Apex structures enhance water drainage on leavesTing Wanga,b, Yifan Sia, Haoyu Daia, Chuxin Lia, Can Gaoa,b, Zhichao Donga,1

, and Lei Jianga,1

aChinese Academy of Sciences Key Laboratory of Bio-inspired Materials and Interfacial Sciences, Technical Institute of Physics and Chemistry, ChineseAcademy of Sciences, 100190 Beijing, China; and bSchool of Future Technology, University of Chinese Academy of Sciences, 101407 Beijing, China

Edited by Peidong Yang, University of California, Berkeley, CA, and approved December 20, 2019 (received for review June 10, 2019)

The rapid removal of rain droplets at the leaf apex is critical forleaves to avoid damage under rainfall conditions, but the generalwater drainage principle remains unclear. We demonstrate thatthe apex structure enhances water drainage on the leaf byemploying a curvature-controlled mechanism that is based onshaping a balance between reduced capillarity and enhanced grav-ity components. The leaf apex shape changes from round to tri-angle to acuminate, and the leaf surface changes from flat to bent,resulting in the increase of the water drainage rate, high-drippingfrequencies, and the reduction of retention volumes. For wet trop-ical plants, such as Alocasia macrorrhiza, Gaussian curvature recon-figuration at the drip tip leads to the capillarity transition fromresistance to actuation, further enhancing water drainage to thelargest degree possible. The phenomenon is distinct from thewidely researched liquid motion control mechanisms, and it offersa specific parametric approach that can be applied to achieve thedesired fluidic behavior in a well-controlled way.

drip tip | biomimetic | capillarity | curvature transition | water shedding

Water drops interacting with substrate are omnipresent in ourlives, and shedding is relevant to numerous fields, including

microfluidics (1), printing (2), spraying (3), transfer (4, 5), andwater collection (6), as well as for the survival of natural species(7–11). Many natural species utilize their surface structures, suchas bumps, spines, or irregular structures with varying curvatureoutlines, to directional transport water and achieve water con-servation and drainage. For instance, the periodic spindle-knotsand joints of spider silk and hierarchical curvatures of pitcherplants transport water to the desired destination (7, 8), and theoriented microstructures of butterfly’s wing and clusters of wetfibers on dog shed undesired water for stable movement and tokeep warm (9, 10). Although these delicate structures performwell, they only influence small volumes of water, in dropletsizes, over limited transport distances. In heavy rain events, itremains a challenge to achieve fast water drainage in bulkvolumes, especially for wet tropical plants that typically havebroad leaves.The drip tip is the most famous evolution of rainforest leaf

with curving apex shapes and orientations for quick drainage (SIAppendix, Table S1). Although biologists found the fast drainageability of the drip tip since the 1980s (12–18), the detailedmechanism of how the apex improves the water drainage re-mains unknown. Besides, it is still not clear whether the curvingstructures of leaf apex affect the drainage process. Hence, insightinto the behavior of droplets interacting with the curvature-varying surface at the apex is needed.Here, we demonstrate the water drainage behavior of leaf is

controlled by the leaf apex structure and the leaf surface cur-vature. The surface curvature variation of the leaf apex reducesthe capillary resistance force, and the bending down of the leafsurface enhances the water gravity component, leading to a highdripping frequency and low water retention. Gaussian curvaturereconfiguration at the leaf drip tip leads to the capillarity changefrom resistance to actuation, which forces droplet detaching atthe transition point and enhancing drainage to the highest de-gree possible. Measurements of leaf apices and water drainageacross the plant leaves from the temperate zone to the wet

tropics support this finding. The leaf of rainforest plant Alocasiamacrorrhiza, which integrates multiple structures and curvatureadvantages at the leaf apex, has a high drainage efficiency.Learning from A. macrorrhiza, we optimize an artificial drain-age device to acquire superior drainage ability than most naturalspecies.

Results and DiscussionWater Drainage Efficiency of Different Leaf Apices. Plant leaves inshape and size vary with temperature and precipitation (19). Theleaf apex is a protruding part of a leaf where water dropletsaccumulate, and droplet separation occurs during drainage. Todetermine the abilities of different leaf apex morphologies onwater drainage control, 44 different species (36 families) ofleaves with various apex angles (20), α, were investigated andclassified (Fig. 1A and SI Appendix, Tables S2 and S3). Theseleaves were collected from the botanical garden in the sub-tropical zone with less precipitation in Beijing, China, andtropical zone with rich precipitation in Guangzhou, China. Basedon the values of apex angles, leaf apices can be roughly classifiedinto two categories: acute (0° < α < 90°) and obtuse (90° < α <180°) (diagram in Fig. 1A) (20). Triangular leaf apices with theacute apex, marked with green dots in Fig. 1A, commonly exist inareas with high precipitation (SI Appendix, Table S2).Besides the category of apex angles, apex shapes, i.e., rounded,

acuminate, and drip tip, are also used for classifying the leafapices (20). The acuminate apex indicates the margin betweenthe apex and the lamina is convex proximally and concave dis-tally, or concave only. The drip tip is one special kind of acu-minate, where the distal portion of the apex abruptly narrows.Four representative leaves with distinct apex shapes are shown in

Significance

Liquid manipulation is of significance not only for industrialspraying and drainage facilities, but also for the survival ofcreatures. Plant leaves perform excellently in rainwater drain-age and leaf drying at the apex to avoid damage. Here wedemonstrate that apex structure enhances water sheddingwith high dripping frequencies and low retention volumes.Based on the understanding of the tiny apex structure in con-trolling water delivery at the plant leaf, the evolutionary lawof leaf apex shape could be further revealed. The shape-controlled liquid manipulation mechanism would improve themicrofluidic and drainage systems.

Author contributions: Z.D. and L.J. designed research; T.W., Y.S., Z.D., and L.J. performedresearch; T.W., Y.S., H.D., C.L., C.G., Z.D., and L.J. contributed new reagents/analytic tools;T.W., Y.S., H.D., C.L., C.G., Z.D., and L.J. analyzed data; and T.W., H.D., Z.D., and L.J. wrotethe paper.

The authors declare no competing interest.

This article is a PNAS Direct Submission.

This open access article is distributed under Creative Commons Attribution-NonCommercial-NoDerivatives License 4.0 (CC BY-NC-ND).1To whom correspondence may be addressed. Email: dongzhichao@iccas.ac.cn orjianglei@iccas.ac.cn.

This article contains supporting information online at https://www.pnas.org/lookup/suppl/doi:10.1073/pnas.1909924117/-/DCSupplemental.

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Fig. 1 B–E, as marked with circles in Fig. 1A and magnified inFig. 1 F–I. The leaf apex maintains rounded shape, such as theapex of Robinia pseudoacacia (Fig. 1B), or grows as an acuminateshape, such as the apex of Magnolia denudata desr. and Colocasiagigantea (Fig. 1 C and D), or grows as the drip tip, such as theapex of A. macrorrhiza (Fig. 1E and Movie S1). The detailedapex morphologies of these four distinct representative speciesalso vary and change from circular (Fig. 1F), elliptical (Fig. 1G),triangular (Fig. 1H), to triangular grooved with curvature tran-sition shape (Fig. 1I), respectively, which may play a role indrainage behavior. In particular, plants live in rainforests withhigh precipitation tend to evolve leaves with large surface areas(Fig. 1 D and E) and acuminate apices (Fig. 1D) or drip tips (Fig.1 E and I and SI Appendix, Fig. S1) at the leaf tips. Biologistsconsider that the drip tip is a classical characteristic of rainforestleaves through long-time evolution (12–16). Numerous biologicalfunctions of leaves can be achieved or even improved via the driptip. For example, rapid water drainage at drip tip will benefit the

drying (12–16) of leaf, which decreases the need for leaf support,reduces organism colonization (17), and facilitates transpiration(21). The underlying mechanism of apex morphology andstructure in determining the drainage efficiency, therefore, isneeded to be demonstrated.Fig. 1J reveals the water drainage ability of the above three apex

shapes. The water weight per square centimeter, ω, on three se-lected leaf surfaces with rounded, acuminate, and drip tip wasmeasured in real time by microbalance under a constant leaf in-clination angle (the angle between the leaf surface and the vertical),β, of 30° and the same heavy mist of ∼90% relative humidity. Theaverage values of the threshold droplet weight, ωThreshold, and waterretention weight, ωRetention, for different leaf apices are shown onthe right side. Notably, the drip tip of A. macrorrhiza performsexcellent water drainage with dripping frequency, dripping volume,and water retention values ∼1.8, ∼1/6, and ∼1/10 than those of R.pseudoacacia with rounded leaf apex, respectively.

Shortened Drainage Time of an Inclined Artificial Triangular Apex. Todetermine the underlying drainage mechanism, we create artificialleaf apices that are simplified into three geometrical models, in-cluding circle, triangle, and pendulous triangle (curved acuminate)models, through 3D printing (22) and laser-cutting techniques (SIAppendix, Table S4). X-radiation and high-speed cameras are usedto record the profile of dripping droplets at the leaf apices. Weinvestigate the effect of apex angle and apex shape on drainagebased on the 2D and 3D leaf morphologies. Apices changingfrom the circular, elliptical, large triangle, to small triangle, haveshown different drainage ability with distinct threshold drop vol-ume, Ωc, and drainage time, τth (Fig. 2). The smallest triangularapex (α = 30°) has the lowest threshold drop volume Ωc compared

Fig. 1. Water drainage on leaves with different leaf apices. (A) The statis-tical plot of the leaf apex angles, α, among different species of leaves inalphabetical order. According to the leaf apex angle, leaf apex can beclassified into obtuse and acute. SI Appendix, Table S1 shows 44 species and36 families of these leaves in alphabetical order. (B–I) Four representativeleaves have various apex shapes: rounded (B), acuminate (C and D), and driptip (E). Moreover, the corresponding optical images of the magnified apicesare shown as round (F), elliptical (G), triangular (H), and triangular withcurvature transition (I), respectively. (J) Water-weight per area, ω, for variousshapes of leaf apices versus time under a heavy mist of 90% relative hu-midity (Left) and the corresponding average threshold droplet weight,ωThreshold, and water-retention weight, ωRetention (Right). The measurementtests are conducted in the same leaf apex area of 2.0 cm2 and the sameinclination angle with β = 30°. The drip tip has a high dripping frequency,low threshold droplet weight, and low water retention. (Scale bar, 1 mm.)

Fig. 2. Sharp triangular apex shortens threshold drainage time under asteep inclination. (A–D) Profiles of water droplets in the threshold state atvarious apex margins from the top view, round α = 180° (A), ellipse (B), largetriangle α = 90° (C), and small triangle α = 30° (D). (E–G) Mechanism of waterdrainage around a sharp apex in a quasistatic state. An accumulating dropletis pinned by the TCL, as marked with a red dashed line. The drop contactwidth, W, changes from threshold state ① to state ② under the balancebetween the front resistance force of capillarity and the driving force of thegravity component. The front resistance force decreases with contact dropwidth, W, and apex angle α. (H) Threshold drainage time, τth, decreases withthe apex angle, α, and inclination angle, β. A sharp triangular apex shortensthe τth value of the accumulating droplet to only 1/4 of that for a circularapex. (Scale bar, 2 mm.)

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to other shapes of apex models under the same inclination angle(β = 30°) (Fig. 2 A–D).In the experiment, water continuously accumulated on the

surface of the leaf apex at a speed of 1.0 mL/min and shed off atthe apex. The shedding of a sessile drop starts on an inclinedsurface with a tilt angle of β, when the gravitational force com-ponent ∼ρgΩcosβ is greater than the capillary force of the dropletat the front of apex ∼ γWcosθ (23–25), where ρ is the liquiddensity, g is gravity, Ω is the droplet volume, γ is the liquid surfacetension, θ is the apparent contact angle, andW is the contact widthof the drop at the front of apex, respectively (Fig. 2 E–G and SIAppendix, Fig. S2 A–C). A critical depinning drop with a thresholddrop volume Ωc and a threshold contact width Wc exists such thatWc/Ωc ∼ cosβ/ j lc2cosθ j , where lc = (γ/ρg)0.5 is the capillary length.Hence, for a surface with a constant contact angle and inclinationangle, a critical shedding droplet has a threshold drop contactwidth of Wc and a threshold volume Ωc. Ωc is proportional to Wc.As α has a positive relationship with Wc, α has a positive re-lationship with Ωc. Besides, a steeply inclined leaf surface with asmall β increases the component of gravity, which induces a lowthreshold drop volume Ωc. Therefore, a leaf with a triangular apexunder specific inclined orientation provides a practical design forreducing the threshold drop volume.The threshold droplet drainage time, τth, is proportional to the

threshold drop volume Ωc and inversely proportional to thedripping frequency. For a dripping droplet with constant flowvelocity and a sharp apex angle, a small Ωc reflects a reducedthreshold dripping time τth. Then, we investigate the effect of theapex angle on water drainage time under the inclination angle ofthe leaf surface between 0° and 85° at a velocity of 1.0 mL/min(Fig. 2H). Through mimicking the apex structures of naturalleaves, we design two-dimension artificial 3D printing leaf apiceswith apex angles in a range between 0° and 180°. τth decreases withα around the apex under the same inclination angle. For instance,the τth value of a flat circular apex is 53.52 ± 1.52 s when β equals80°, whereas the τth value of a flat triangular apex (FTA, α = 30°)is only 13.86 ± 0.33 s under the same β. A nearly 75% reductionoccurs when the leaf apex changes from circular to acute. Besides,a small α leads to a low apex sticking force and little water re-tention, thus enhancing the drainage efficiency (SI Appendix, Fig.S3). A narrow acute apex or a steeply inclined orientation leads toa small threshold drop volume and a short dripping time. Notably,more than 80% of the leaves in Fig. 1A have triangular apices.

Reduced Water Drainage Retention on Pendulous Triangular GroovedApices. Based on the analysis in Fig. 2H, a steeply inclined tri-angular leaf apex is the best state for drainage. However, thenatural leaves give another answer. We observe and record theinclination angles of four species of leaves, Alocasia macrorrhiza(Lour.) Scott, M. denudate desr., Sophora japonica Linn., andCornus controversa at different times of the day under sunshineand rainfall, respectively (SI Appendix, Fig. S4). The leaves ofsubtropical zone possess a relatively large inclination angle, β, ina range of 50°–70° under sunshine for better photosynthesis anda relatively small inclination angle, β, in a range of 20°–40° forbetter drainage and avoiding damage (26, 27). In the rainforestwith frequent heavy rainfall, the leaf of A. locasia macrorrhiza(Lour.) tilts a large β1 in the base to guarantee maximum sunlightexposure, and a small β2 in the apex to satisfy the need for fastdrainage (28, 29). The leaf surface curves from a 2D shape to a3D shape (the pendulous shape) at the leaf apex, which couldimpose an instability force on the threshold droplet.The artificial leaf apex model is designed to be a version of the

pendulous triangular shape or pendulous grooved triangularshape (Fig. 3). Threshold droplets form a stable triple contactline (TCL) at a given threshold drop contact width W on a flattriangular apex (FTA) (Fig. 3A). When the bending position ofthe pendulous triangular apex (PTA) is located at the threshold

contact line, the drop remains stable (Fig. 3B). If the bendingposition is located above the threshold contact line, the dropletwould experience a sudden instability induced by the increasedgravitational force component and decreased resistant capillarityat the bending point. The reconstructed balance forces the waterdroplet to drip off with a smaller volume than for other flat apexshapes (Fig. 3C and SI Appendix, Fig. S2E).In addition to a lower threshold dripping volume, the PTA

also reduces water retention on the leaf surface compared withflat triangular apex. The water retention on the surface persquare meter, wRe, at flat triangular apex and PTA were mea-sured under the same apex angle range and the inclination anglerange (15° < α < 180°, 30° < β < 60°) at a water injection velocityof 1.0 mL/min (Fig. 3D). The wRe on a pendulous circular apex(α = 180°, β1 = 60°, β2 = 0°) is only 3/5 of that on a flat circularapex (α = 180°, β = 60°). A PTA (α = 15°, β1 = 60°, β2 = 0°)further reduces wRe to 1/21 of that on the flat circular apex (α =180°, β = 60°).The use of PTA in design was common in Eastern Asia ancient

architectures, including tile art, especially for the drip tiles on theedges of eaves (30, 31). Drip tile shows a similar function withthe pendulous triangular leaf apex. The upper surface of the driptile possesses a slight inclination of the tile surface in base with alarge β1 to converge and transport water. The downward triangularapex possesses a steep inclination with a small β2 to achieve fastrainwater runoff, decrease the requirements for roof support, andreduce the erosion of the architecture base (Fig. 3E).Significantly, leaf veins in plants form the gradient of groove,

further promoting water shedding under capillary force (32, 33).The large gradient of the groove reduces the threshold drainagetime (Fig. 3F). The gradient of the conical groove is controlledby the ratio of the large side of radius R and a small side of radiusr (r = 5 mm), the radius ratio (R/r) of which changes from 1 to 4(SI Appendix, Fig. S5A). We design the PTA with a groovestructure by folding along the artificial apex axis with a folding

Fig. 3. PTGA reduces drainage retention. (A–C) The resistance force of anaccumulating droplet on an FTA, α = 30° and β1 = 30°) (A) and a PTA, α = 30°,β1 = 30°, and β2 = 0°) (B and C) at the threshold moment. The thresholddroplet is pinned by the TCL, as marked with a red dotted line. (D) The waterdrainage retention per area, wRe, decreases with the apex angle, α, andinclination angle, β, for the FTA and PTA. (E) Drip tile has a similar structurewith a PTA structure of leaf for fast drainage. (F) Threshold drainage timedecreases with the gradient of the conical channel. The two conical endshave a large radius of R and a small radius of r. The small side of the channelhas a radius of 5 mm. (G) Compared with FTA and PTA, the PTGA has thelowest water retention (wRe).

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angle of 120° (SI Appendix, Fig. S5B). Pendulous triangulargrooved apex (PTGA) can significantly increase the drainageefficiency by reducing water retention and decreasing thresholddrainage time. The pendulous triangular grooved apex (α = 15°,β1 = 60°, β2 = 0°) reduces the water retention, wRe, to 0.250 ±0.100 kg/m2, and with only 4% the retention of the flat circularapex (Fig. 3G). Therefore, the PTGA improves drainage effi-ciency by reducing water retention and threshold drainage time.The understanding of this drainage mechanism from leaf apexand architecture drip tile can be used to design drainage facilitiesand solve problems in printing and open-air fluidic systems (34–36) that are currently faced.

Gaussian Curvature Inversion at the Drip-Tip Apex PromotesThreshold Droplet Shedding. The leaf drip tip of A. macrorrhizain rainforest areas has a long and curvy acuminate apex with aPTGA structure (Fig. 4A). Water drains at the leaf drip tip anddetaches at the groove outlet (Fig. 4C). Detailed observations ofthe drip tip apex are shown in Fig. 4 B and D–F and Movie S1.For the drip tip of A. macrorrhiza, the microchannel curvesoutward in a concave open state, and the end of the apex curvesinward in a convex closed state (Fig. 4B). At the closing outlet ofthe groove, the concave groove transforms into a convex apexfrom the front and top view (Fig. 4 B and D), with the curvaturechanging from negative to positive along with the groove outlet

from the radial cross-sectional view (Fig. 4E). The curve outlineof the drip tip shows a positive curvature from the axial cross-sectional view (Fig. 4F). The Gaussian curvature along with driptip changes from negative to positive correspondingly (SI Ap-pendix, Fig. S6).As water drains along the surface of the drip tip, the gradually

closing groove with outward concave shape forces water to thedistal end of the apex (blue arrow in Fig. 4G). The structuretransition from the concave surface to the convex surface at theoutlet of the groove rapidly reverses its curvature, resulting in theentrapment of an air wedge with a minimum drop contact widthW (Fig. 4G). The reduced air–water–solid contact line promotesdroplet shedding (SI Appendix, Fig. S2F). Besides, local pressurecaused by surface curvature makes a difference in water shed-ding from the cross-sectional view (Fig. 4H). The upward pres-sure difference drives water from the neck to the upper site,ΔPupper = P2 − P1 = γ (1/R2 − 1/R1), and the downward pressuredifference drives water from the neck to the lower site, ΔPdown =P2 − P3 = γ (1/R3 − 1/R2), where Pi is the local pressure, Ri is thelocal radius of the droplet perpendicular to the plane. Gaussiancurvature transition changes from negative at the concave grooveto positive at the convex apex along the drip tip. The inversion ofGaussian curvature at the drip tip leads to the reverse of theLaplace pressure direction, promoting the necking, separation,and shedding of threshold droplets at the curvature transitionpoint (Fig. 4 F–I). Significantly, the inversion of Gaussian cur-vature results in the local pressure difference (or the inversion ofLaplace pressure difference) and droplet necking and detachingoff, which is different from the liquid spontaneously breaking upinto droplets. These two processes are similar in the result butdifferent in origin.

Fast Water Drainage of the Bioinspired Drip-Tip Apex.X-ray scan canmeasure the drip-tip apex morphologies in detail and reconstructthe drip-tip morphologies for the biomimetic 3D printed model(Fig. 5A and SI Appendix, Fig. S5C). Considering the curvature

Fig. 4. Gaussian curvature inversion at the drip tip promotes thresholddroplet shedding. (A) The integration of the drainage structure of the pen-dulous grooved triangular apex and the spatial curvature inversion at the drip-tipapex of the rainforest leaf A. macrorrhiza. (C) The drip-tip apex with curvatureinversion at the outlet of the groove structure promotes the detachment of thethreshold droplet. (B and D) Magnified scanning electron microscope (SEM)images of the structure of the drip-tip apex from the front and top views.The surface of drip tip changes from the concave shape to the convex shapealong the groove outlet at the apex. (E and F) X-ray slices of the structure ofthe drip tip from the cross-sectional view. A Gaussian curvature transitionfrom negative of the concave shape to the positive of the convex shapeoccurs. (G) Force mapping of the threshold droplet under curvature inversionat the outlet of the groove. Minimum resistance of the threshold dropletwith a characteristic length ofW at the concave–convex curvatures transitionpoint. (H and I) Threshold droplet detaches at the outlet of the groove witha separation point at concave–convex curvatures inversion point from topand side views under pressure-driven process. (Scale bar, 300 μm.)

Fig. 5. Fast water drainage efficiency of the bioinspired drip tip. (A) Illus-tration of bioinspired drip-tip model. (B) Fast water drainage at the artificial tipwith curvature transition from concave shape to convex shape. Tip with curva-ture transition performs better droplet shedding than the tip with one curva-ture, with a fixed separation point and a nearly ∼56% reduction of the drainagetime under an injection liquid speed of 1200 mL/h. (C) Water weight per area, ω,of artificial leaf apices versus time under a heavy mist of approximately ∼90%relative humidity (Left) and the corresponding average threshold droplet weightand water-retention weight (Right). The bioinspired drip-tip apex (BDTA) yields ahigh dripping frequency, small dripping volume, and low water retention. LTA,large triangular apex; STA, small triangular apex. (Scale bar, 3 mm.)

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inversion from concave shape to convex shape is the unique partof the drip tip, we demonstrate the effect of the curvature in-version structure on enhancing water drainage by a control ex-periment (Fig. 5B). Comparing the leaf tip with only one convexcurvature, the artificial tip with concave–convex curvatures canreduce nearly 56% of the drainage time. Besides, the lower theconvex shape of the drip tip than the concave shape can enhancethe water drainage with a much shorter drainage time (SI Ap-pendix, Fig. S7).To better evaluate the drainage performance, we compare the

drainage efficiency of the bioinspired drip tip and other artificialtips (Fig. 5C). The experiment was performed under a heavy mistof ∼90% relative humidity and a large inclination angle β = 30°to mimic the real-leaf apex dripping process. The sample surfacehas the same area of 2.0 cm2. The water weight per squarecentimeter, ω, of the PTGA reflects excellent drainage ability,with a dripping frequency that is ∼6 times greater than that for alarge triangular apex and water retention that is ∼1/4 that for alarge triangular apex. The bioinspired drip tip apex (BDTA)performs even better on drainage, with a dripping frequency ∼12times greater than that for a large triangular apex and with waterretention ∼1/6 than that for a large triangular apex. The averagevalues of the threshold droplet weight and water retentionweight for different artificial leaf apices are shown in Fig. 5 C,Right. The bioinspired drip-tip apex has the least amount ofaccumulated water at the leaf apex over the longest time, whichshows the best drainage efficiency among all of the artificialapices studied. Following the movement of the raindrops at plantleaves, the special posture and unique structure of the leaf apexinspire us to perform biomimetic researches that optimize arti-ficial leaf apex structures with superior drainage abilities.

ConclusionsWater drainage is enhanced by the apex under the instabilityforce. The enhanced instability is caused by the reduced capillaryresistance of droplet at acuminate apex shape and the increasedgravity component of a droplet under steep apex orientation.

The multiple steric curvature structures and efficient drainagemodes of leaf apices can be studied to understand the roles of“tiny” structures in plant survival control and provide in-spiration for upgrading the drainage facilities and anticorrosionarchitecture. Such information may offer insights into the fieldof water collection with rapid water shedding or transport.Moreover, a different perspective for solving scientific andtechnological problems, such as those related to printing andopen-air fluidic applications, may be developed to achieve high-speed spatial water delivery. Besides, the relevant results can beused to developing cutting-edge commercial drop and fiberfabrication processes, such as fog/drug nano/microdroplet fab-rication via spraying and fiber construction via electrospinning.

Materials and MethodsDifferent species of plant leaves were collected from the subtropical zone ofBeijing and the tropical zone of Guangzhou. The leaf surfaces were cleanedby water and 50% ethanol concentration in sequences and dried by N2 (SIAppendix, Fig. S8). Flat artificial apices and bioinspired drip tip were madeby a 3D printer (Formlabs 2), and the artificial PTA and PTGA were made bylaser cutting (LSC30 CO2 laser). Fog-shedding experiment was conducted underthe dense fog mist with ∼90% relative humidity, and water weight wasmeasured on the microbalance in real time. The selected surface area of theartificial leaf apex is 2 cm2. The water-shedding experiment was captured by ahigh-speed camera (iSPEED-3, Olympus). Water flowed from a nozzle per-pendicular to the leaf surface at a constant injection speed. The water-flowvelocity was controlled by a microinjection gear pump (Harvard). The effect ofincoming drop on the dripping drop at the drip-tip leaf (SI Appendix, Fig. S9)and the influence of leaf wettability (37, 38) on drainage at the apex (SI Ap-pendix, Table S3 and Fig. S10) were illustrated. Other detailed experimentalprocedures are illustrated in SI Appendix, SI Materials and Methods.

Data Availability Statement. All data discussed in the paper is available withinthe SI Appendix and Dataset S1.

ACKNOWLEDGMENTS. This work was supported by the National NaturalScience Foundation (21703270, 21988102), the National Key R&D Program ofChina (2018YFA0208501, 2017YFA0206901), and the Key Research Programof the Chinese Academy of Sciences (KJZD-EW-M01).

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