Journal of Experimental Botany, Vol. 54, No. 385, pp.

1295±1303, April 2003

DOI: 10.1093/jxb/erg127

RESEARCH PAPER

Quantitative assessment to the structural basis of water

repellency in natural and technical surfaces

P. Wagner, R. FuÈrstner, W. Barthlott and C. Neinhuis1

Botanisches Institut und Botanischer Garten, Rheinische Friedrich-Wilhelms-UniversitaÈt Bonn, Meckenheimer
Allee 170, D-53115 Bonn, Germany

Received 5 June 2002; Accepted 10 January 2003

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Abstract Introduction
Many plant surfaces are water-repellent because of a The surfaces of plants, especially those of leaves, exhibit a
complex 3-dimensional microstructure of the epider- great number of structural types that have been studied and
mal cells (papillae) and a superimposed layer of classi®ed in detail by scanning electron microscopy in the
hydrophobic wax crystals. Due to its surface tension, last 30 years (reviews in Baker and Parsons, 1971;
water does not spread on such surfaces but forms Barthlott, 1981, 1990; Barthlott and Ehler, 1977;
spherical droplets that lie only on the tips of the Barthlott and Wollenweber, 1981; Bukovac et al., 1981;
microstructures. Studying six species with heavily Holloway and Baker, 1974; Jeffree, 1986). Three general
microstructured surfaces by a new type of confocal levels of structuring have been identi®ed in this respect:
light microscopy, the number, height, and average the general cell's shape (primary-), cuticular folds (sec-
distance of papillae per unit area were measured. ondary-) and epicuticular wax crystals (tertiary-structure).
These measurements were combined with those of Surface structures are also important in a functional
an atomic force microscope which was used to respect. Water-repellency has been of particular interest
measure the exposed area of the ®ne-structure on because it is a major factor in spray application processes
individual papillae. According to calculations based (Baker et al., 1983; Boize, 1976; Bukovac et al., 1979;
upon these measurements, roughening results in a Kadota and Matsunaka, 1986; Kuzych and Meggitt, 1983;
reduction of the contact area of more than 95% com- Watanabe and Yamaguchi, 1991b; Wirth et al., 1991;
pared with the projected area of a water droplet. By Zabkiewicz et al., 1988). Therefore, the wettability of leaf
applying water/methanol solutions of decreasing surfaces has been studied in detail from the beginning of
surface tension to a selection of 33 water-repellent the last century (Boyce and Berlyn, 1988; Crisp, 1963;
species showing different types of surface struc- Engel, 1939; Fogg, 1947; Hall and Burke, 1974; Holloway,
tures, the critical value at which wetting occurs was 1970; Linskens, 1950; Moilliet, 1963; Rentschler, 1971;
determined. The results impressively demonstrated Watanabe and Yamaguchi, 1991a; Ziegenspeck, 1942).
the importance of roughening on different length From many studies it became apparent that a particular
scales for water-repellency, since extremely papillose microroughness, especially due to epicuticular wax crys-
surfaces, having an additional wax layer, are able to tals, is the structural basis of extreme water-repellency of
resist up to 70% methanol. Surfaces that lack papillae surfaces. Due to their surfaces tension water droplets form
or similar structures on the same length scale are spheres lying only on the tips of the structures. The
much more easily wetted. principal connections between surface roughness and
water-repellency were worked out by Cassie and Baxter
(1944), as well as Wenzel (1936). Later, the wetting
Key words: Confocal microscopy, epicuticular wax, plant properties of surfaces were subject to intensive studies in
cuticle, water-repellency. physics, as well as in biology, and reviewed several times
1
Present address and to whom correspondence should be sent: Botanisches Institut, Technische UniversitaÈt Dresden, D-01069 Dresden, Germany.
Fax: +49 351 463 37032. E-mail: [email protected]

Journal of Experimental Botany, Vol. 54, No. 385, ã Society for Experimental Biology 2003; all rights reserved
1296 Wagner et al.
(Adam, 1963; Adamson, 1990; Bico et al., 1999; Arti®cial surfaces
de Gennes, 1985; Holloway, 1969, 1970). Nowadays, In addition to the plant surfaces, several arti®cial microstructured
water-repellency has gained much interest because it hydrophobic surfaces were tested. They can be divided into
(a) replicates of leaf surfaces (Fig. 1 f) and (b) technical surfaces
represents the basis for a self-cleaning property of such made of metal with electrochemically deposited microstructures
surfaces called the `lotus effect' (Barthlott and Neinhuis, (Fig. 1g, h).
1997) which can also be transferred into technical Replicates were made of a negative form worked out with a
applications (further informations see: www.lotus-effect. 2-component silicone moulding mass (President light body, ColteÁne,
Switzerland). After drying, the negative is ¯exible and rubber-like.
com). Into this a conventional lacquer (Acryllack, seidenmatt weiû, Karl
In an earlier study of about 300 plant species, Neinhuis Knauber, Germany) or a liquid polymer (polyether, ZK 2068-026,
and Barthlott (1997) showed that a wide range of different BASF, Germany) was ®lled, which resulted in an almost perfect
surface morphologies is suitable to serve as a basis for a replicate of a leaf's surface up to details in the range of 1 mm when
water-repellent and self-cleaning surface. Although, intui- dry. Epicuticular wax crystals could not be replicated because of
their poor mechanical stability.
tively, some species appeared to be more optimized to non- The metal surfaces used for the expermiments are fabricated for
wetting (especially those with prominent epidermal printed electronic circuits representing copper-foils with a smooth

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papillae covered with an additional layer of wax crystals) upper surface and a heavily structured lower surface (Bolta,
conventional contact angle measurements did not allow Germany, Circuit Foil, Luxemburg). The surfaces of some of these
samples are strongly reminiscent of a microstructured leaf surface.
any clear differentiation between individual morphologies. Becromal (Frolyt, Freiberg, Germany) is an electrochemically
In addition, the real contact area between a water droplet microstructured aluminium surface used for condensator manufac-
and a rough surface has never been determined in an turing.
experimental approach, although it is generally accepted Both replicates and metal foils were gold sputtered for 40 s and
that the reduction in contact area and the enclosure of hydrophobized with 1-hexadecanethiol (Merck-Schuchard,
Hohenbrunn, Germany)/heptane (Carl Roth, Karlsruhe, Germany).
air between microstructures is the most important factor The metal surfaces were additionally hydrophobized with Dynasilan
to induce non-wetting (Dettre and Johnson, 1964; (Sivento, Rheinfelden, Germany) or Antispread (Dr Tillwich, Horb-
Herminghaus, 2000; Holloway, 1970). Ahldorf, Germany), both are ¯uorinated agents. In Dynasilan a SiO-
Therefore a new approach to address both questions was group adheres to the surface while the molecule's F3C-end is
exposed to the air. Antispread polymerizes on the surface.
adopted. The ®rst approach is based on the assumption that Hexadecanethiol is a non-¯uorized agent, which the SH-group
certain morphologies are better adapted to non-wetting, i.e. binds to the gold atoms of the surface while exposing the
if the surface tension of a liquid is reduced, some species hydrophobic CH3-group.
should be wetted more easily than others. To address this
question, a range of species was tested against water± Specimen preparation
methanol mixtures with decreasing surface tension to Samples of the leaf surfaces were cut into 535 mm in size avoiding
determine the critical value at which wetting occurs. the leaf's veins, dehydrated and ®xed according to the liquid-
substitution method (Ensikat and Barthlott, 1993), allowing long-
In a second approach, an attempt was made to determine term investigation of the prepared plant specimens with CLM or
the contact area between a water droplet and a rough AFM. In addition, SEM investigation of the same sample is possible
surface base compared with the projected area of the for more than 30 min.
droplet. This was done by measuring and calculating the The samples were ®xed to microscope-slides for CLM-investiga-
tion or an AFM-specimen holder using 2-component epoxide-glue
exposed area of differently structured surfaces combining (UHU plus schnellfest, UHU, BuÈhl, Germany), sputter-coated (SCD
confocal white light microscopy and atomic force micro- 034, Balzers Union, Wiesbaden, Germany) with gold of approxi-
scopy, since the minimization of the contact area was mately 10 nm thickness in order to increase electrical conductivity
supposed to be one of the major reasons for extreme water- (SEM) or light re¯ection (CLM).
repellency.
Wetting with water±methanol solutions
In order to ®nd out differences between surfaces that are water-
repellent (contact angles >150°), but show differently microstruc-
tured surfaces, water±methanol mixtures were applied. Methanol is a
Materials and methods suitable medium because it is miscible with water in any concen-
tration, lowers the surface tension (Fig. 2) and does not alter the wax
Plant material
ultrastructure, which has been demonstrated earlier (Neinhuis and
All plants were taken from the Botanical Garden of the University of Edelmann, 1996). The specimens have been ®xed to a tilted surface
Bonn (Germany) and represent a selection of microstructured plants (q=25°) with double-sided adhesive tape to allow droplets to bounce
(Table 1) based on the list published by Neinhuis and Barthlott off the surface. A single use syringe (vol=10 ml) has been attached to
(1997). For the combined investigation with confocal light a tripod in a way that the tip of the drain tube (inner diameter: 0.70
microscopy (CLM) and atomic force microscopy (AFM) six species mm; Braun Sterican Gr. 12, 0.70330 mm) was ®xed 10 mm above
(marked in bold in Table 1) were chosen due to their prominent the specimen surface. The methanol was added to distilled water,
primary sculpture and extreme water-repellency (Fig. 1a±e). In total, weight in mass-percentage, in steps of 5%. Values of the critical
31 species were taken without further preparation for the wetting test surface tension were taken as follows: individual droplets of water±
with methanol±water mixtures. methanol mixtures (approximately 10 ml) were dropped onto the test
 Wetting of structured surfaces 1297
Table 1. list of plants (accession numbers of the Botanical Garden, Bonn, are given) treated with water±methanol mixtures
*, Abaxial leaf side only; bold, measured with the CLM and AFM; (+), plant has been grown for this investigation.
Species Family Acc. no. Primary structure Epicuticle wax type
Acacia dealbata Link Mimosaceae 00146 papillose tubules
Alchemilla mollis (Buser) Rothm. Rosaceae 10092 trichomes rodlets
Alocasia macrorrhiza (L.) G. Don. Araceae 01194 papillose* wax®lm
Apocynum cannabinum L. Apocynyceae 06118 convex platelets
Argemone mexicana L. Papaveraceae 03318 convex tubules
Berberis gagnepainii Schneid. Berberidaceae 04155 convex tubules
Berberis julianae Schneid. Berberidaceae 01914 convex tubules
Berberis verruculosa Hemsl. & Wils. Berberidaceae 12285 convex tubules
Brassica oleracea L. Brassicaceae + smooth dendritic rodlets
Chondrilla juncea L. Asteraceae 03411 trichomes platelets
Colocasia esculenta (L.) Schott. Araceae 04069 papillose platelets
Coronilla coronata L. Fabaceae 03151 convex tubules
Crambe maritima L. Brassicaceae 03333 smooth polymorph

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Daphniphyllum humile Maxim. Daphniphyllaceae 00515 papillose tubules
Dicentra formosa (Haw.) Walp. Fumariaceae 03303 convex tubules
Drimys winteri Forst. & Forst. Winteraceae 00768 convex tubules
Eucalyptus macrocarpa Hook Myrtaceae 00612 convex tubules
Euphorbia atropurpurea Brouss. Euphorbiaceae 13695 convex platelets
Euphorbia characias L. Euphorbiaceae 14076 convex platelets
Euphorbia myrsinites L. Euphorbiaceae 08048 papillose platelets
Ginkgo biloba L. Ginkgoaceae 01894 convex tubules
Hebe albicans (Petrie) Ckn. Scrophulariaceae 00728 papillose platelets
Iris japonica Thunb. Iridaceae 00283 convex platelets
Liriodendron chinense (Hemsl.) Sarg. Magnoliaceae 15354 papillose* transv. ridged rodlets
Marsilea drummondii A.Br. Marsileaceae 00225 convex platelets
Nelumbo nucifera (Willd.) Pers. Nelumbonaceae 11705 papillose tubules
Neptunia plena (L.) Benth. Mimosaceae 15656 papillose platelets
Oryza sativa L. Poaceae 08616 papillose platelets
Papaver atlanticum (Ball) Coss. Papaveraceae 03315 papillose tubules
Thalictrum ¯avum (Desf.) Battand. Ranunculaceae 02700 convex tubules
Tropaeolum majus L. Tropaeolaceae + convex tubules
Xanthosoma nigrum (Vell.) Mansf. Araceae 16126 papillose* platelets
Xanthosoma spec. Araceae 01070 papillose* platelets

surface according to Fig. 3 and (a) bounced off if the critical surface Measurement techniques
tension was not reached or (b) partially wetted the test surface and In order to measure the surface's primary sculpture from a leaf by
got stuck if the critical surface tension was reached. Those values are CLM usually 10 (20 on Alocasia macrorrhiza) epidermal papillae
presented in Fig. 7. Convergently to the surface tension, the overall have been selected and surveyed in X-, Y- and a diagonal direction
mass of the droplets decreased from 10.4 mg (pure distilled water) to over the papilla's centre. Each survey results in a 2D pro®le, which
3.8 mg (solution with about 90% methanol) due the lower density of has been used for measuring the papilla's height from top to bottom.
methanol. The experiments have been carried out at room tempera- From each species two digital images have been surveyed. The
ture (approximately 23 °C). On the basis of preceding tests contact results of 20 individual papillae have been statistically averaged and
angles were not measured because methanol evaporates too fast from the standard deviation (sd) has been calculated. To investigate the
small droplets and therefore no standardized conditions could be papillas' lateral distances a free survey-line has been laid from top to
established. top, for example, the highest point in the image. About ®ve to eight
neighbouring cells surrounding a central papilla have been measured
Microscopy and the results have been statistically calculated as mentioned above.
The CLM used is a recently developed new type of white light From A. macrorrhiza only one digital image was available, so 20
confocal microscope (m-surf, NanoFocus, Duisburg, Germany) epidermal papillae were surveyed from this image. Each sample of
which has been designed speci®cally for the 3D-investigation of the other species has been measured as described above.
microstructures (Fig. 4). The digital images were analysed by A square of usually 20 mm has been scanned by AFM (Fig. 5).
NanoFocus software. More detailed information is given by Jordan These images have been analysed using the bearing-application
et al. (1998). included in the microscope's software (NanoScope IIIa, vers.
A standard AFM (NanoScope IIIa, Digital Instruments, 4.23r3). Bearing analysis reveals which area of an investigated
Mannheim, Germany) was used applying the tapping-mode. The surface lies above or below any arbitrarily chosen height. A level of
needles used were one-piece and made of silicon, type: Nanoprobe 1.0 was chosen, 0.5 mm below the top of the papillae: these two
SPMÔ, TESP, L=125 mm, F0=298±312 kHz. heightsÐor better depthsÐwere chosen according to the presumed
All specimens have been additionally investigated by SEM elastic deformation of a virtual water droplet of two different masses.
(Cambridge Stereoscan 200, Oxford, UK; LEO 440, LEO, Such a water droplet lying on a few tips of a microstructured surface
Oberkochen, Germany). is deformed by its own weight, but held in shape by its surface
1298 Wagner et al.

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Fig. 1. SEM photographs of microstructured water-repellent surfaces. (A) Nelumbo nucifera adaxial leaf surface (ad), (B) Liriodendron chinense
abaxial leaf surface (ab), (C) Euphorbia myrsinites ad, (D) Colocasia esculenta ad, (E) Alocasia macrorrhiza ab, (F) replicate of A. macrorrhiza,
(G) copper-foil Bolta 18 mm B0, (H) copper-foil circuit foil 35 mm NT-TO.

tension. So it bulges its lower part to a certain depth into the Results and discussion
microstructure (Fig. 6), lying on the secondary or tertiary structure of
the epidermal papillae. According to this deformation the contact The results of the wetting tests with methanol±water
area was calculated according to the two different bearing heights. mixtures are given in Fig. 7. Depending on the amount of
 Wetting of structured surfaces 1299

Fig. 2. Surface tension of water±methanol mixtures. Increasing the

mass-percentage of methanol in water the surface tension of the
mixtures is reduced by about 66%.

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Fig. 4. Confocal light microscopic picture of the abaxial leaf surface
of Alocasia macrorrhiza used for measuring average distance and
height of epidermal papillae. The numbers in the corners indicate the
length scale, the grey scale indicates the height of papillae.

Fig. 3. Testing device for the application of individual droplets of

water±methanol mixtures on different plant and technical surfaces.

structuring, individual plant surfaces repel mixtures

containing up to 75% methanol, in a single measurement
of C. esculenta up to 80% (not indicated in the ®gure).
There is a gradient in the cell shape directly related to the
repellency from left to right. While all Berberis species as
well as Alchemilla mollis exhibit ¯at or only slighly
convex outer epidermal cell walls (apart from the layer of
wax crystals that was present in all species), those species Fig. 5. Atomic force microscopic picture of the tip of an individual
with high methanol repellency (for example, Nelumbo papilla of Alocasia macrorrhiza leaves showing the ®ne structure
nucifera, Colocasia esculenta) are characterized by very based on cuticular folds.
pronounced papillae (Fig. 1a, d). On the other hand there is
no evidence that high methanol resistance depends on a columns indicate the occurence of small wax tubules
speci®c wax chemistry and micromorphology. The grey composed of the secondary alcohol nonacosan-10-ol
1300 Wagner et al.
which are found among surfaces characterized by different effective, the replicate of the leaf surface of A. macrorrhiza
amounts of structuring at the cell level and therefore of shows a considerably lower methanol resistance as com-
different methanol resistance. pared to the natural surface (30% versus 70% of the leaf).
In addition to the plant surfaces some arti®cial surfaces In earlier papers dealing with water repellency, the
have been tested. The percentage values up to which a surface roughness has attracted much attention, because it
water±methanol droplet was repelled from the surface are is obviously the reason for contact angles close to 180°
shown in Table 2. the very high values of Bolta 18 mm (Bico et al., 1999; Busscher et al., 1984; Cassie and
BO-foil (Fig. 1g), which show a methanol-tolerance of up Baxter, 1944; Dettre and Johnson, 1964; Holloway, 1970;
to 90%, are remarkable, in that only almost pure methanol Wenzel, 1936). In these papers the actual kind of
wets the surface. The results of the other arti®cial surfaces roughness has been recorded only as long as they were
are within the range of the tested plants. However, periodic, for example, grids or columns or were supposed
although the arti®cial hydrophobization should be more to be fractal (Onda et al., 1996; Shibuichi et al., 1996,
1998). Natural surfaces rarely show periodic structures,
some have been regarded as self-af®ne, but generally they

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exhibit a random distribution of structures on different
length scales. On the other hand, investigations of water-
repellent surfaces that included a detailed description of
the structures, were not correlated with the wetting
properties since contact angle measurements of the water
droplets did not allow differentiation between individual
topologies (Holloway, 1970; Neinhuis and Barthlott,
1997). The results presented here show the effectiveness
of combined structures which have been theoretically
postulated earlier (Herminghaus, 2000). In particular, the
formation of papillae is a good design to increase water-
Fig. 6. The drawing visualizes schematically the contact lines of two
virtual water droplets with different masses on a papillose surface repellency. This may be due to the fact that waxes in
used to calculate the actual contact areas in relation to the projected general are not very hydrophobic because of many
areas of the droplets. hydrophilic functional groups (e.g. ±OH, ±COOH,

Table 2. Methanol resistance of hydrophobic technical surfaces

Note that the Bolta copper surface shows very high values while replicates of the lower leaf surface of Alocasia macrorrhiza are considerably
lower than the original leaf surface independent of the type of hydrophobization.
Gold Thiol Antispread Dynasilan
CF 35 mm NT-TO 25% ± 30%
CF 35 mm NT-TW HTE 25% 50% 20%
Becromal CD888F 86V 25% ± 20%
Bolta 18 mm BO 70% >90% 80%
Replicate made of Polyether ± ± ± 30%a
BASF ZK 2068-026
Replicate made of Knauber 30% ± ±
acryllaquer
a
Not hydrophobized due to the hydrophobic nature of the polymer.

Table 3. Average number of papillae mm±2 and the calculated remaining contact area
The calculated contact area of a virtual water droplet of two different masses, modelled by the two bearing depths, is given in (%) in relation to
a smooth surface of 1 mm2. All values are average (except A. macrorrhiza).
Species Papillae on 1 mm2 Contact area (%) according to a depth (d) of

(d)=1.0 mm (d)=0.5 mm
A. macrorrhiza 2002 5.13 1.50
C. esculenta 2662 7.20 1.71
E. myrsinites 1265 2.74 0.41
L. chinense 737 4.88 1.44
N. nucifera 3431 6.95 1.77
X. spec 967 6.40 1.12
 Wetting of structured surfaces 1301
±CHO). As demonstrated by Herminghaus (2000), speci®c structures have an enormous in¯uence on the wetting
combinations of structures on different length scales can properties, which can be seen from those experiments
result in water-repellent surfaces (not ultra-phobic) where the same type of hydrophobic chemical has been
although the intrinsic contact angle of a speci®c chemical applied. However, the contradictory behaviour of the
on the surface is below 90°. From the biological point of replicates from leaves of A. macrorrhiza still remains to be
view, an explanation for the evolution of such `over- explained.
optimized' surfaces may be found in the fact, that plants In the second approach the surface structures of six of
(especially under wetland conditions) often have to deal the most extremely water- and methanol-repellent plant
with water that may be polluted with oily or amphiphilic surfaces have been investigated by CLM and AFM. The
substances. These substances originate from decaying papillae representing a length scale of some 10 mm show
organic matter and may decrease the surface tension. To an aspect ratio (height/distance) smaller than one (Fig. 8).
achieve this goal, water-repellency caused by wax crystals Statistically tested, there is no correlation between this
in the range of 0.5±5 mm seems not to be suf®cient, but parameter and the resistance against high methanol
needs an additional surface structure in the range of 20±50 concentrations (X versus Y scatter plot: r2=0.0071).

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mm, which is provided by the shape of the outer epidermal Using the digital images of the CLM it was possible to
cell walls. Other reasons for developing water-repellent calculate the average number of papillae mm±2 (Table 3). It
surfaces have been discussed by Neinhuis and Barthlott is remarkable that, although all surfaces are water-repel-
(1997). lent, the papillae density varies between 737 and 3431. At
The resistance of up to 90% methanol in the case of one the same time, the aspect ratio does not show such a
of the copper foils may be explained by the combination of variation. In X. spec. and L. chinense with comparatively
a highly effective surface structure that is hardly distin- high skittle-like papillae (27.8 and 38.6 mm) the cells
guishable from plant surface, together with a ¯uorinated correspondingly have a greater lateral distance compared
hydrophobic chemical that minimizes the surface energy. with those with lower papillae.
As in the plant surfaces, the metals show a similar These results are in good accordance with the theoretical
behaviour in that respect, that individual combinations of background about water-repellency of microstructured

Fig. 7. Resistance of leaf surfaces against wetting with water±methanol mixtures. Leaf surfaces without papillose epidermal cells (on the left) are
more easily wetted than those with prominent papillae (right). Grey columns mark wax tubules composed of nonacosan-10-ol indicating that the
high methanol resistance is independent of the individual ®ne structure of the wax layer but mainly depends on the sculpturing of the outer
epidermal cell wall.
1302 Wagner et al.

Fig. 8. Aspect ratio of the epidermal cells of six highly non-wettable Fig. 9. Papillae density (1 mm±2) in relation to methanol resistance.

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plant surfaces compared to the highest repelled methanol- Surfaces with a high amount of smaller papillae show a better
concentration showing no obvious correlation. resistance than those with lower numbers. Note that A. macrorrhiza
(wax ®lm on heavily structured secondary structure) is equally
resistant as those surfaces characterized by wax crystals.

surfaces that has been discussed in detail elsewhere (Dettre

and Johnson, 1964; Herminghaus, 2000; Holloway, 1970). that a well designed water-repellent surface is optimized
A very thin air-layer trapped in the depressions between only in a narrow range of aspect ratios and papillae
the papillae is regarded to be the main reason for water- densities. Individual morphologies of contributing struc-
repellency. Increasing the distance between the papillae on tures are less important to achieve this goal as can be seen
the one hand increases the water/air contact area. At the from the surface of A. macrorrhiza. Here, water-repellency
same time, the water droplet may penetrate more easily is based on papillose epidermal cells and cuticular folds.
into the depressions by its own weight, high impact The latter are covered by a thin amorphous wax ®lm and
velocity or due to a reduced surface tension and wets the not by wax crystals. The number of papillae as well as
surface. Increasing the height of the papillae decreases the aspect ratios and the contact area are comparable to the
risk because the droplet dramatically increases its surface other species and therefore the values for methanol
before reaching the bottom between the papillae. resistence are in the same range.
Looking at the number of papillae per unit area shows
only a weak correlation (r2=0.7213) with the methanol
resistance of a particular surface (Fig. 9) due to the small Acknowledgements
amount of data. However, a higher amount of smaller
The authors gratefully acknowledge funding of the project by the
papillae seems to be more effective in terms of water- Deutsche Bundesstiftung Umwelt, technical support and the per-
repellency than surfaces with larger but less numerous mission to use the confocal light microscopy by NanoFocus.
papillae.
Furthermore, the contact area of a water droplet on the
tips of the epidermal papillae in relation to the projected References
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