). Sample preparation for SEM of plant surfaces. Micron , 39 (7), 1049–1061. Rascio, N. (2002). The underwater life of secondarily aquatic plants: Some problems and solutions. Critical Reviews in Plant Sciences , 21 (4), 401–427. Reynolds, J. F., Kemp, P. R., Tenhunen, J. D., and Lange, O. L. (1989). Modeling the effects of drought on the water use of vegetation. Agricultural and Forest Meteorology , 46 (1–2), 193–209. Riederer, M., and Schreiber, L. (2001). Protecting against water loss: Analysis of the barrier properties of plant cuticles. Journal of Experimental Botany , 52 (363), 2023–2032. Riedl, R., and Riedl, M. (2004). The life of the wetland plants . Springer. Sakamoto, Y., and Yamaguchi, T. (2017). Cuticular features and leaf wettability of aquatic plants. Plant Biology , 19 (6), 928–935. Schrader, J., Shi, P., Royer, D. L., Peppe, D. J., Gallagher, R. V., Li, Y., Wang, R., and Wright, I. J. (2021). Leaf size estimation based on leaf length, width and shape. Annals of Botany , 128 (4), 395–406. https://doi.org/10.1093/aob/mcab078 Sikorska, D., Papierowska, E., Szatyłowicz, J., Sikorski, P., Suprun, K., and Hopkins, R. J. (2017). Variation in leaf surface hydrophobicity of wetland plants: The role of plant traits in water retention. Wetlands , 37 (5), 997–1002. https://doi.org/10.1007/s13157-017-0924-2 Smith, W. K., and McClean, T. M. (1989). Adaptive relationship between leaf water repellency, stomatal distribution, and gas exchange. American Journal of Botany , 76 (3), 465–469. StatSoft, Inc (2011) STATISTICA (data analysis software system) version10. Viz, C. A., and Pacada, M. R. (2022). Methods in trichome density analysis using optical microscopy. Botanical Methods , 4 (1), 19–25. Vogel, S. (1988). Life's devices: The physical world of animals and plants . Princeton University Press. Wang, C., Liu, Y., and Li, X. (2010). Influence of stomatal density on leaf hydrophobicity. Journal of Experimental Botany , 61 (6), 1569–1574. Wang, Z., Zhang, J., and Li, Y. (2014). Surface free energy and wettability of rice leaf surfaces. Langmuir , 30 (5), 1397–1403. Wang, Y., Zhang, C., Hu, C., and Zeng, J. (2021). Variation in stomatal traits and its relation to leaf wettability in plants from a subtropical region. Ecological Indicators , 121 , 107135. Zambrano, J., et al. (2019). Leaf surface properties and their implications for foliar water uptake in tropical montane epiphytes. Plant, Cell and Environment , 42 (2), 434–445. Zhu, C., et al. (2018). Quantitative analysis of stomatal density using nail polish impression method. Journal of Plant Research , 131 (3), 499–508. Additional Declarations No competing interests reported. Cite Share Download PDF Status: Published Journal Publication published 19 Feb, 2026 Read the published version in Wetlands Ecology and Management → Version 1 posted Editorial decision: Revision requested 09 Jun, 2025 Reviews received at journal 08 Jun, 2025 Reviews received at journal 04 Jun, 2025 Reviewers agreed at journal 01 Jun, 2025 Reviewers agreed at journal 29 May, 2025 Reviewers agreed at journal 28 May, 2025 Reviewers invited by journal 27 May, 2025 Editor assigned by journal 27 May, 2025 Submission checks completed at journal 26 May, 2025 First submitted to journal 23 May, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. 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(All the images are in 1000x magnification).\u003c/p\u003e","description":"","filename":"Picture3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6731857/v1/b5ce3757dc91b8508e9075c4.jpg"},{"id":83685696,"identity":"86a59559-e87a-4bf4-bf7e-12362cc2dd64","added_by":"auto","created_at":"2025-05-30 18:00:38","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":683929,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003ea) b)\u003c/strong\u003e SEM images of adaxial and abaxial leaf surfaces of \u003cem\u003eNymphaea nouchali \u003c/em\u003eshowing undulating and irregular epidermis with no micro or nano scaling. \u003cstrong\u003ec) d)\u003c/strong\u003eSEM images of adaxial and abaxial leaf surfaces of \u003cem\u003eCryptocoryne retrospiralis \u003c/em\u003eshowing uneven undulating surface, sunken stomata but with no wax crystals. \u003cstrong\u003ee) f)\u003c/strong\u003e SEM images of adaxial and abaxial leaf surfaces of \u003cem\u003eIpomoea aquatica\u003c/em\u003e showing smooth surface with cuticular striations but no wax crystals \u003cstrong\u003eg) h)\u003c/strong\u003e SEM images of adaxial and abaxial leaf surfaces of \u003cem\u003eHygrophila auriculata\u003c/em\u003e with smooth surface and several stomata embedded within shallow depressions and surrounded by smooth cuticular ridges. \u003cstrong\u003ei) j)\u003c/strong\u003e SEM images of adaxial and abaxial leaf surfaces of Eriocaulon\u003cem\u003ecuspidatum\u003c/em\u003e showing uniform surface topography with occasional stomatal striations. \u003cstrong\u003ek) l)\u003c/strong\u003e SEM images of adaxial and abaxial leaf surfaces of \u003cem\u003eTypha angustifolia\u003c/em\u003e showing uneven and micropatterned structures but lacking wax crystals.\u003c/p\u003e","description":"","filename":"Picture4.jpg","url":"https://assets-eu.researchsquare.com/files/rs-6731857/v1/1e29a05a348f7e071512e30d.jpg"},{"id":103251034,"identity":"90a84c87-73ab-4953-99dc-73219dfe7590","added_by":"auto","created_at":"2026-02-23 16:01:58","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":3825427,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6731857/v1/35f00eba-35f2-4183-9762-2826cfd482da.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Decoding Leaf Wettability: The Role of Physicochemical Leaf Traits in Hydrophytic Plants","fulltext":[{"header":"Introduction","content":"\u003cp\u003eLeaf wettability refers to the interaction of water with respect to the leaf surface, either by spreading or forming droplet, primarily measured by by the contact angle (CA) formed between a water droplet and the leaf surface. A high contact angle (greater than 90\u0026deg;) indicates hydrophobicity, leading to the formation of water beads that quickly shed from the surface, whereas a low contact angle (less than 90\u0026deg;) signifies hydrophilicity, allowing water to spread across the surface (Koch and Barthlott \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). According to Aryal and Neuner (\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) and Wang et al. (\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) the wettability of leaves can be classified as follows: Leaves with contact angles less than 40\u0026deg; are categorized as super hydrophilic, reflecting an exceptionally strong affinity for water. Surfaces with contact angles between 40\u0026deg; and 90\u0026deg; are regarded as highly wettable. When the angle lies between 90\u0026deg; and 110\u0026deg;, the surface is considered wettable, whereas angles from 110\u0026deg; to 130\u0026deg; suggest moderate hydrophobicity. Leaf surfaces exhibiting contact angles in the range of 130\u0026deg; to 150\u0026deg; are identified as hydrophobic, while those exceeding 150\u0026deg; are termed superhydrophobic, indicating pronounced water repellency. This classification system provides a useful framework for comparing the wettability of different plant species under varying environmental conditions. This property is influenced by both the chemical composition and the micro- to nanostructure of the leaf surface, including the presence of epicuticular waxes and specialized structures like trichomes (Bhushan and Jung \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2011\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eWith the rise in extreme weather events like heavy rainfall and droughts (Allan \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2011\u003c/span\u003e), the interaction between leaves and atmospheric water has become increasingly relevant. Plant leaves not only interact directly with rainfall but also serve as condensation surfaces for fog and dew, thereby influencing the manner in which water reaches the ground (Dunkerley \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The retention of surface water on leaves not only alters the hydrological cycle but also has a diverse impact on the plant itself (Dawson and Goldsmith \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The effects on plants are both positive and negative. Depending on the conditions surface water retention can block transpiration and photosynthesis (Aparecido et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2016\u003c/span\u003e,\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Berry and Goldsmith \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) whereas in some cases foliar water uptake can boost photosynthesis and growth (Eller et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Carmichael et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The formation of water films on leaf surfaces can obstruct stomatal openings, thereby impeding CO₂ diffusion and reducing photosynthetic efficiency (Smith and McClean \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e1989\u003c/span\u003e). Given that CO₂ diffuses approximately 10,000 times slower in water than in air, even thin water layers can significantly limit gas exchange (Brewer \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1991\u003c/span\u003e). Water films can alter the leaf's boundary layer, affecting transpiration rates. While a continuous water film may reduce transpiration by increasing resistance to water vapor diffusion, it can also lead to uncontrolled water loss if the cuticle is compromised (Brewer and Smith \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1997\u003c/span\u003e). ​ In epiphytic Bromeliads, wettable leaves play a specialized role in absorption of water and nutrient (Zambrano et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Hydrophobic leaf surfaces can deter pathogen establishment by minimizing water retention, which is essential for the germination of many fungal spores. Conversely, hydrophilic surfaces that retain water may create favorable conditions for pathogen proliferation (Holder \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2007\u003c/span\u003e).​ The \"lotus effect,\" characterized by superhydrophobic surfaces with self-cleaning properties, allows water droplets to remove contaminants and pathogens as they roll off the leaf, thereby maintaining optimal photosynthetic surfaces and reducing disease incidence (Barthlott and Neinhuis \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1997\u003c/span\u003e).​ Now in general, it appears that water presence for brief periods on the leaf surfaces can be beneficial but prolonged wetness often proves disadvantageous. Therefore, the balance of leaf wettability is crucial, with adaptations varying depending on environmental conditions and ecological niches.\u003c/p\u003e \u003cp\u003eWetland habitats are characterized by unique environmental conditions such as constant or periodic waterlogging and high humidity. These conditions pose significant challenges to plants, including reduced gas exchange due to water films on leaf surfaces and increased susceptibility to pathogens that thrive in moist environments (Reynolds et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e1989\u003c/span\u003e; Evans et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e1992\u003c/span\u003e; Huang et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). To cope with excessive water, wetland plants have evolved various morphological and physicochemical adaptations. One crucial adaptation is the development of leaf surface traits that influence wettability, thereby affecting water retention and promoting efficient water shedding (Sikorska et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).​ The leaf wettability depends upon various factors like the epicuticular wax layer thickness and composition which play a major role in minimizing leaf wetting (Holloway \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e1969\u003c/span\u003e; Koch et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). Trichomes with higher densities often enhance hydrophobicity (Brewer et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1991\u003c/span\u003e; Pan et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Stomatal density and spatial arrangement affect surface roughness and influence contact angle measurements (Brewer and Nu\u0026ntilde;ez \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Surface free energy (SFE) affects the interaction between water and the leaf, with lower SFE correlating with higher hydrophobicity (Wang et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Additionally, nano- and microstructural features like wax crystals and surface ridges further enhance water repellency by increasing roughness and modifying surface chemistry (Sikorska et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSo, in the previously done research on the wettability of plant leaves from various habitats the focus has been given to individual trait or general principle of wettability rather than systematically correlating multiple physicochemical and structural characteristics with wettability across diverse wetland taxa. In this study, we investigate the relationship between leaf wettability and key physicochemical traits in nine wetland (aquatic) plant species native to the Western Ghats of Maharashtra, India. Both adaxial and abaxial surfaces were assessed for contact angles and also analyzed a set of traits including leaf length, width, length to width ratio, area, thickness, stomatal and trichome densities, water adhesion, surface structure, and surface free energy. Scanning electron microscopy (SEM) was employed to examine micro- and nano-scale surface features. This study aims to identify the key leaf traits influencing hydrophobicity and to investigate their role in facilitating adaptive strategies in wetland plant species.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cp\u003eUsing random sampling method for the selection, nine pond dwelling aquatic plants were selected for analyzing leaf wettability and the factors influencing it. The young and fully matured leaves after collecting were transported to the laboratory in the air tight packets so that they remain as fresh as possible. A total of 5 leaves were analyzed per plant species to determine the selected traits used in this study. Leaf wettability was determined measuring the contact angles of both adaxial and abaxial surfaces by the sessile drop method with the help of goniometer (rame hart goniometer). A 10 \u0026micro;l droplet of distilled water was deposited on the leaf surface employing a syringe and contact angle was determined by analyzing the image captured using CA meter software. For calculating surface free energy, we used 1-bromonapthalene as the nonpolar solvent. Using the Owens and Wendt (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1969\u003c/span\u003e) method where the total surface free energy is divided into two components: a dispersive (non-polar) component and a polar component, surface free energy was estimated by measuring the contact angles of at least two different liquids typically one polar (distilled water) and one non-polar (1-bromonapthalene) whose surface free energy is known on the solid surface. The formula to calculate SFE is as following:\u003cdiv id=\"Equa\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equa\" name=\"EquationSource\"\u003e\n$$\\:{\\gamma\\:}_{s}=\\:{\\gamma\\:}_{s}^{d}+\\:{\\gamma\\:}_{s}^{p}$$\u003c/div\u003e\u003c/div\u003e\u003cdiv id=\"Equb\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equb\" name=\"EquationSource\"\u003e\n$$\\:{\\gamma\\:}_{l}\\left(1+cos\\:\\theta\\:\\right)=\\:2\\:\\left.\\left(\\:\\sqrt{\\left\\{{\\gamma\\:}_{s}^{d}\\cdot\\:{\\gamma\\:}_{l}^{d}\\right\\}}+\\:\\sqrt{\\left\\{{\\gamma\\:}_{s}^{p}\\cdot\\:{\\gamma\\:}_{l}^{p}\\right\\}}\\right.\\right)$$\u003c/div\u003e\u003c/div\u003e\u003c/p\u003e \u003cp\u003e \u003cspan class=\"InlineEquation\"\u003e \u003cspan class=\"mathinline\"\u003e\\(\\:{\\gamma\\:}_{s}\\:\\)\u003c/span\u003e \u003c/span\u003e: Total surface free energy of the solid, ​\u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:{\\gamma\\:}_{s}^{d}\\)\u003c/span\u003e\u003c/span\u003e: Dispersive component of the solid's surface free energy, \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:{\\gamma\\:}_{s}^{p}:\\)\u003c/span\u003e\u003c/span\u003e Polar component of the solid's surface free energy, \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:{\\gamma\\:}_{l}\\)\u003c/span\u003e\u003c/span\u003e \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\::\\:\\text{T}\\text{o}\\text{t}\\text{a}\\text{l}\\:\\text{s}\\text{u}\\text{r}\\text{f}\\text{a}\\text{c}\\text{e}\\:\\text{f}\\text{r}\\text{e}\\text{e}\\:\\text{e}\\text{n}\\text{e}\\text{r}\\text{g}\\text{y}\\:\\text{o}\\text{f}\\:\\text{t}\\text{h}\\text{e}\\:\\text{l}\\text{i}\\text{q}\\text{u}\\text{i}\\text{d}\\)\u003c/span\u003e\u003c/span\u003e, \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:{\\gamma\\:}_{l}^{d}\\)\u003c/span\u003e\u003c/span\u003e: Dispersive component of the liquid's surface free, energy, \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:{\\gamma\\:}_{l}^{p}\\)\u003c/span\u003e\u003c/span\u003e: Polar component of the liquid's surface free energy, \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:\\theta\\:\\)\u003c/span\u003e\u003c/span\u003e: Contact angle of the liquid on the solid\u003c/p\u003e \u003cp\u003eFor surface structure analysis, air-dried leaf samples were placed on mounting stubs and gold sputtering is done, then imaged using a JEOL JSM-IT200 scanning electron microscope to assess structural differences in relation to contact angle (Pathan et.al. \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Leaf length and width were measured using a ruler, and fresh weight was recorded using a standard weight box. The length-to-width ratio was subsequently calculated. Leaf area was derived from length and width measurements, adjusted by a shape correction factor, following the method described by Schrader et al. (\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Leaf thickness was measured using a micrometer gauge. Leaf adhesion on both the adaxial and abaxial surfaces was determined using the formula \u003cb\u003eW = (W₁ \u0026ndash; W₀)/S\u003c/b\u003e, where \u003cem\u003eW₁\u003c/em\u003e represents leaf weight after wetting, \u003cem\u003eW₀\u003c/em\u003e is the weight before wetting, and \u003cem\u003eS\u003c/em\u003e denotes leaf\u0026rsquo;s surface area (Wang et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Trichome density (trichomes/mm\u0026sup2;) on both adaxial and abaxial surfaces was manually evaluated using a dissecting light microscope with observations taken from ten sites per leaf (Viz and Pacada \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Clear nail polish was thinly applied to both upper and lower leaf surfaces and left to dry. The dried film was peeled off with adhesive tape, mounted on slides, and examined under a light microscope to count stomata per mm\u0026sup2;(Zhu et al. \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). The parameters were analyzed for their correlation with the adaxial and abaxial contact angles to determine their effect on wettability. Pearson correlation coefficient was computed for each parameter against the contact angles of both surfaces, with statistical significance at p\u0026thinsp;\u0026lt;\u0026thinsp;0.05, with the help of Statistica 10 software (StatSoft Inc., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2011\u003c/span\u003e).\u003c/p\u003e"},{"header":"Results ","content":"\u003cp\u003eA total of nine wetland plant species belonging to three habitats i.e. submerged, floating, and emergent based on leaf\u0026rsquo;s position, were assessed for various morphological and surface traits in relation to leaf wettability, as quantified by contact angle (CA) measurements on both the leaf surfaces. On the adaxial surface, contact angles ranged between 61\u0026deg; and 143\u0026deg;, while on abaxial surface they varied from 47\u0026deg; to 124\u0026deg;, indicating a broad spectrum of wettability among the sampled species. Out of the nine species examined, only three i.e. \u003cem\u003eColocasia esculenta\u003c/em\u003e, \u003cem\u003eNelumbo nucifera\u003c/em\u003e from emergent habitat and \u003cem\u003ePistia stratiotes\u003c/em\u003e from floating habitat exhibited hydrophobic properties (CA\u0026thinsp;\u0026gt;\u0026thinsp;90\u0026deg;), whereas the remaining six species displayed hydrophilic surfaces (CA\u0026thinsp;\u0026lt;\u0026thinsp;90\u0026deg;). Additionally, in all but one species assessed, the adaxial surface showed contact angle more than the abaxial surface. All other physicochemical parameters were measured whose values are mentioned in the Table \u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e. Pearson\u0026rsquo;s correlation analysis was employed to evaluate the strength, direction and significance of the associations between leaf traits (e.g., width, trichome density, surface free energy) and contact angle values.\u003c/p\u003e \u003cp\u003eTable\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e shows the data of all the parameters studied. This data was then used to find out Pearson\u0026rsquo;s correlation coefficient values between the traits of leaf studied and the adaxial and abaxial contact angles measured at p\u0026thinsp;\u0026lt;\u0026thinsp;0.05, using Statistica 10 software.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eLeaf trait measurements of wetland plants used to assess correlations with contact angle and surface wettability:\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"20\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c12\" colnum=\"12\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c13\" colnum=\"13\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c14\" colnum=\"14\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c15\" colnum=\"15\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c16\" colnum=\"16\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c17\" colnum=\"17\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c18\" colnum=\"18\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c19\" colnum=\"19\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c20\" colnum=\"20\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePlant name\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHabitat\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eLength\u0026thinsp;\u0026plusmn;\u0026thinsp;SD (cm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eWidth\u0026thinsp;\u0026plusmn;\u0026thinsp;SD (cm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eL/W ratio\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eWeight\u0026thinsp;\u0026plusmn;\u0026thinsp;SD (g)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eThickness\u0026thinsp;\u0026plusmn;\u0026thinsp;SD(mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eArea (mm\u0026sup2;)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eAdaxial CA\u0026thinsp;\u0026plusmn;\u0026thinsp;SD (\u0026deg;)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003eAbaxial CA\u0026thinsp;\u0026plusmn;\u0026thinsp;SD (\u0026deg;)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c11\"\u003e \u003cp\u003eWater adhesion\u0026thinsp;\u0026plusmn;\u0026thinsp;SD (adaxial)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c12\"\u003e \u003cp\u003eWater adhesion\u0026thinsp;\u0026plusmn;\u0026thinsp;SD (abaxial)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c13\"\u003e \u003cp\u003eAdaxial CA\u0026thinsp;\u0026plusmn;\u0026thinsp;SD (\u0026deg;) nonpolar\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c14\"\u003e \u003cp\u003eAbaxial CA\u0026thinsp;\u0026plusmn;\u0026thinsp;SD (\u0026deg;) nonpolar\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c15\"\u003e \u003cp\u003eSFE adaxial (mj/m2)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c16\"\u003e \u003cp\u003eSFE abaxial (mj/m2)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c17\"\u003e \u003cp\u003eStomatal density adaxial (per mm\u0026sup2; )\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c18\"\u003e \u003cp\u003eStomatal density abaxial ( per mm\u0026sup2; )\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c19\"\u003e \u003cp\u003eTrichome density adaxial (per mm\u0026sup2;)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c20\"\u003e \u003cp\u003eTrichome density abaxial (per mm\u0026sup2;)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eColocasia esculenta\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eemergent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e18.68\u0026thinsp;\u0026plusmn;\u0026thinsp;0.67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e12.52\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1.49\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e3.768 \u0026plusmn; 0.19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.272\u0026thinsp;\u0026plusmn;\u0026thinsp;0.007\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e16371\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c9\"\u003e \u003cp\u003e121.25 \u0026plusmn; 2.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c10\"\u003e \u003cp\u003e95.55 \u0026plusmn; 1.52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.61\u0026thinsp;\u0026plusmn;\u0026thinsp;0.23 g/m\u0026sup2;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e1.22\u0026thinsp;\u0026plusmn;\u0026thinsp;0.45 g/m\u0026sup2;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c13\"\u003e \u003cp\u003e76\u0026thinsp;\u0026plusmn;\u0026thinsp;3.42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c14\"\u003e \u003cp\u003e45.3\u0026thinsp;\u0026plusmn;\u0026thinsp;2.90\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e17.18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e22.5\u0026thinsp;\u0026plusmn;\u0026thinsp;1.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e148.75\u0026thinsp;\u0026plusmn;\u0026thinsp;1.66\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eNelumbo nucifera\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eemergent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e15.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.64\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e18.46 \u0026plusmn; 0.88\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.83\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e6.84 \u0026plusmn; 0.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.318 \u0026plusmn; 0.007\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e22458\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c9\"\u003e \u003cp\u003e143.8 \u0026plusmn; 2.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c10\"\u003e \u003cp\u003e124.2 \u0026plusmn; 1.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e0.97\u0026thinsp;\u0026plusmn;\u0026thinsp;0.09 g/m\u0026sup2;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e1.33\u0026thinsp;\u0026plusmn;\u0026thinsp;0.31 g/m\u0026sup2;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c13\"\u003e \u003cp\u003e62.2\u0026thinsp;\u0026plusmn;\u0026thinsp;3.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c14\"\u003e \u003cp\u003e46.6\u0026thinsp;\u0026plusmn;\u0026thinsp;2.75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e28.75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e29.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e498.75\u0026thinsp;\u0026plusmn;\u0026thinsp;5.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTypha angustifolia\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eemergent\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e144.02 \u0026plusmn; 2.98\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e2.8 \u0026plusmn; 0.30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e51.43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e7.36 \u0026plusmn; 0.17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e1.296 \u0026plusmn; 0.018\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e25808\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c9\"\u003e \u003cp\u003e79.5 \u0026plusmn; 2.4\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c10\"\u003e \u003cp\u003e76.45 \u0026plusmn; 1.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e6.58\u0026thinsp;\u0026plusmn;\u0026thinsp;0.39 g/m\u0026sup2;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e7.05\u0026thinsp;\u0026plusmn;\u0026thinsp;0.44 g/m\u0026sup2;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c13\"\u003e \u003cp\u003e30.7\u0026thinsp;\u0026plusmn;\u0026thinsp;2.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c14\"\u003e \u003cp\u003e21.35\u0026thinsp;\u0026plusmn;\u0026thinsp;2.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e42.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e45.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e417.5\u0026thinsp;\u0026plusmn;\u0026thinsp;4.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e393.75\u0026thinsp;\u0026plusmn;\u0026thinsp;3.65\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eIpomoea aquatica\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003efloating\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e9.02 \u0026plusmn; 0.68\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e5.62 \u0026plusmn; 0.07\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e0.94 \u0026plusmn; 0.05\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.31 \u0026plusmn; 0.007\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e3549\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c9\"\u003e \u003cp\u003e75.15 \u0026plusmn; 0.58\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c10\"\u003e \u003cp\u003e47.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e14.09\u0026thinsp;\u0026plusmn;\u0026thinsp;0.42 g/m\u0026sup2;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e19.72\u0026thinsp;\u0026plusmn;\u0026thinsp;0.73 g/m\u0026sup2;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c13\"\u003e \u003cp\u003e16.9\u0026thinsp;\u0026plusmn;\u0026thinsp;1.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c14\"\u003e \u003cp\u003e27.2\u0026thinsp;\u0026plusmn;\u0026thinsp;1.63\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e47.09\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e59.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e156.25\u0026thinsp;\u0026plusmn;\u0026thinsp;2.21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e333.75\u0026thinsp;\u0026plusmn;\u0026thinsp;3.75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eNymphaea nouchali\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003efloating\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e21.96 \u0026plusmn; 1.59\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e18.6 \u0026plusmn; 0.316\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1.18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e11.49 \u0026plusmn; 0.54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.42 \u0026plusmn; 0.006\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e32268\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c9\"\u003e \u003cp\u003e63.1 \u0026plusmn; 0.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c10\"\u003e \u003cp\u003e52 \u0026plusmn; 1.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e13.94\u0026thinsp;\u0026plusmn;\u0026thinsp;0.83 g/m\u0026sup2;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e16.42\u0026thinsp;\u0026plusmn;\u0026thinsp;0.19 g/m\u0026sup2;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c13\"\u003e \u003cp\u003e44.05\u0026thinsp;\u0026plusmn;\u0026thinsp;2.79\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c14\"\u003e \u003cp\u003e32.35\u0026thinsp;\u0026plusmn;\u0026thinsp;2.94\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e46.18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e55.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e397.5\u0026thinsp;\u0026plusmn;\u0026thinsp;2.36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003ePistia stratiotes\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003efloating\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e11.12 \u0026plusmn; 0.23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e8.36 \u0026plusmn; 0.13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1.33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e7.86 \u0026plusmn; 0.13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.718 \u0026plusmn; 0.017\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e6322\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c9\"\u003e \u003cp\u003e95.64 \u0026plusmn; 0.58\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c10\"\u003e \u003cp\u003e113.4 \u0026plusmn; 0.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e36.38\u0026thinsp;\u0026plusmn;\u0026thinsp;0.72 g/m\u0026sup2;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e104.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.87 g/m\u0026sup2;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c13\"\u003e \u003cp\u003e73.1\u0026thinsp;\u0026plusmn;\u0026thinsp;3.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c14\"\u003e \u003cp\u003e51.2\u0026thinsp;\u0026plusmn;\u0026thinsp;2.56\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e21.68\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e29.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e65.2\u0026thinsp;\u0026plusmn;\u0026thinsp;2.52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e25.8\u0026thinsp;\u0026plusmn;\u0026thinsp;1.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e29\u0026thinsp;\u0026plusmn;\u0026thinsp;1.78\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eCryptocoryne retrospiralis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003esubmerged\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e23.12\u0026thinsp;\u0026plusmn;\u0026thinsp;0.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e1.48 \u0026plusmn; 0.074\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e15.62\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e1.63 \u0026plusmn; 0.028\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.292 \u0026plusmn; 0.011\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e2258\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c9\"\u003e \u003cp\u003e62.85\u0026thinsp;\u0026plusmn;\u0026thinsp;1.75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c10\"\u003e \u003cp\u003e54.6\u0026thinsp;\u0026plusmn;\u0026thinsp;1.77\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e10.62\u0026thinsp;\u0026plusmn;\u0026thinsp;0.76 g/m\u0026sup2;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e15.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.38 g/m\u0026sup2;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c13\"\u003e \u003cp\u003e45.6\u0026thinsp;\u0026plusmn;\u0026thinsp;2.58\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c14\"\u003e \u003cp\u003e16.2\u0026thinsp;\u0026plusmn;\u0026thinsp;2.71\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e46.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e56.9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e98.75\u0026thinsp;\u0026plusmn;\u0026thinsp;2.52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e91.25\u0026thinsp;\u0026plusmn;\u0026thinsp;1.46\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eEriocaulon cuspidatum\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003esubmerged\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e4.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.109\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e0.92 \u0026plusmn; 0.09\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e4.45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e0.208 \u0026plusmn; 0.007\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e1.14 \u0026plusmn; 0.048\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e268\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c9\"\u003e \u003cp\u003e84.2 \u0026plusmn; 1.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c10\"\u003e \u003cp\u003e58.5 \u0026plusmn; 2.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e52.24\u0026thinsp;\u0026plusmn;\u0026thinsp;0.33 g/m\u0026sup2;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e55.97\u0026thinsp;\u0026plusmn;\u0026thinsp;0.67 g/m\u0026sup2;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c13\"\u003e \u003cp\u003e11.1\u0026thinsp;\u0026plusmn;\u0026thinsp;1.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c14\"\u003e \u003cp\u003e9.7\u0026thinsp;\u0026plusmn;\u0026thinsp;3.22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e45.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e55.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e8.75\u0026thinsp;\u0026plusmn;\u0026thinsp;0.89\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eHygrophila auriculata\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003esubmerged\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e \u003cp\u003e6.24 \u0026plusmn; 0.35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e \u003cp\u003e1.24 \u0026plusmn; 0.048\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e5.03\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c6\"\u003e \u003cp\u003e0.22 \u0026plusmn; 0.015\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c7\"\u003e \u003cp\u003e0.31 \u0026plusmn; 0.011\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c8\"\u003e \u003cp\u003e511\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c9\"\u003e \u003cp\u003e61.8 \u0026plusmn; 1.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c10\"\u003e \u003cp\u003e60.05 \u0026plusmn; 1.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c11\"\u003e \u003cp\u003e29.41\u0026thinsp;\u0026plusmn;\u0026thinsp;0.51 g/m\u0026sup2;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c12\"\u003e \u003cp\u003e35.29\u0026thinsp;\u0026plusmn;\u0026thinsp;0.29 g/m\u0026sup2;\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c13\"\u003e \u003cp\u003e34.2\u0026thinsp;\u0026plusmn;\u0026thinsp;1.76\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c14\"\u003e \u003cp\u003e39.2\u0026thinsp;\u0026plusmn;\u0026thinsp;2.90\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c15\"\u003e \u003cp\u003e49.49\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c16\"\u003e \u003cp\u003e49.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c17\"\u003e \u003cp\u003e226.25\u0026thinsp;\u0026plusmn;\u0026thinsp;4.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c18\"\u003e \u003cp\u003e298.75\u0026thinsp;\u0026plusmn;\u0026thinsp;4.21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c19\"\u003e \u003cp\u003e7.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c20\"\u003e \u003cp\u003e4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.63\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eThe data in Table \u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e had been used to find out the correlation between the various leaf parameters and the measure of wettability i.e. contact angle. The correlation coefficient ( r ) values show how strong is the relationship between the two parameters taken into consideration.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eof every parameter against adaxial contact angles. of every parameter against abaxial contact angle.\u003c/p\u003e \u003cp\u003eFigures \u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e1\u003c/span\u003e and \u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e2\u003c/span\u003e show Pearson\u0026rsquo;s correlation matrix heatmaps representing the linear relationships between key leaf parameters and contact angles on the adaxial (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e1\u003c/span\u003e) and abaxial (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e2\u003c/span\u003e) surfaces. The color gradients reflect the strength (r-values) and direction of the correlations, with values ranging from \u0026minus;\u0026thinsp;1 (strong negative) to +\u0026thinsp;1 (strong positive).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eUsing these Pearson\u0026rsquo;s correlation coefficient values we found out the significance of all the parameters affecting the contact angles i.e. wettability of the leaves. The values for Pearson correlation coefficient and p- values which shows significance of the correlation are mentioned in the Table \u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003ePearson Correlation Coefficients (r) and p-values Between Leaf Parameters and Contact Angles.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eParameter\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003er-value (Adaxial)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003ep-value (Adaxial)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003er-value (Abaxial)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003ep-value (Abaxial)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSFE abaxial\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-0.975\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.000\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSFE adaxial\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003e-0.80\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.010\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWidth\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.561\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.116\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.480\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.191\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWater adhesion adaxial\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e-0.35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.365\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eStomatal density abaxial\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-0.36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.345\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eArea\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.277\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.471\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.264\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.493\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eL/W ratio\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e-0.235\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.543\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-0.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.782\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWeight\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.197\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.611\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.393\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.295\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWater adhesion abaxial\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.160\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.681\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eStomatal density adaxial\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.703\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eThickness\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e-0.111\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.776\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-0.010\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.979\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLength\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e-0.094\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.810\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.02\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.961\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTrichome density abaxial\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0.47\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0.203\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTrichome density adaxial\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.017\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.966\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"5\"\u003e\u003cb\u003eNote: Significant results (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05) are displayed in bold.\u003c/b\u003e\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eAmong all measured parameters, \u003cb\u003esurface free energy (SFE)\u003c/b\u003e showed the strongest correlations with contact angle. SFE on both adaxial and abaxial surfaces was \u003cb\u003enegatively correlated\u003c/b\u003e with contact angle values, indicating that leaves with lower surface energy exhibited higher water repellency. For example, \u003cem\u003eColocasia esculenta\u003c/em\u003e, with low abaxial SFE (33 mJ/m\u0026sup2;), showed high contact angles (adaxial: 121.25\u0026deg;, abaxial: 95.55\u0026deg;), suggesting strong hydrophobicity. Conversely, \u003cem\u003eIpomoea aquatica\u003c/em\u003e, which had the highest abaxial SFE (59.3) exhibited the lowest abaxial contact angle (47.3\u0026deg;), reflecting greater wettability. No consistent or significant correlations were found between contact angle and leaf length, width, thickness, area, or weight. In this study, trichome densities did not show significant associations with contact angle, likely due to their low or absent values across most species. But in case of \u003cem\u003ePistia stratiotes\u003c/em\u003e, a floating hydrophyte, the hydrophobic behavior is the result of presence of trichomes ( density more than 25 mm⁻\u0026sup2;) on both of its surfaces. Regarding stomatal density, no significant association between stomatal density and contact angle on either surface was found as per the Pearson correlation analysis.\u003c/p\u003e \u003cp\u003eScanning electron microscopy (SEM) was used to examine the microstructural features of both adaxial and abaxial leaf surfaces for all nine wetland plant species. The SEM images revealed distinct variations in surface texture, presence/absence of trichomes, stomatal exposure, and cuticular structures, which were analyzed in relation to contact angle measurements to infer their influence on leaf wettability.The two hydrophobic species, \u003cem\u003eColocasia esculenta\u003c/em\u003e and \u003cem\u003eNelumbo nucifera\u003c/em\u003e, showed specialized surface features that are consistent with high contact angle values. The surfaces of these leaves displayed a hierarchical surface architecture, including micro- and nanoscale roughness combined with dense epicuticular wax crystals. In \u003cem\u003eNelumbo nucifera (\u003c/em\u003eFig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e3\u003c/span\u003e; \u003cem\u003ecand d)\u003c/em\u003e, papillate epidermal cells and dense wax tubules were observed\u0026mdash;hallmarks of the \"lotus effect,\" which is known to promote superhydrophobicity by minimizing contact area between water and the surface (Barthlott and Neinhuis \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Koch et al. 2009). Similarly, the morphology of \u003cem\u003eColocasia esculenta (\u003c/em\u003eFig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e3\u003c/span\u003e; \u003cem\u003ea and b)\u003c/em\u003e, leaf exhibits a two-tier surface structure characterized by hexagonal microcavities resembling a honeycomb-like pattern (Kumar and Bhardwaj \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) with uniform wax coverage and elevated surface ridges that contribute to its water-repellent properties. The surface structures of \u003cem\u003ePistia stratiotes (\u003c/em\u003eFig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e3\u003c/span\u003e;\u003cem\u003ee and f)\u003c/em\u003e show the waxy trichomes that help bead up the water drops.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e The other six plants\u0026rsquo; contact angle in this study is below 90 degrees showing their affinity towards water. Their SEM analysis revealed surface structures consistent with those hydrophilic surfaces reported in previous studies.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe scanning electron micrographs (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e4\u003c/span\u003ea\u0026ndash;l) reveal consistent structural adaptations among the studied wetland species, supporting their hydrophilic surface behavior. Across all samples, the absence of micro- and nanoscale hierarchical features, including wax crystal deposition, correlates with low contact angles and increased wettability. In \u003cem\u003eNymphaea nouchali\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e4\u003c/span\u003ea, b), both leaf surfaces show an undulating and irregular epidermis with no epicuticular projections. This simple surface structure promotes close contact with water, enhancing hydrophilicity. Similarly, \u003cem\u003eCryptocoryne retrospiralis\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e4\u003c/span\u003ec, d) has an uneven surface with sunken stomata but lacks wax crystals, which likely aids in water absorption and gas exchange in moist or submerged environments. \u003cem\u003eIpomoea aquatica\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e4\u003c/span\u003ee, f) exhibits a smooth surface with fine cuticular striations, which may help water spread evenly while reducing air trapping. The absence of wax further supports its wettable nature. In \u003cem\u003eHygrophila auriculata\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e4\u003c/span\u003eg, h), the stomata are located in shallow depressions surrounded by smooth cuticular ridges, likely guiding water movement without blocking gas exchange. \u003cem\u003eEriocaulon cuspidatum\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e4\u003c/span\u003ei, j) shows a mostly uniform surface with slight striations near the stomata, consistent with water-retaining surfaces adapted for diffusion-based gas exchange. Lastly, \u003cem\u003eTypha angustifolia\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e4\u003c/span\u003ek, l) displays uneven, micropatterned surfaces without waxes, suggesting a structural adaptation that maintains hydrophilicity while providing surface stability in wet habitats.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThis study investigated the relationship between leaf surface traits and wettability in nine wetland plant species by analyzing contact angles on both adaxial and abaxial surfaces. The results revealed a broad spectrum of wettability, with only three species exhibiting hydrophobic characteristics and the remainder showing hydrophilic behavior. Surface free energy (SFE) was found to be the most consistent predictor of contact angle, while traits like stomatal and trichome density showed inconsistent or weak correlations. These findings provide insight into how structural and physicochemical leaf traits contribute to ecological adaptations in wetland environments.\u003c/p\u003e\n\u003ch3\u003eHydrophobicity and leaf positioning\u003c/h3\u003e\n\u003cp\u003eInterestingly, except \u003cem\u003ePistia stratiotes\u003c/em\u003e, the floating hydrophyte, whose hydrophobic behavior is attributed to the trichomes present on it, the other hydrophobic species are characterized by leaves that are held above the water surface rather than being submerged or floating. This leaf position suggests an ecological adaptation that favors water repellency, likely to facilitate self-cleaning (lotus effect) and prevent prolonged water retention, which can block gas exchange or promote microbial colonization (Barthlott and Neinhuis \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Koch et al. 2009). The elevated and hydrophobic leaves of \u003cem\u003eNelumbo nucifera\u003c/em\u003e are particularly well-known for their superhydrophobic nature, attributed to hierarchical surface structuring and wax crystals, which significantly reduce surface free energy and promote water rolling (Bhushan and Jung \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). Similarly, \u003cem\u003eColocasia esculenta\u003c/em\u003e has been reported to exhibit a highly water-repellent surface, potentially serving to maintain gas exchange and light capture in wetland conditions (Ensikat et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). These surface adaptations can be viewed as a response to the plants\u0026rsquo; aerial leaf posture, which exposes them to rain and surface splash, increasing the need for self-cleaning mechanisms (Barthlott et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2007\u003c/span\u003e).\u003c/p\u003e\n\u003ch3\u003eAdaptation of hydrophilic surfaces in aquatic plants\u003c/h3\u003e\n\u003cp\u003eIn contrast, the remaining hydrophilic species, often with submerged or floating leaves, do not exhibit such repellency. This observation supports earlier studies showing that there is reduced selective pressure for hydrophobic surfaces in aquatic or semi-aquatic environments where leaves remain consistently wet or underwater (Vogel \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e1988\u003c/span\u003e; Riedl and Riedl \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). In such species, hydrophilic surfaces may even aid in water absorption or nutrient exchange, depending on environmental conditions. This surface polarity in wettability is well-documented in foliage, often reflecting functional specialization. The adaxial surface, being more exposed to environmental factors like rainfall and sunlight, may evolve more hydrophobic traits to facilitate rapid droplet shedding and self-cleaning. The abaxial surface, typically more shaded and involved in gas exchange through stomata, may maintain a more hydrophilic profile to support physiological functions (Neinhuis and Barthlott \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Brewer \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1991\u003c/span\u003e).\u003c/p\u003e\n\u003ch3\u003eContribution of surface free energy to wettability\u003c/h3\u003e\n\u003cp\u003eThe inverse relationship between SFE and contact angle aligns with established principles of surface chemistry, where lower SFE facilitates water repellency (Bhushan and Jung \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). This relationship emphasizes the significance of surface physicochemical properties in determining leaf wettability across species with different ecological strategies.\u003c/p\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eTrichomes as determinants of water repellency\u003c/h2\u003e \u003cp\u003eHairy leaf surfaces, such as those of \u003cem\u003eAlchemilla vulgaris\u003c/em\u003e, are highly water-repellent because their stiff hairs keep water droplets elevated, preventing contact with the leaf surface and creating a fakir-like effect (Otten and Herminghaus \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). Brewer et al. (\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1991\u003c/span\u003e) found that plant species with dense trichomes (more than 25 mm⁻\u0026sup2;) tended to be more water-repellent, though some still retained droplets. Similarly, Neinhuis and Barthlott (\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1997\u003c/span\u003e) observed that leaves with wax-covered trichomes were extremely water-repellent, while those with non-waxy trichomes only resisted water briefly after droplet contact.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eInconsistent influence of stomatal density\u003c/h3\u003e\n\u003cp\u003eThis observation aligns with the findings of Lott (\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), who reported that stomatal density in five plant species did not significantly influence leaf wettability, possibly due to limited sample size or species-specific traits. All plants in Lott\u0026rsquo;s study had stomata only on the abaxial surface, as is typical in many plant species. However, this finding contrasts with studies by Brewer and Nu\u0026ntilde;ez (\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2007\u003c/span\u003e), who observed that surfaces with higher stomatal densities tended to be less wettable, possibly due to the microstructural complexity that encourages droplet retention. Unlike our findings, Wang et al. (\u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) found a positive correlation between stomatal density and contact angle in species studied in Xi\u0026rsquo;an, China\u0026mdash;suggesting that in some cases, higher stomatal density may increase hydrophobicity by contributing to surface microtexture or by trapping air, similar to the Cassie\u0026ndash;Baxter wetting regime. Taken together, these contrasting reports suggest that the influence of stomatal density on wettability is context-dependent and may be modulated by other surface features such as cuticular wax, trichomes, and surface roughness. In this study, the lack of a clear trend indicates that stomatal density alone may not be a reliable predictor of hydrophobicity across diverse wetland species.\u003c/p\u003e\n\u003ch3\u003eSurface structure of hydrophilic leaves\u003c/h3\u003e\n\u003cp\u003eHydrophilic leaf surfaces typically lack the complex micro- and nano-scale structures found on superhydrophobic leaves. They often appear smooth or slightly rough under SEM and show little to no wax crystalloids, which contributes to lower contact angles and increased water spreading (Neinhuis and Barthlott \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1997\u003c/span\u003e). The cuticle on these leaves may be thin, cracked, or composed of more polar compounds, allowing greater wettability (Riederer and Schreiber \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). In some cases, visible features such as stomata or trichomes can retain water, further promoting hydrophilic behavior (Brewer and Smith \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1997\u003c/span\u003e). These surfaces also tend to have higher surface free energy due to the absence of hydrophobic waxes (Koch et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). As a result, contact angles are typically below 90\u0026deg;, indicating that water readily spreads rather than beads up (Mittal and Davis \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2007\u003c/span\u003e).The collective absence of epicuticular wax crystals\u0026mdash;a key determinant of hydrophobicity\u0026mdash;across all species is a defining trait of these hydrophytes. These traits are not coincidental; rather, they represent evolutionary responses to persistently wet environments where maintaining a thin water layer on the leaf surface is advantageous. Such features may enhance nutrient absorption, support foliar respiration during submersion, and minimize damage from waterlogging (Rascio \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Sakamoto and Yamaguchi \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThese findings reinforce the view that hydrophilic leaf surface structures are ecologically advantageous in aquatic and semi-aquatic environments. They contrast starkly with the superhydrophobic adaptations found in xerophytic and canopy-dwelling species, where water repellence is vital for self-cleaning and pathogen resistance (Barthlott and Neinhuis \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Koch et al. 2009).\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eThis research undertaken examined the relationship between leaf\u0026rsquo;s physicochemical traits and wettability in hydrophytes living in three different ecological conditions i.e. emergent, floating, and submerged. Three plants from each were studied. Contact angle measurements revealed that only three species\u0026mdash;\u003cem\u003eNelumbo nucifera\u003c/em\u003e, \u003cem\u003eColocasia esculenta\u003c/em\u003e (emergent), and \u003cem\u003ePistia stratiotes\u003c/em\u003e (floating) exhibited hydrophobic behavior, while the remaining species displayed hydrophilic surfaces. Notably, the two emergent species had aerial leaves that avoid direct water contact, and the hydrophobicity of \u003cem\u003ePistia stratiotes\u003c/em\u003e was attributed to the presence of dense, waxy trichomes.\u003c/p\u003e \u003cp\u003eAll other plant species showed wetting behavior. Among the various morphological and anatomical leaf traits analyzed, surface free energy (SFE) emerged as the strongest predictor of contact angle. The inverse correlation between SFE and contact angle reinforces the importance of physicochemical properties over purely structural features in determining leaf wettability. All other leaf parameters studied i.e. leaf length, width, area, water adhesion etc. did not have stark impact in determining wettability. SEM analysis further showed that most hydrophilic species lacked micro- and nanoscale hierarchical surface features and epicuticular wax crystalloids\u0026mdash;structures typically associated with water repellency in terrestrial species. These findings suggest that leaf wettability in wetland plants is a complex trait shaped by both ecological context and adaptive function. While hydrophobic surfaces in emergent and floating species may aid in self-cleaning and gas exchange, hydrophilic surfaces in submerged or consistently wet leaves may facilitate foliar water uptake, nutrient absorption, or sustained physiological activity under saturated conditions. Thus, wettability patterns reflect functional adaptations aligned with each species\u0026rsquo; microhabitat and life form. This indicates that leaf wettability is closely linked to habitat-specific ecological pressures and contributes to the functional diversity of plant responses in wet environments. Further research is needed to explore how these traits interact with environmental variables across diverse wetland habitats.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003e\u003cstrong\u003eCompeting interests\u003c/strong\u003e:\u003c/h2\u003e\n\u003cp\u003eThe authors have no financial or nonfinancial interests to disclose.\u003c/p\u003e\n\u003ch2\u003eFunding:\u003c/h2\u003e\n\u003cp\u003eno funding\u003c/p\u003e\n\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\n\u003cp\u003eAll authors contributed significantly to the research and manuscript preparation. STB was responsible for conceptualization, methodology, and experimentation, literature review, drafting the original manuscript. MSN designed the experiments, collected data, and performed formal analysis. TDD conducted the curated data, and contributed to reviewing and editing the manuscript. RVM plant material collection, data validation, critical manuscript review. NVP handled final manuscript editing, proofreading, supervision, and correspondence. All authors reviewed and approved the final version of the manuscript.\u003c/p\u003e\n\u003ch2\u003eAcknowledgement\u003c/h2\u003e\n\u003cp\u003eThe authors are thankful to the Principal, The New College, Kolhapur, for providing the laboratory facilities. STB is also thankful to University Grants Commission, New Delhi, [ UGC Ref. No.: 835/ (CSIR-UGC NET JUNE 2019] for providing the financial assistance through UGC-JRF award for research work.\u003c/p\u003e\n\u003ch2\u003eData availability:\u003c/h2\u003e\n\u003cp\u003eThe datasets generated during and/or analyzed during the current study are available from the corresponding author on reasonable request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAllan, R. P. (2011). 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Quantitative analysis of stomatal density using nail polish impression method. \u003cem\u003eJournal of Plant Research\u003c/em\u003e, \u003cem\u003e131\u003c/em\u003e(3), 499\u0026ndash;508.\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"wetlands-ecology-and-management","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"wetl","sideBox":"Learn more about [Wetlands Ecology and Management](https://www.springer.com/journal/11273)","snPcode":"11273","submissionUrl":"https://submission.nature.com/new-submission/11273/3","title":"Wetlands Ecology and Management","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"contact angle, hydrophobicity, scanning electron microscopy, surface free energy, wettability","lastPublishedDoi":"10.21203/rs.3.rs-6731857/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6731857/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eLeaf wettability refers to the extent to which water adheres on leaf surfaces., which can have positive or negative effect on leaf, plant and ecosystem functioning. Hydrophytic plants have developed morphological adaptations to cope up with the excess presence of water. In this study, we analyzed contact angles and key physicochemical leaf traits including length, width, area, thickness, water adhesion, stomatal density, trichome density, surface structure, and surface free energy of nine wetland plant species belonging to three different habitats i.e. floating, submerged and emergent, on the basis of leaf position, from the Western Ghats of Maharashtra. Measurements were taken for both the adaxial and abaxial surfaces to determine the correlations between these traits and leaf wettability. Among the examined traits, surface free energy emerged as the primary determinant of hydrophobicity, exhibiting a strong negative correlation with contact angle. Contact angles ranged from 61\u0026deg; to 143\u0026deg; on the adaxial surface and from 47\u0026deg; to 124\u0026deg; on the abaxial surface. While most other physical traits did not show significant correlation, the presence of trichomes in \u003cem\u003ePistia stratiotes\u003c/em\u003e a floating hydrophyte enabled the formation of water beads on its surface. Additionally, Scanning Electron Microscopy (SEM) was employed to investigate the microstructural and nano-structural features of both sides of the leaf, providing further insights into their role in wettability. Studying the adaptations of the hydrophytes to their environment reveals that incorporating wettability and nano structural surface traits, along with other plant characteristics, enhances our understanding of plant\u0026ndash;water interactions in wetland ecosystems.\u003c/p\u003e","manuscriptTitle":"Decoding Leaf Wettability: The Role of Physicochemical Leaf Traits in Hydrophytic Plants","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-05-30 17:52:33","doi":"10.21203/rs.3.rs-6731857/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-06-09T06:55:54+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-06-09T01:01:48+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-06-04T05:14:38+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"152004445390294785431169767377302621599","date":"2025-06-01T22:34:53+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"283298310272579688603060206644835459839","date":"2025-05-29T23:40:19+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"89337243786040595724112238787330137650","date":"2025-05-28T05:06:19+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-05-27T22:16:30+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-05-27T22:13:10+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-05-26T04:34:56+00:00","index":"","fulltext":""},{"type":"submitted","content":"Wetlands Ecology and Management","date":"2025-05-23T10:05:57+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"wetlands-ecology-and-management","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"wetl","sideBox":"Learn more about [Wetlands Ecology and Management](https://www.springer.com/journal/11273)","snPcode":"11273","submissionUrl":"https://submission.nature.com/new-submission/11273/3","title":"Wetlands Ecology and Management","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"1c9245bf-cfb9-45c2-8df1-ebe6097618dd","owner":[],"postedDate":"May 30th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2026-02-23T16:00:28+00:00","versionOfRecord":{"articleIdentity":"rs-6731857","link":"https://doi.org/10.1007/s11273-026-10119-x","journal":{"identity":"wetlands-ecology-and-management","isVorOnly":false,"title":"Wetlands Ecology and Management"},"publishedOn":"2026-02-19 15:57:07","publishedOnDateReadable":"February 19th, 2026"},"versionCreatedAt":"2025-05-30 17:52:33","video":"","vorDoi":"10.1007/s11273-026-10119-x","vorDoiUrl":"https://doi.org/10.1007/s11273-026-10119-x","workflowStages":[]},"version":"v1","identity":"rs-6731857","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6731857","identity":"rs-6731857","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}