Downloaded from http://rsif.royalsocietypublishing.

org/ on March 11, 2015

rsif.royalsocietypublishing.org

Removal mechanisms of dew via

self-propulsion off the gecko skin
Gregory S. Watson1, Lin Schwarzkopf2, Bronwen W. Cribb3, Sverre Myhra4,
Marty Gellender5 and Jolanta A. Watson1

Research
Cite this article: Watson GS, Schwarzkopf L,
Cribb BW, Myhra S, Gellender M, Watson JA.
2015 Removal mechanisms of dew via
self-propulsion off the gecko skin. J. R. Soc.
Interface 12: 20141396.
http://dx.doi.org/10.1098/rsif.2014.1396

Received: 22 December 2014

Accepted: 16 February 2015

Subject Areas:
biomaterials, biophysics
Keywords:
lizard, gecko, condensation, nanostructures,
dew, contaminants

Authors for correspondence:

Gregory S. Watson
e-mail: gwatson1@usc.edu.au
Jolanta A. Watson
e-mail: jolanta.watson@griffith.edu.au

Electronic supplementary material is available

at http://dx.doi.org/10.1098/rsif.2014.1396 or
via http://rsif.royalsocietypublishing.org.

School of Science and Engineering, University of the Sunshine Coast, Maroochydore DC, Queensland 4558,
Australia
2
School of Marine and Tropical Biology, James Cook University, Townsville, Queensland 4811, Australia
3
Centre for Microscopy and Microanalysis and School of Biological Sciences, The University of Queensland,
St Lucia, Queensland 4072, Australia
4
The University of Oxford, Begbroke Science Park, Sandy Lane, Yarnton OX5 1PF, UK
5
Previously Queensland Department of Environment and Heritage Protection, GPO Box 2454, Brisbane,
Queensland 4001, Australia
Condensation resulting in the formation of water films or droplets is an unavoidable process on the cuticle or skin of many organisms. This process
generally occurs under humid conditions when the temperature drops below
the dew point. In this study, we have investigated dew conditions on the skin
of the gecko Lucasium steindachneri. When condensation occurs, we show that
small dew drops, as opposed to a thin film, form on the lizards scales. As the
droplets grow in size and merge, they can undergo self-propulsion off
the skin and in the process can be carried away a sufficient distance to freely
engage with external forces. We show that factors such as gravity, wind and
fog provide mechanisms to remove these small droplets off the gecko skin
surface. The formation of small droplets and subsequent removal from the
skin may aid in reducing microbial contact (e.g. bacteria, fungi) and limit
conducive growth conditions under humid environments. As well as providing
an inhospitable microclimate for microorganisms, the formation and removal of
small droplets may also potentially aid in other areas such as reduction and
cleaning of some surface contaminants consisting of single or multiple
aggregates of particles.

1. Introduction
Geckos have received considerable interest, generally focusing on the adhesion
properties of the small structures (setae) on their feet (typically Gekko gecko;
[17]). The remaining regions of the lizard body have received very little attention
in relation to specific functions which may be associated with the nano- and
microstructuring [8,9]. This is somewhat surprising as geckos have interesting
and intricate microstructuring on the skin regions. Foot microstructuring in
geckos may also be evolutionarily linked to the body skin microstructuring
(e.g. substructure of scales) [10].
One prominent feature of this group of lizards is the small hairs, typically
called spines, spinules or microspinules spaced 0.20.7 mm apart and up to several micrometres in height [11,12]. A previous study has shown that these spines
can be water-repellent and suggested that they may serve as a self-cleaning surface where rain may carry away contaminants [8]. Maintaining the surface of the
gecko skin free from a water film may be beneficial for a number of reasons. For
example, the growth of many microorganisms is enhanced by increased water
availability and proliferation may result from such wetting conditions. Indeed,
studies have shown that some lizards are susceptible to various external contaminants that can cause serious skin problems and diseases [13]. High humidity
conditions and low temperatures have been shown to act as potential factors in
the development of such reptile bacterial infections [14]. Thus, the wetting behaviour of the skin (the mechanical barrier and potential portal) plays an important
role in particle and microbial resistance.

& 2015 The Author(s) Published by the Royal Society. All rights reserved.

Downloaded from http://rsif.royalsocietypublishing.org/ on March 11, 2015

In this study, we describe the intricate topographical

surface structure of the body of the gecko (L. steindachneri)
and investigate the wetting formation under humid conditions and potential pathways by which condensates can be
transported from the bodys surface.

2. Experimental procedure
2.1. Gecko capture and preparation
Box-patterned geckos (L. steindachneri) were captured at night
by hand, from the Mingela Ranges (S 20 080 0600 E 146 520 3200 ),
Queensland (QLD), Australia. The Mingela Ranges are semiarid, with a long-term (50 year) median rainfall of 493 mm,
and 62.5 rain days per year on average [19]. Only healthy,
adult lizards were returned to the laboratory, held in plastic containers with a heat source for thermoregulation, paper towel as
substrate (changed weekly), a small tree branch and water ad
libitum. They were fed domestic European crickets (Acheta
domestica) three times a week. Geckos were allowed to shed
twice, so that any oils were completely absent from their skin
before experiments were conducted. Lizards were euthanized
to ensure the skins surface was intact, and undamaged.
Lizard skins were surgically separated by scalpel, cut into smaller sections (15  15 mm2) and attached to glass slides for
droplet experiments.

2.2. Characterization of surface micro- and

nanostructure
Scanning electron microscope (SEM) imaging was performed
on a small section of lizard skin (approx. 3  5 mm2) mounted

2.3. Condensation experiments

The sectioned skin samples were mounted on a cooled copper
element maintained at a temperature below the dew point
(16208C). The surface temperature of the skin was monitored
with a dual-temperature meter HT-L13 with thermocouples
(type K) and an infrared thermometer MT300. Typical conditions of dew formation were achieved in the laboratory with
humidity of 7585% and ambient temperatures of 24278C.

2.4. Droplet cleaning mechanism observations

Droplet formation and dynamics were captured using a Nikon
(V1) digital camera (video mode at 400 and 1200 frames per
second (fps), 640  240 and 320  120 pixel resolution, respectively, frame rates and acquisition times of 2.50 and 0.833 ms per
frame and video play-back at 30 fps or 15 fps). Videos (4064)
were taken observing condensation (duration ranged from 5
to 15 s), with another 336 videos observing small water droplets
impacting with the gecko skin surface (dispensed and dispersed by spray bottle). The camera housing was attached to
a Canon macro lens (EF-S 60 mm ultrasonic) for wide-field,
low-magnification observations, whereas a custom-made optical microscope equipped with Olympus microscope lenses
(MDPlan) of 4 and 10 were used for higher magnifications
(effectively up to approx. 300). Contamination experiments
were carried out using silica (KOBO MSS-500/20average
diameter approx. 20 mm) and polymethylmethacrylate (Bangs
Laboratories BB03Nmean diameter 83 mm) beads. These
were seeded on the skin by distributing a thin coating (single
and clumped) of particles onto a clean glass slide and then
gently tapping the inverted slide over the skin.

3. Results and discussion

3.1. Topographical characterization of the gecko skin
Adult box patterned geckos L. steindachneri (figure 1a) typically
achieve a length of approximately 55 mm with a mass of
approximately 2.5 g and live in a range of areas including
semi-arid regions and are typically ground-dwelling [20]. The
optical images of the dorsal regions (figure 1a,b) show significant pigmentation which may serve as camouflage for the
lizard. Previous studies have investigated the oberhautchen
(outer thin layer) of numerous lizard skins and determined
that they are composed of keratin [2125]. X-ray photon electron spectroscopy data of the gecko skin in our study also
suggest keratin is a component of the dorsal and abdominal
regions where detailed scans showed binding energies of
sulfur and nitrogen consistent with an organic environment.
The micro- and nanostructure on the dorsal and ventral
scales was examined at higher resolutions using a SEM. On
the dorsum, the scales were up to 200 mm in diameter with a
similar centre-to-centre spacing and height, typically over
50 mm, and on the abdomen they were approximately 250 mm
in diameter with a similar spacing (figure 1b,c). The areas
between the scales comprise regions where the skin is heavily
folded (figure 1d). The structuring on the scales at the microand nanolevel consists of spinules (hairs) and a basal layer.
Scales have spinules with lengths of approximately 500 nm to

J. R. Soc. Interface 12: 20141396

(1) To what extent does this self-propulsion process occur in

nature?
(2) Are surfaces with more intricate and complex hierarchical structures than simple insect cuticles used for such
functionality?
(3) Does the process aid in species survival?
(4) Importantly, what specific mechanisms are used and
what factors are involved to aid this process in total
removal of water droplets from the surface (e.g. other
environmental forces, habit, shape of organism)?

on an aluminium stub with carbon-impregnated double-sided

adhesive, sputter coated with 710 nm of platinum, then
imaged using a JEOL 7001 field emission SEM at 8 kV.

rsif.royalsocietypublishing.org

We have investigated a ground-dwelling gecko (Lucasium

steindachneri) that lives in semi-arid habitats, where contact via
rain is limited. For these geckos, rain may be totally absent
between skin-shedding events; however, these lizards live in an
environment in which humid air and low overnight temperatures
can potentially result in condensed liquid water (dew). Thus, the
behaviour of the condensate on the lizard skin is a possible source
of water contamination and thus the focus of our study.
Recently, it has been shown on some natural and artificial
superhydrophobic surfaces that when small condensing water
droplets merge, as they grow in size, then the combined single
droplet is self-propelled off the surface [1517]. This process
arises from a change in surface energy from the transformation
(coalescence) event [15,17]. Interestingly, this process has also
been shown to aid in removal of small contaminating particles
from surfaces and is thus a potential mechanism for self-cleaning
[17]. To date, the only reported examples of the self-propulsion
process on an animal surface has been on insect wings [17,18].
Obvious questions that arise from these studies include

Downloaded from http://rsif.royalsocietypublishing.org/ on March 11, 2015

(a)

rsif.royalsocietypublishing.org
J. R. Soc. Interface 12: 20141396

(b)

(c)

200 mm
(d )

300 mm
(e)

30 mm

2 mm

Figure 1. Images reveal the gecko and the multi-level micro- and nanostructuring on its body. (a) Photograph of the box-patterned gecko Lucasium steindachneri.
(b) Optical image shows the microstructure of the outer skin on the dorsum of the gecko and (c) abdominal region. These regions consisted primarily of dome-shaped
scales in a relatively close-packed hexagonal patterning. (d) Topographical SEM image of the epidermal fold (scales) and areas between scales on the dorsal region of the
lizard, and (e) microstructure on the dorsum showing a micro/nanostructuring consisting of spinules and a base layer patterning. (Online version in colour.)
over 4 mm (figure 1e). The spinules are spherically capped with a
radius of curvature less than 30 nm. The density of spinules is
high: over 400 per 10  10 mm. The structuring below the spinules comprises a network of intersecting ribs forming a
honeycomb-like basal layer (figure 1e). Lenticular sense organs
are also visible on the scale edges (figure 1d). This microand nanostructuring is of sufficient density and roughness to
indicate that the surface may have anti-wetting properties.

3.2. Wettability and dew formation under natural

condensation conditions
The intricate patterning of the gecko spinule array topography facilitates a hydrophobic interaction with water. Static
water contact angles confirm this and are generally within
the range of 151 1558 (figure 2a) on the dorsal and abdominal regions of the gecko, which is at the higher end of
reported values on lizard skin [8,9] and superhydrophobic
in nature. Hysteresis values were comparable to those of
the lotus leaf. There are a number of theories to express the
superhydrophobic condition, all of which have certain
assumptions and limitations [2630]. Cassie & Baxter [27]
express the superhydrophobic state in terms of a number
of interfaces: a liquid air interface with the ambient

environment surrounding the droplet and a surface under

the droplet involving solid air, solid liquid and liquid air
interfaces. Equation (3.1) expresses the contact angle (uC)
formed with a rough surface:
cos uC f1 cos u1  f2

(3:1)

where f1 is the total area of solid under the drop per unit projected area under the drop, f2 is defined in an analogous way
of viewing the liquidair component and u1 is the contact
angle on a smooth surface of the same material.
Equation (3.1) necessitates the surface to have the required
roughness to allow air in topographically favoured regions
such as troughs and surface depressions. Thus, topographies
which increase the airwater interface and minimize the
solidliquid contact area may lead to higher contact angles.
While the use of the original CassieBaxter equation (equation
(3.1)) is often preferable to other variations of the expression, it
does require knowledge of the extent of penetration of the
liquid into the depressions of the surface [30]. If the contact
angle is known, however, then equation (3.1) can be used to predict as a first approximation the extent of penetration of the liquid
into the troughs of a rough surface. From our measured geometric values and approximating the spinule terminus as a
spherical cap (area 2prh, where h is the central distance from

Downloaded from http://rsif.royalsocietypublishing.org/ on March 11, 2015

(a)

(e)

(b)

(f)
65 mm

1 mm

60 mm

300 mm

(d )
300 mm

Figure 2. Images of various sized static water droplets interacting with the gecko skin and snapshots of droplets self-propelling off the gecko skin. (a) Small droplet
of water (approx. 3 mm in diameter) on the dorsal region of the gecko Lucasium steindachneri. The droplet maintains a near-spherical shape and contact angle
(above 1508). (b d ) Condensation formed early in the morning under natural environmental conditions on the dorsal region of the gecko skin showing droplet
distribution, density and size range. Parts (b,c) were taken at a slight angle from the plane of the surface. Part (d ) was obtained along the surface plane. Natural
condensation was observed during a period where temperatures were 21 348C and 8 238C (average day-time and night-time temperatures, respectively: humidity
up to 94%). (e) Condensation drops autonomously propelling off the skin (geckovescence) with the surface oriented horizontally (see also the accompanying
electronic supplementary material, movie S1, which shows that the propelled droplets clearly leave the surface and move under the influence of gravity).
(f ) Self-propulsion of water droplets from the abdominal region of the gecko skin. Droplets in the range of 10 80 mm propel off the surface (see also electronic
supplementary material, movie S2). (Online version in colour.)
the top of the sphere to the point at which liquid will invade) and
u1 as 1158 and using a droplet invasion of 12.5 nm will calculate a
predicted contact angle of approximately 1758. The experimentally determined value of 1511558 suggests that an area of
approximately 40 times this value is required to bring the contact
angle within the measured range. Thus, the droplet may invade
to a greater extent. Inclined or slightly bent spinules near the apex
may significantly increase the contact area with resting bodies
on the surface. In addition, owing to the dome-shaped scale topography and folds on the skin surface, water droplets may interact
with the spinule shafts to varied degrees. Indeed, the spinules
may undergo bending (especially near the tips), which may
also increase contact area. If we make a very basic approximation
of the spinules as cylindrical spring structures and consider only
the narrower end regions of the tapered shafts as the structural
elements, then we can view a tip displacement by modification
of a cantilever bending equation [31]
Dsp

4L3 Fdrop
3pNsp Rs 4 E

(3:2)

where L is the length (400 nm used in our case where a significant

fraction of the bending will presumably occur), E is the elastic

modulus of the material (1.5 GPa), Fdrop is the total droplet

force (a value of the radius, Rs, of 30 nm is used, reflecting
the lower region of the cylindrical element (spinule)). For a
water droplet of 2 mm diameter (approx. 4 ml) resting on the
skin and assuming an equally distributed force contact on the
end of around 100 000 spinules (based on spinule density and
droplet contact area determined by microscopic imaging directly
above the droplet), a deflection of over 5 nm would take place for
each individual spinule near the tip.
It should be noted that the above description makes a
number of assumptions and ignores numerous other parameters/factors that may cause the deviation of the contact
angle from predictions, such as the non-homogeneity of the
surface, defects, chemical variations and the natural bending
itself as it appears in figure 1e. Nevertheless, the abovementioned illustrative example provides some informative
predictions for water interactions with surface projections of
the scale as exhibited by the gecko and presents a useful comparative behaviour with smaller droplet interactions and
other organisms.
The spinule topography on the gecko is different from the
structure found on many other lizards [3235]. Some other
organisms, however, do have a dense spinule (hair) array

J. R. Soc. Interface 12: 20141396

(c)

rsif.royalsocietypublishing.org

600 mm

Downloaded from http://rsif.royalsocietypublishing.org/ on March 11, 2015

To assess whether naturally occurring condensed droplets, such

as those shown in figure 2, when merging will form larger droplets and remain pinned to the skins surface, or alternatively be
self-propelled off the surface from changes in energy during
coalescence, we conducted condensation experiments. By cooling the gecko skin below the dew point, surface condensation
was induced forming droplets in the same size range as those
shown in figure 2bd. As they grew, some merged and then
self-propelled from the skin surface. Figure 2e shows a snapshot
in time of micro-droplets propelling autonomously off the skin
(see the accompanying electronic supplementary material,
movie S1). The fully merged propelled droplets were typically
approximately 40120 mm in diameter.
A recent paper [16] has shown that on a fabricated surface
as the subcooling value increases (termed supersaturation by
the authors) the micro/nanostructures fail to maintain the

3.3.1. Direct self-propulsion of droplets off the skin surface via

the droplet merging process (single jumping event)
For illustrative purposes, the droplet propulsion process can be
viewed in a very simplified manner by considering changes in
the droplet surface energies. If we consider the case where two
small water droplets (not necessarily of equal size) coalesce on a
superhydrophobic surface to form a larger droplet, the maximum height, Hmax, that can be attained by a droplet can be
determined by integrating the velocity of a droplet over its
time-of-flight resulting in equation (3.3):
"
(
)#
rw Rm
9ra CD fw
(1 f)2
Hmax
ln 1 2 2
1
(3:3)
3ra CD
Rm rw g (1 f 3 )2=3
where ra,w are the densities of air and water, respectively; Rm is
the radius of the merged single droplet; CD is the drag coefficient of the droplet in air; fw is the water surface tension; g is
the gravitational acceleration (9.8 m s22); and f is the ratio of
drop diameters.
Equation (3.3) sets an upper limit for the maximum possible
height that can be reached assuming all the released surface
energy is converted into kinetic energy of the droplet and ignoring viscous forces and adhesional forces of the droplet to the
surface. It also assumes that the droplet is propelled vertically
and that the drag coefficient remains constant as the droplet
decelerates. Interestingly, the maximum height calculated
from equation (3.3) varies little over the range of smaller droplet
diameters. The height achieved, however, is significantly

J. R. Soc. Interface 12: 20141396

3.3. Self-propulsion of condensing droplets from

coalescence

jumping-induced removal mechanism owing to an increase

in the number of nucleation sites. This leads to flooding and
a loss of superhydrophobicity and the formation of highly
pinned Wenzel droplet morphologies. In our case, humidity
and temperature changes, while potentially large, generally
occur over a relatively long period of time in the natural
environment of the gecko, and thus flooding seems unlikely
to occur. Observation of droplet numbers as shown in
figure 2 supports this premise; however, how the gecko
structuring performs in relation to fabricated surfaces in this
area is of interest and worthy of exploration in future studies.
To investigate the range of droplet sizes involved, videos at
higher magnification were collected of self-propulsion on both
the dorsal and abdominal skin of the gecko. A sequence of
images from the abdominal region of the gecko skin where droplets of varying size (from 10 mm to over 80 mm in diameter)
merge and are self-propelled from the surface is seen in
figure 2f (see also electronic supplementary material, movie S2).
This mechanism demonstrates that small water droplets can
be projected from the skin surface with no external forces. If the
lizard, however, is orientated in a horizontal position with
respect to gravity as demonstrated in figure 2e, then the selfpropelled droplet may eventually return to the skin under the
influence of gravity. It may remain there if no other forces are
acting on it and the droplet propulsion was orthogonal
(normal) to the overall plane of the lizards skin. However, the
ultimate destination of propelled droplets will depend on
numerous factors (e.g. lizards in their natural environment will
adopt several orientations and will also be exposed to other
environmental influences and forces). With this in mind, we
explored a number of skin surface orientations and observed
the overall dynamics of the droplet-propelled process in different
situations. We have identified a number of mechanisms that may
aid in complete transfer of droplets from the surface of the skin.

rsif.royalsocietypublishing.org

on their surface. Many insects, for example, have hairs on their

cuticle and especially wing surfaces [36] such as those found on
lacewings, craneflies and termites [18,3740]. In these cases,
however, the hairs on the wings are significantly longer and
spaced over an order of magnitude further apart (over 10 mm
in length and spacing from 5 to 15 mm apart). This is also typical
of many other insect species where hairs are separated by tens
of micrometres. The much larger spacing and lower density of
hairs on insects provides a shedding mechanism to remove
water quickly so as to keep wings free from liquid, thus maintaining functionality. The nanostructures (e.g. nano-grooves,
nano-channels) on the hairs of many insects have been shown
to aid in anti-wetting [37,38]. The gecko, while not having
such channel structuring on the hairs, does however exhibit a
much higher density of hairs. Thus, it appears that the high
hair density on the gecko skin is sufficient to minimize the interaction time with water in various forms without the aid of finer
structuring. Small-scale structuring found on insect surfaces at
the submicrometre dimensions, of a similar lateral scale to the
spinule spacing as seen on the gecko, has been associated
with cleaning/self-cleaning of the cuticle surfaces [39,41]. Comparatively, large water droplets such as the example in figure 2a
can become mobile with minor tilting of the surface (a few
degrees) and thus are easily displaced from their equilibrium
state. This not only facilitates water removal from the surface,
but also facilitates the extrication of collected contaminants
from the skin.
In more arid regions, a significant period may pass without rain and thus the most likely form of water contamination
will arise from atmospheric condensation on the lizard skin.
When we exposed gecko skin samples to natural environmental condensation conditions, dew formed on the surface
in the form of small droplets as opposed to a continuous
film (figure 2bd ). The dew droplet population that formed
on the gecko skin consisted of many small densely packed
spherically shaped, micrometre-sized droplets. The surfacebound droplets on the skin were typically in the range of
10 100 mm with each scale region often housing multiple
droplets. From examination of figure 2b d, it is clear that
the density of dew droplets (related to the spacing of nucleation sites) and the growth of droplets from the condensation
process will facilitate merging of two or multiple clusters of
small droplets.

Downloaded from http://rsif.royalsocietypublishing.org/ on March 11, 2015

(a) 0 ms

3.34 ms

6.67 ms

0 ms

(b)

4.17 ms

0.83 ms

4.17 ms

5 ms

1.67 ms

5.83 ms

2.5 ms

6.67 ms

3.34 ms

7.5 ms

8.34 ms

1
1.67 ms

5 ms

9.17 ms
2

2.5 ms

5.83 ms

10.83 ms
0.47 mm

Figure 3. (a) Lateral sweeping motion of self-propelled droplets along the surface arising from coalescence of multiple droplets of varying sizes (electronic supplementary material, movie S4). (b) Large mobile self-propelled droplet moving laterally with a high kinetic energy sufficient to maintain direction and scavenge
smaller droplets (electronic supplementary material, movie S5). (Online version in colour.)

reduced when coalescence occurs with droplets of dissimilar

sizes. The droplet size distribution from condensation, such as
those seen in figure 2, ensures droplets of different sizes will
coalesce frequently, although droplets of similar sizes will also
merge (as those seen in figure 2e). Droplets exceeding the dimensions of the lizard scales (larger than 200 mm) are rare (figure 2),
thus droplets smaller than this will most often be involved in the
self-propulsion process. This size range of droplets upon
coalescence will also overcome adhesional forces from the surface, and, once free, these smaller droplets are less influenced
by gravitational forces than larger droplets.
Equation (3.3) predicts a maximum height approaching
20 mm; however, the majority of droplets were propelled
0.52 mm (figure 2e and electronic supplementary material,
movie S1). A more comprehensive view of the propulsion process has been presented by Peng et al. [42]. They view the initial
total kinetic energy of the coalesced droplet as Eitk DEs 2
Evis 2 Eh 2 Ew 2 Ecah, where Eitk is the initial total kinetic
energy of the coalesced droplet, DEs is the surface energy
released by the droplet coalescence, Evis is the viscous dissipation in the droplet, Eh is the gravitational energy change
during droplet coalescence and Ew is the work of adhesion.
Ecah is the energy consumed overcoming the contact angle
hysteresis.
The authors state that approximately 25% of the energy
released by the droplet coalescence is converted to the effective kinetic energy in the vertical motion of the coalesced
droplet jumping from the surface. Thus, the higher propelled
heights predicted in equation (3.3) can be attributed to the
omission of other factors such as viscous dissipation. The
droplet propulsion process was also measured at 1200 fps,
allowing droplet speed to be measured accurately. The
typical initial velocities of droplets were below 0.4 m s21.
The merging of two or more droplets can cause selfpropulsion off a surface (electronic supplementary material,
movies S1 S2). If the gecko is orientated in a vertical position, then the single propulsion of a droplet can result in
complete removal from the surface. It is evident that if the
gecko is inclined at near-vertical angles (approaching 908),
then propelled droplets will be able to traverse large distances and are more likely to be totally removed from the

surface (see electronic supplementary material, figure S1

and movie S3surface inclined at approx. 708). If the lizard
is orientated in a horizontal or near-horizontal stance, then
the underlying body and side regions of the gecko will also
be amenable to this mechanism. The small size of the gecko
and the rounded cross-sectional profile of the lizard body
will also facilitate complete removal via a single propulsion
of a water droplet. Once droplets are self-propelled from
the skin in these situations gravity can assist in complete
removal of the droplets. Thus, no other external force is
required for droplet removal. As the distance that a droplet
can be propelled is significant in relation to the smallest lateral dimensions of this lizard then this mechanism can, in
principle, remove droplets from anywhere on the surface of
the skin.

3.3.2. Self-propelled impacting droplets which laterally sweep

the surface and facilitate removal of surface-bound
condensationremoval facilitated by sufficiently high
kinetic energy of impacting droplets
Merging droplets are typically propelled in near-vertical directions in relation to the skin surface (for example, electronic
supplementary material, movie S1). Some proportion of droplets, however, is propelled at angles facilitating motion
along the plane of the skins surface rather than directly out
of plane. This lateral sweeping of droplets across the surface
is often facilitated by the coalescence of multiple droplets of
varying sizes (figure 3a and electronic supplementary material,
movie S4). This may potentially be aided by the skin topography where droplets that merge are initially resting on scales
and thus are on an inclined surface (figure 2f ). Sufficiently
large droplets merging can be driven along the surface with
sufficient momentum to remove themselves from the surface
and in the process collect smaller droplets along the way as
shown in figure 3a,b (see also electronic supplementary
material, movie S5). The process may be viewed as an extended
form of the lotus effect (impacting and rolling droplets), where
the initiating step is via self-propulsion(s) of condensation as
opposed to a falling or rolling droplet from rain. Thus, the
removal process may take the form of transformation of the

J. R. Soc. Interface 12: 20141396

0.83 ms

rsif.royalsocietypublishing.org

100 mm

Downloaded from http://rsif.royalsocietypublishing.org/ on March 11, 2015

(b)

(a)

0 ms

5 ms

8.34 ms

3.34 ms

5.83 ms

9.17 ms

1.67 ms

200 mm
4.17 ms

22.5 ms

12.5 ms

23.34 ms

15.83 ms

24.17 ms

4.17 ms

6.67 ms

10 ms

Figure 4 Small water droplets merging with surface drops and self-propelling off the surface of the gecko skin. (a) A small impacting droplet of 20 mm with a
velocity of 0.013 m s21 impacts with a stationary condensed droplet on the surface and the resulting change in surface energy propels the coalesced droplet above
the surface at a speed of over seven times greater than the impacting droplet (electronic supplementary material, movie S6). (b) A small impacting droplet (highlighted by the arrow) contacting two stationary condensed droplets on the surface which are in very close proximity. The two surface-bound droplets merge from the
process and are self-propelled off the skin (electronic supplementary material, movie S7).
self-propulsion process to that resembling a rolling process
along the surface as in the lotus effect.

3.3.3. Impacting droplets facilitate self-propulsion off the skin

surface (impacting droplets from fog and small selfpropelled droplets)
The situation where small falling water droplets impact stationary surface-bound droplets is shown in figure 4a,b. A falling
droplet of 20 mm diameter with a velocity of 0.013 m s21 impacting a stationary surface-bound droplet of approximately 70 mm
is illustrated in figure 4a and also in the electronic supplementary material, movie S6. Figure 4b shows a very small droplet
(highlighted by the arrow) travelling at a slow velocity,
making contact with two stationary surface droplets of
approximately 100 and 70 mm in close proximity to each other,
resulting in self-propulsion off the surface (see also electronic
supplementary material, movie S7).

Fog conditions occur regularly in areas inhabited by this

particular gecko (observation from collection and weather
data for regions of habitat (Bureau of Meteorology [19])) and
often coincide when high humidity conditions take place. Fog
as well as previously self-propelled droplets may collide with
the skin and merge with existing dew droplets, resulting in a
newly merged droplet self-propelling off the surface. The process may be aided by kinetic energy of the impacting mobile
droplet; however, for slow moving small droplets the surface
energy changes are the primary driving force for propulsion
of the final droplet. The change in surface energy from coalescence in figure 4a results in a self-propelled droplet approaching
0.1 m s21. Small, mobile impacting droplets may also facilitate
self-propulsion by adding to the volume of stationary droplets
forcing them to merge with neighbouring droplets (figure 4b
and electronic supplementary material, movie S7). Propulsion
arising from small impacting droplets may also be responsible
for future coalescence and subsequent propulsion events.

J. R. Soc. Interface 12: 20141396

2.5 ms

0.83 ms

21.67 ms

rsif.royalsocietypublishing.org

70 mm

0 ms

Downloaded from http://rsif.royalsocietypublishing.org/ on March 11, 2015

rsif.royalsocietypublishing.org

III

IV

II
I

gecko back

Figure 5. Schematic summary of the mechanisms that can potentially totally remove small water droplets off the gecko skin. (I) Direct self-propulsion of droplets off
the skin surface (near vertical projection). (II) Multiple coalescence of droplets driven along the plane of the surface (sweeping) with the droplet scavenging laterally
along the surfacecan result in lotus-type collection. (III) Impacting droplets from fog or other falling droplets facilitating self-propulsion off the skin surface.
(IV) Wind-assisted removal. (Online version in colour.)
(a)

0 ms

3.33 ms (b)

70 mm

0.83 ms

4.17 ms

1.67 ms

23.33 ms

2.5 ms

24.17 ms

100 mm

Figure 6. (a) A condensed water droplet of approximately 100 mm in diameter encapsulating a silica particle (highlighted by the arrow) of 15 mm in diameter. The droplet
and its contaminant are self-propelled off the surface. (b) Before and after (self-propulsion) frames show droplets which house silica particles. (Online version in colour.)

3.3.4. Wind-assisted removal of self-propelled droplets

A significant population of propelled droplets have sufficient
momentum to be transported outside the skinair boundary
layer (figure 2e), allowing them to be exposed to external
environmental forces (such as light breezes). Importantly,
the self-propulsion releases the droplet from the surfaceair
interface and associated adhesive forces and exposes it to
forces that could potentially transport the drop significant distances away. As the droplets are extremely small, their mass
will be sufficiently low that very slight external wind forces
are able to fully remove the droplet from the skin once they
have self-propelled above the surface. For a 50 mm diameter
water condensate, a wind force of less than 1 nN is required
to oppose the gravitational force. For a 200 mm droplet, the
force increases to approximately 40 nN. Electronic supplementary material, movie S8, illustrates the susceptibility of
droplets to removal if exposed to light breezes over the surface.
Droplets can easily be transported several centimetres away
from the point of origin. Wind forces may also initiate selfpropulsion of droplets by transporting small droplets laterally
on the surface forcing coalescence.

A summary of the mechanisms discussed above is shown

schematically in figure 5, all of which can totally remove
water droplets from the skin.

3.3.5. Other potential benefits of self-propulsion of condensed

droplets
While the formation of high-contact-angle small micrometresized droplets and subsequent self-propulsion provides a
landscape under dew conditions which can potentially limit
the growth of microorganisms such as bacteria and fungi
on the lizard skin, does the phenomenon offer any other
possible attributes for the organism? It should be noted that
the self-propulsion of water droplets on artificial superhydrophobic surfaces has shown to be a way of controlling heat
transfer [16] and this may be relevant as some water-repellent
reptile surfaces may contribute to thermoregulation.
The self-propelled droplet-assisted mechanisms as seen in
figure 5 may also possibly aid in removal of some contaminants
from the gecko skin. We have made a preliminary investigation to
see if contaminants (e.g. silicon and polymethylmethacrylate

J. R. Soc. Interface 12: 20141396

Fg

Downloaded from http://rsif.royalsocietypublishing.org/ on March 11, 2015

(a)

(c)

rsif.royalsocietypublishing.org

150 mm

1 mm

150 mm

Figure 7. (a) Before and after snapshots of a small clump of PMMA particles being removed off the surface. (b) A large clump of PMMA particles self-propelling
along the surface until coming to rest nested up against another large clump (electronic supplementary material, movie S10). (c) A relatively large area being
cleaned on the surface from numerous droplets/clumps of particles. (Online version in colour.)
particles) could be removed by condensation and self-propulsion
processes. Figure 6a shows that a small silica particle encapsulated in a droplet can be removed via the droplet propulsion
mechanism. Figure 6b shows groups of droplets (two and three
droplets which house silica beads) demonstrating self-prolusion.
The gecko skin surface was also seeded with polymethylmethacrylate beads (PMMAmean diameter of 83 mm with a
contact angle of 608) and exposed to condensation conditions.
Large particles and particle clumps were transported along
the surface with only small amounts of condensed water surrounding them (electronic supplementary material, movie S9).
Particle clumps rearrange owing to meniscus forces, and the
energy was released in the form of jumping conglomerates of
particles (figure 7a,b and electronic supplementary material,
movie S10).
While a previous study has shown a single jump of hydrophilic glass beads from a superhydrophobic wing surface [17],
we have found that small clumps of these less hydrophilic
PMMA particles (in heavily contaminated regions) tended
to undergo sequential aggregation with a series of selfpropulsions and can be finally propelled along or off the surface.
The aggregate seen in figure 7b makes intermittent contact with
the surface during the propulsion event by bouncing across the
surface. Interestingly, PMMA clumps comprising only a few
particles can be propelled from the skin surface (figure 7a). As
well as nucleation occurring on the skin surface, figure 7a
demonstrates that nucleation of droplets can occur on individual particles. It is apparent from figure 7a that only small
condensed water drops or films relative to the particle size can

facilitate self-propulsion off the surface. The lateral sweeping

off of particles/droplets along the plane of the skin surface is
shown in figure 7c (see also electronic supplementary material,
movie S11). In this case, multiple droplets/clumps containing
numerous particles can be transported along the surface leaving
the skin free from contamination.
A few recent studies have investigated self-cleaning of the
gecko foot [43,44] and suggested that adhesive forces attracting
a dirt particle to the substrate and hyperextension are two significant factors for cleaning. Of note in our study are the similarities
as well as the significant differences in foot structuring and spinule structure (radius of curvature, multiple projections, etc.;
see visual comparison in electronic supplementary material,
figure S2).

4. Conclusion
The dense array of spinules on the gecko skin provides an architecture suited for removal of large water droplets. It also
provides an ideal construction for the self-propulsion of small
condensed droplets. Dew on some surfaces may promote the
growth of pathogens such as fungi and bacteria and may
enable spores to develop. In humid environments, this temporary water formation or thin film may result in proliferation
resulting in larger populations of microorganisms. As the dew
on the superhydrophobic gecko skin is a temporary phenomenon (constant self-propulsion of droplets and small-sized
droplets leading to quick evaporation in sunlight/heated

J. R. Soc. Interface 12: 20141396

(b)

Downloaded from http://rsif.royalsocietypublishing.org/ on March 11, 2015

removal of just a few particles with minimal water/droplet

coverage and expansive cleaning from lateral mobile droplets
brushing the surface clean. While we have identified removal
mechanisms, the efficiency of cleaning the surface based on
droplet and condensate processes for removing particles
(e.g. minor contamination of biological matter, bacteria, soil
particles) requires further investigation in subsequent studies.

Ethics statement. This work was conducted under ethics approval

A1676, and QNP permit WITK05209908.
scientific and technical assistance, of the Australian Microscopy and
Microanalysis Research Facility at the Centre for Microscopy and
Microanalysis, The University of Queensland, where the scanning
electron microscope images in figure 1d,e were collected.

References
1.

Russell AP. 2002 Integrative functional morphology

of the Gekkotan adhesive system (Reptilla: Gekkota).
Integr. Comp. Biol. 42, 1154 1163. (doi:10.1093/
icb/42.6.1154)
2. Stark AY, Badge I, Wucinich NA, Sullivan TW,
Niewiarowski PH, Dhinojwala A. 2013 Surface
wettability plays a significant role in gecko adhesion
underwater. Proc. Natl Acad. Sci. USA 110,
63406345. (doi:10.1073/pnas.1219317110)
3. Huber G, Mantz H, Spolenak R, Mecke K, Jacobs K,
Gorb SN, Arzt E. 2005 Evidence for capillarity
contributions to gecko adhesion from single spatula
nanomechanical measurements. Proc. Natl Acad. Sci.
USA 102, 16 293 16 296. (doi:10.1073/pnas.05063
28102)
4. Huber G, Gorb SN, Hosoda N, Spolenak R, Arzt E.
2007 Influence of surface roughness on gecko
adhesion. Acta Biomater. 3, 607 610. (doi:10.1016/
j.actbio.2007.01.007)
5. Autumn K et al. 2002 Evidence for van der Waals
adhesion in gecko setae. Proc. Natl Acad. Sci. USA
99, 12 25212 256. (doi:10.1073/pnas.192252799)
6. Sun W, Neuzil P, Kustandi TS, Oh S, Samper VD.
2005 The nature of the gecko lizard adhesive force.
Biophys. J. 89, L14L17. (doi:10.1529/biophysj.105.
065268)
7. Bhushan B. 2009 Biomimetics: lessons from
naturean overview. Phil. Trans. R. Soc. A 367,
14451486. (doi:10.1098/rsta.2009.0011)
8. Hiller UN. 2009 Functional surfaces in biology.
Berlin, Germany: Springer Science.
9. Spinner M, Gorb SN, Westhoff G. 2013 Diversity of
functional microornamentation in slithering geckos
Lialis (Pygopodidae). Proc. R. Soc. B 280, 20132160.
(doi:10.1098.rspb.2013.2160)
10. Russell AP. 1976 Morphology and biology of reptiles.
Linnean Society Symposium Series 3. London, UK:
Academic Press.
11. Ruibal R. 1968 The ultrastructure of the surface of
lizard scales. Copeia 4, 698703. (doi:10.2307/
1441836)
12. Rosenberg HI, Russell AP, Cavey MJ. 1992
Development of the subdigital adhesive pads of

13.

14.

15.

16.

17.

18.

19.
20.

21.

22.

Ptyodactylus guttatus (Reptilia: Gekkonidae).

J. Morphol. 211, 243258. (doi:10.1002/jmor.
1052110302)
Pare JA, Sigler L, Rosenthal KL, Mader DR. 2006
Reptile medicine and surgery. London, UK: Saunders
Elsevier.
Hoppmann E, Barron HW. 2007 Dermatology in
reptiles, topics in medicine and surgery. J. Exot. Pet
Med. 16, 210224. (doi:10.1053/j.jepm.2007.
10.001)
Boreyko JB, Chen CH. 2009 Self-propelled dropwise
condensate on superhydrophobic surfaces. Phys.
Rev. Lett. 103, 184501. (doi:10.1103/PhysRevLett.
103.184501)
Miljkovic N, Enright R, Nam Y, Lopez K, Dou N, Sack
J, Wang EN. 2013 Jumping-droplet-enhanced
condensation on scalable superhydrophobic
nanostructured surfaces. Nano Lett. 13, 179187.
(doi:10.1021/nl303835d)
Wisdom KM, Watson JA, Qu X, Liu F, Watson GS,
Chen CH. 2013 Self-cleaning of superhydrophobic
surfaces by self-propelled jumping condensate. Proc.
Natl Acad. Sci. USA 110, 7992 7997. (doi:10.1073/
pnas.1210770110)
Watson JA, Cribb BW, Hu HM, Watson GS. 2011
A dual layer hair array of the brown lacewing:
repelling water at different length scales.
Biophys. J. 100, 1149 1155. (doi:10.1016/j.bpj.
2010.12.3736)
Bureau of Meteorology. 2014 See http://www.bom.
gov.au/.
Wilson S. 2005 A field guide to the reptiles of
Queensland. Sydney, Australia: New Holland
Publishers.
Molina R, Jovancic P, Jocic D, Bertran E, Erra P. 2003
Surface characterization of keratin fibres treated by
water vapour plasma. Surf. Interface Anal. 35,
128 135. (doi:10.1002/sia.1510)
Toni M, Dalla Valle L, Alibardi L. 2007 The
epidermis of scales in gecko lizards contains
multiple forms of b-keratins including basic
glycine proline serine-rich proteins. J. Proteome
Res. 6, 1792 1805. (doi:10.1021/pr060626+)

23. Fraser RDB, Parry DAD. 1996 The molecular

structure of reptilian keratin. Int. J. Biol. Macromol.
19, 207 211. (doi:10.1016/0141-8130(96)01129-4)
24. Dalla Valle L, Nardi GB, Bonazza G, Zuccal C, Emera
D, Alibardi L. 2010 Forty keratin-associated bproteins (b-proteins) form the hard layers of scales,
claws, adhesive pads in the green anole lizard,
Anolis carolinensis. J. Exp. Zool. B 314, 11 32.
(doi:10.1002/jez.b.21306)
25. Dalla Valle L, Nardi A, Toffolo V, Niero C, Toni M,
Alibardi L. 2007 Cloning and characterization of
scale b-keratins in the differentiating epidermis of
geckoes show they are glycine prolineserine-rich
proteins with a central motif homologous to avian
keratins. Dev. Dyn. 236, 374388. (doi:10.1002/
dvdy.21022)
26. Wenzel RN. 1936 Resistance of solid surfaces to
wetting by water. Ind. Eng. Chem. 28, 988 994.
(doi:10.1021/ie50320a024)
27. Cassie ABD, Baxter S. 1944 Wettability of porous
surfaces. Trans. Faraday Soc. 49, 546 551. (doi:10.
1039/TF9444000546)
28. Herminghaus S. 2000 Roughness-induced nonwetting. Europhys. Lett. 52, 165170. (doi:10.
1209/epl/i2000-00418-8)
29. Gao L, McCarthy TJ. 2007 How Wenzel and Cassie
were wrong. Langmuir. 23, 3762 3765. (doi:10.
1021/la062634a)
30. Milne AJB, Amirfazli A. 2012 The Cassie equation: how
it is meant to be used. Adv. Coll. Int. Sci. 170, 4855.
(doi:10.1016/j.cis.2011.12.001)
31. Gere JM, Timoshenko SP. 1984 Mechanics
of materials. Independence, KY: Thomson
Brooks/Cole.
32. Peterson JA, Bezy RL. 1985 The microstructure and
evolution of scale surfaces in Xantusiid lizards.
Herpetologica 41, 298324. (http://www.jstor.org/
stable/3892277)
33. Peterson JA. 1984 The microstructure of the scale
surface in iguanid lizards. J. Herp. 18, 437467.
(doi:10.2307/1564106)
34. Harvey MB. 1993 Microstructure, ontogeny, and
evolution of scale surfaces in xenosaurid lizards.

J. R. Soc. Interface 12: 20141396

Acknowledgements. The authors acknowledge the facilities, and the

10

rsif.royalsocietypublishing.org

conditions), this may limit or minimize pathogen attachment

and growth. It is also worth mentioning that self-propelled droplets will lead to a surface inhibiting a water film formation on
the skin. We have identified four major processes that can be
employed in assisting removal of small condensed droplets.
External environmental forces such as gravity, wind (by ambient air flow) and impacting small droplets (e.g. fog) and
laterally projected droplets can assist the self-propulsion mechanism for full removal from the surface. The sweeping motion of
droplets from multiple drops merging is an interesting process
as it does not need to fully rely on gravity as the motion is lateral
along the surface.
We have also shown that the skin surface is conducive to
cleaning by these mechanisms, which have the potential to
carry away small contaminants. Of particular interest is the

Downloaded from http://rsif.royalsocietypublishing.org/ on March 11, 2015

ordered cuticular nano-arrays on insect wings.

Biophys. J. 94, 3352 3360. (doi:10.1529/biophysj.
107.109348)
42. Peng B, Wang S, Lan Z, Xu W, Wen R, Ma X. 2013
Analysis of droplet jumping phenomenon with
lattice Boltzmann simulation of droplet coalescence.
Appl. Phys. Lett. 102, 151601. (doi:10.1063/1.479
9650)
43. Hansen WR, Autumn K. 2005 Evidence for selfcleaning in gecko setae. Proc. Natl Acad. Sci. USA
102, 385389. (doi:10.1073/pnas.0408304102)
44. Hu S, Lopez S, Niewiarowski PH, Xia Z. 2012
Dynamic self-cleaning in gecko setae via digital
hyperextension. J. R. Soc. Interface 9, 2781 2790.
(doi:10.1098/rsif.2012.0108)

11

J. R. Soc. Interface 12: 20141396

38. Watson GS, Cribb BW, Watson JA. 2010 How micro/
nanoarchitecture facilitates anti-wetting: an elegant
hierarchical design on the termite wing. ACS Nano
4, 129 136. (doi:10.1021/nn900869b)
39. Hu HM, Watson JA, Cribb BW, Watson GS. 2011
Fouling of nanostructured insect cuticle: adhesion of
natural and artificial contaminants. Biofouling 27,
1125 1137. (doi:10.1080/08927014.2011.637187)
40. Watson GS, Cribb BW, Hu HM, Watson JA. 2011
Contrasting micro/nano architecture on termite
wings: two divergent strategies for optimising
success of colonisation flights. PLoS ONE 6, 110.
(doi:10.1371/journal.pone.0024368)
41. Watson GS, Myhra S, Cribb BW, Watson JA. 2008
Putative function(s) and functional efficiency of

rsif.royalsocietypublishing.org

J. Morphol. 216, 161 177. (doi:10.1002/jmor.

1052160205)
35. Harvey MB, Gutberlet RL. 1995 Microstructure,
evolution, and ontogeny of scale surfaces in cordylid
and gerrhosaurid lizards. J. Morphol. 226, 121
139. (doi:10.1002/jmor.1052260202)
36. Wagner T, Neinhuis C, Barthlott W. 1996
Wettability and contaminability of insect wings
as a function of their surface sculptures. Acta Zool.
77, 213225. (doi:10.1111/j.1463-6395.1996.
tb01265.x)
37. Hu HM, Watson GS, Cribb BW, Watson JA. 2011 Non
wetting wings and legs of the cranefly aided by
fine structures of the cuticle. J. Exp. Biol. 214,
915920. (doi:10.1242/jeb.051128)