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1.  Biomimetics inspired surfaces for drag reduction and oleophobicity/philicity 
Summary
The emerging field of biomimetics allows one to mimic biology or nature to develop nanomaterials, nanodevices, and processes which provide desirable properties. Hierarchical structures with dimensions of features ranging from the macroscale to the nanoscale are extremely common in nature and possess properties of interest. There are a large number of objects including bacteria, plants, land and aquatic animals, and seashells with properties of commercial interest. Certain plant leaves, such as lotus (Nelumbo nucifera) leaves, are known to be superhydrophobic and self-cleaning due to the hierarchical surface roughness and presence of a wax layer. In addition to a self-cleaning effect, these surfaces with a high contact angle and low contact angle hysteresis also exhibit low adhesion and drag reduction for fluid flow. An aquatic animal, such as a shark, is another model from nature for the reduction of drag in fluid flow. The artificial surfaces inspired from the shark skin and lotus leaf have been created, and in this article the influence of structure on drag reduction efficiency is reviewed. Biomimetic-inspired oleophobic surfaces can be used to prevent contamination of the underwater parts of ships by biological and organic contaminants, including oil. The article also reviews the wetting behavior of oil droplets on various superoleophobic surfaces created in the lab.
doi:10.3762/bjnano.2.9
PMCID: PMC3148050  PMID: 21977417
aquatic animals; biomimetics; drag; lotus plants; shark skin; superhydrophobicity; superoleophobicity
2.  Non-Contaminating Camouflage: Multifunctional Skin Microornamentation in the West African Gaboon Viper (Bitis rhinoceros) 
PLoS ONE  2014;9(3):e91087.
The West African Gaboon viper (Bitis rhinoceros) has an extraordinary coloration of pale brown and velvety black markings. The velvety black appearance is caused by a unique hierarchical surface structures which was not found on the pale brown scales. In the present study we examined the wettability of the vipeŕs scales by measuring contact angles of water droplets. Velvet black scale surfaces had high static contact angles beyond 160° and low roll-off angles below 20° indicating an outstanding superhydrophobicity. Our calculations showed that the Cassie-Baxter model describes well wettability effects for these surfaces. Self-cleaning capabilities were determined by contaminating the scales with particles and fogging them until droplets formed. Black scales were clean after fogging, while pale scales stayed contaminated. Black scales feature multifunctional structures providing not only water-repellent but also self-cleaning properties. The pattern of nanoridges can be used as a model for surface-active technical surfaces.
doi:10.1371/journal.pone.0091087
PMCID: PMC3944882  PMID: 24599379
3.  Robust adhesion of flower-like few-layer graphene nanoclusters 
Scientific Reports  2012;2:511.
Nanostructured surface possessing ultrahigh adhesion like “gecko foot” or “rose petal” can offer more opportunities for bionic application. We grow flower-like few-layer graphene on silicon nanocone arrays to form graphene nanoclusters, showing robust adhesion. Their contact angle (CA) is 164° with a hysteresis CA of 155° and adhesive force for a 5 μL water droplet is about 254 μN that is far larger than present reported results. We bring experimental evidences that this great adhesion depends on large-area plentiful edges of graphene nanosheets tuned by conical nanostructure and intrinsic wetting features of graphene. Such new hierarchical few-layer graphene nanostructure provides a feasible strategy to understand the ultra-adhesive mechanism of the “gecko effect” or “rose effect” and enhance the wettability of graphene for many practical applications.
doi:10.1038/srep00511
PMCID: PMC3397321  PMID: 22803004
4.  Stable superhydrophobic surface of hierarchical carbon nanotubes on Si micropillar arrays 
Nanoscale Research Letters  2013;8(1):412.
It is of great importance to construct a stable superhydrophobic surface with low sliding angle (SA) for various applications. We used hydrophobic carbon nanotubes (CNTs) to construct the superhydrophobic hierarchical architecture of CNTs on silicon micropillar array (CNTs/Si-μp), which have a large contact angle of 153° to 155° and an ultralow SA of 3° to 5°. Small water droplets with a volume larger than 0.3 μL can slide on the CNTs/Si-μp with a tilted angle of approximately 5°. The CNTs growing on planar Si wafer lose their superhydrophobic properties after exposing to tiny water droplets. However, the CNTs/Si-μp still show superhydrophobic properties even after wetting using tiny water droplets. The CNTs/Si-μp still have a hierarchical structure after wetting, resulting in a stable superhydrophobic surface.
doi:10.1186/1556-276X-8-412
PMCID: PMC3874759  PMID: 24098965
Carbon nanotube; Hierarchical architecture; Superhydrophobic surface
5.  Under-water superoleophobic Glass: Unexplored role of the surfactant-rich solvent 
Scientific Reports  2013;3:1862.
Preparing low energy liquid-repellant surfaces (superhydrophobic or superoleophobic) have attracted tremendous attention of late. In all these studies, the necessary liquid repellency is achieved by irreversible micro-nano texturing of the surfaces. Here we show for the first time that a glass surface, placed under water, can be made superoleophobic (with unprecedented contact angles close to 180 degrees and roll off angles only a few fractions of 1 degree) by merely changing the surfactant content of the water medium in which the oil (immiscible in water) has been dispersed. Therefore, we propose a paradigm shift in efforts to achieve liquid-repellant systems, namely, altering the solvent characteristics instead of engineering the surfaces. The effect occurs for a surfactant concentration much larger than the critical micelle concentration, and is associated to strong adsorption of surfactant molecules at the solid surface, triggering an extremely stable Cassie-Baxter like conformation of the oil droplets.
doi:10.1038/srep01862
PMCID: PMC3659319  PMID: 23689477
6.  Capillary origami: superhydrophobic ribbon surfaces and liquid marbles 
Summary
In the wetting of a solid by a liquid it is often assumed that the substrate is rigid. However, for an elastic substrate the rigidity depends on the cube of its thickness and so reduces rapidly as the substrate becomes thinner as it approaches becoming a thin sheet. In such circumstances, it has been shown that the capillary forces caused by a contacting droplet of a liquid can shape the solid rather than the solid shaping the liquid. A substrate can be bent and folded as a (pinned) droplet evaporates or even instantaneously and spontaneously wrapped on contact with a droplet. When this effect is used to create three dimensional shapes from initially flat sheets, the effect is called capillary origami or droplet wrapping.
In this work, we consider how the conditions for the spontaneous, capillary induced, folding of a thin ribbon substrate might be altered by a rigid surface structure that, for a rigid substrate, would be expected to create Cassie–Baxter and Wenzel effects. For smooth thin substrates, droplet wrapping can occur for all liquids, including those for which the Young’s law contact angle (defined by the interfacial tensions) is greater than 90° and which would therefore normally be considered relatively hydrophobic. However, consideration of the balance between bending and interfacial energies suggests that the tendency for droplet wrapping can be suppressed for some liquids by providing the flexible solid surface with a rigid topographic structure. In general, it is known that when a liquid interacts with such a structure it can either fully penetrate the structure (the Wenzel case) or it can bridge between the asperities of the structure (the Cassie–Baxter case).
In this report, we show theoretically that droplet wrapping should occur with both types of solid–liquid contact. We also derive a condition for the transition between the Cassie–Baxter and Wenzel type droplet wrapping and relate it to the same transition condition known to apply to superhydrophobic surfaces. The results are given for both droplets being wrapped by thin ribbons and for solid grains encapsulating droplets to form liquid marbles.
doi:10.3762/bjnano.2.18
PMCID: PMC3148044  PMID: 21977426
capillary origami; Cassie; contact angle; superhydrophobic; Wenzel
7.  Influence of Cuticle Nanostructuring on the Wetting Behaviour/States on Cicada Wings 
PLoS ONE  2012;7(4):e35056.
The nanoscale protrusions of different morphologies on wing surfaces of four cicada species were examined under an environmental scanning electron microscope (ESEM). The water contact angles (CAs) of the wing surfaces were measured along with droplet adhesion values using a high-sensitivity microelectromechanical balance system. The water CA and adhesive force measurements obtained were found to relate to the nanostructuring differences of the four species. The adhesive forces in combination with the Cassie-Baxter and Wenzel approximations were used to predict wetting states of the insect wing cuticles. The more disordered and inhomogeneous surface of the species Leptopsalta bifuscata demonstrated a Wenzel type wetting state or an intermediate state of spreading and imbibition with a CA of 81.3° and high adhesive force of 149.5 µN. Three other species (Cryptotympana atrata, Meimuna opalifer and Aola bindusara) exhibited nanostructuring of the form of conically shaped protrusions, which were spherically capped. These surfaces presented a range of high adhesional values; however, the CAs were highly hydrophobic (C. atrata and A. bindusara) and in some cases close to superhydrophobic (M. opalifer). The wetting states of A. bindusara, C. atrata and M. opalifer (based on adhesion and CAs) are most likely represented by the transitional region between the Cassie-Baxter and Wenzel approximations to varying degrees.
doi:10.1371/journal.pone.0035056
PMCID: PMC3335046  PMID: 22536351
8.  Superhydrophobic Surface Based on a Coral-Like Hierarchical Structure of ZnO 
PLoS ONE  2010;5(12):e14475.
Background
Fabrication of superhydrophobic surfaces has attracted much interest in the past decade. The fabrication methods that have been studied are chemical vapour deposition, the sol-gel method, etching technique, electrochemical deposition, the layer-by-layer deposition, and so on. Simple and inexpensive methods for manufacturing environmentally stable superhydrophobic surfaces have also been proposed lately. However, work referring to the influence of special structures on the wettability, such as hierarchical ZnO nanostructures, is rare.
Methodology
This study presents a simple and reproducible method to fabricate a superhydrophobic surface with micro-scale roughness based on zinc oxide (ZnO) hierarchical structure, which is grown by the hydrothermal method with an alkaline aqueous solution. Coral-like structures of ZnO were fabricated on a glass substrate with a micro-scale roughness, while the antennas of the coral formed the nano-scale roughness. The fresh ZnO films exhibited excellent superhydrophilicity (the apparent contact angle for water droplet was about 0°), while the ability to be wet could be changed to superhydrophobicity after spin-coating Teflon (the apparent contact angle greater than 168°). The procedure reported here can be applied to substrates consisting of other materials and having various shapes.
Results
The new process is convenient and environmentally friendly compared to conventional methods. Furthermore, the hierarchical structure generates the extraordinary solid/gas/liquid three-phase contact interface, which is the essential characteristic for a superhydrophobic surface.
doi:10.1371/journal.pone.0014475
PMCID: PMC3012683  PMID: 21209931
9.  Superhydrophobic nature of nanostructures on an indigenous Australian eucalyptus plant and its potential application 
In this preliminary study, the morphology and nanostructured features formed by the epicuticular waxes of the mottlecah (Eucalyptus macrocarpa) leaf were investigated and quantified. The surface features formed by the waxes give the leaf remarkable wetting and self-cleaning properties that enhance the plant’s survival in an arid climate. This paper also provides experimental evidence of the self-assembly properties of the epicuticular waxes. Analysis of the water contact angle measurements gave a mean static contact angle of 162.00 ± 6.10 degrees, which clearly indicated that the mottlecah’s leaf surface was superhydrophobic. Detailed field emission scanning electron microscopy examination revealed that the surface was covered by bumps approximately 20 μm in diameter and regularly spaced at a distance of around 26 μm. The bumps are capped by nanotubules/pillars with an average diameter of 280 nm at the tips. Self-cleaning experiments indicated that the mottlecah’s leaf could be effectively cleaned by a fine spray of water droplets that rolled over the surface picking up contaminants. Field emission scanning electron microscopy investigation of extracted epicuticular waxes revealed that the waxes were capable of self-reassembly and formed features similar to those of the original leaf surface. Furthermore, also reported is a simple technique for surface treating one side of a planar surface to produce a superhydrophobic surface that can be used as a planar floatation platform for microdevices.
doi:10.2147/NSA.S24834
PMCID: PMC3781715  PMID: 24198490
superhydrophobic; nanopillars; epicuticular waxes; self-cleaning; floatation; micro-fluidic
10.  Flower thermoregulation facilitates fertilization in Asian sacred lotus 
Annals of Botany  2009;103(7):1159-1163.
Background and Aims
The thermoregulatory flower of the Asian sacred lotus (Nelumbo nucifera) can maintain a relatively stable temperature despite great variations in ambient temperature during anthesis. The thermoregulation has been hypothesized to offer a direct energy reward for pollinators in lotus flowers. This study aims to examine whether the stable temperature maintained in the floral chamber influences the fertilization process and seed development.
Methods
An artificial refrigeration instrument was employed to cool flowers during the fertilization process and post-fertilization period in an experimental population. The effect of temperature on post-pollination events was also examined by removing petals in two field populations.
Key Results
Treatments with low floral temperature did not reduce stigma receptivity or pollen viability in undehisced anthers. Low temperature during the fertilization period significantly decreased seed set per flower but low temperature during the phase of seed development had no effect, suggesting that temperature regulation by lotus flowers facilitated fertilization success. Hand-pollination treatments in two field populations indicated that seed set of flowers with petals removed was lower than that of intact flowers in north China, where ambient temperatures are low, but not in south China, confirming that reducing the temperature of carpels did influence post-pollination events.
Conclusions
The experiments suggest that floral thermoregulation in lotus could enhance female reproductive success by facilitating fertilization.
doi:10.1093/aob/mcp051
PMCID: PMC2707905  PMID: 19282320
Nelumbo nucifera; Asian sacred lotus; beetle-pollination syndrome; fertilization process; post-pollination events; pollen viability; stigma receptivity; thermoregulation
11.  Superhydrophobic Photosensitizers. Mechanistic Studies of 1O2 Generation in the Plastron and Solid/Liquid Droplet Interface 
Journal of the American Chemical Society  2013;135(50):18990-18998.
We describe here a physical-organic study of the first triphasic superhydrophobic sensitizer for photooxidations in water droplets. Control of synthetic parameters enables the mechanistic study of “borderline” two- and three-phase superhydrophobic sensitizer surfaces where 1O2 is generated in compartments that are wetted, partially wetted, or remain dry in the plastron (i.e., air layer beneath the droplet). The superhydrophobic surface is synthesized by partially embedding silicon phthalocyanine (Pc) sensitizing particles to specific locations on polydimethylsiloxane (PDMS) posts printed in a square array (1 mm tall posts on 0.5 mm pitch). In the presence of red light and oxygen, singlet oxygen is formed on the superhydrophobic surface and reacts with 9,10-anthracene dipropionate dianion (1) within a freestanding water droplet to produce an endoperoxide in 54–72% yields. Control of the 1O2 chemistry was achieved by the synthesis of superhydrophobic surfaces enriched with Pc particles either at the PDMS end-tips or at PDMS post bases. Much of the 1O2 that reacts with anthracene 1 in the droplets was generated by the sensitizer “wetted” at the Pc particle/water droplet interface and gave the highest endoperoxide yields. About 20% of the 1O2 can be introduced into the droplet from the plastron. The results indicate that the superhydrophobic sensitizer surface offers a unique system to study 1O2 transfer routes where a balance of gas and liquid contributions of 1O2 is tunable within the same superhydrophobic surface.
doi:10.1021/ja410529q
PMCID: PMC3930608  PMID: 24295210
12.  Surface Structure and Wetting Characteristics of Collembola Cuticles 
PLoS ONE  2014;9(2):e86783.
The cuticles of the arthropods Collembola (springtails) are known to be superhydrophobic, displaying such properties as water-repellence and plastron formation; overhanging surface structures have been suggested as the source of these properties. Superhydrophobicity is closely related to surface structuring and other surfaces with overhanging structures have been shown to possess robust superhydrophobic properties. In effort to correlate the wetting performance and surface structuring of the cuticles, from both a technical and evolutionary point of view, we investigated a selection of Collembola species including species from several families and covering habitats ranging from aquatic to very dry. The observed contact angles of wetting was in general larger than those predicted by the conventional models. Not all the studied Collembola were found to have superhydrophobic properties, indicating that superhydrophobicity is common, but not a universal trait in Collembola. Overhanging structures were found in some, but not all Collembola species with superhydrophobic cuticles; which leads to the conclusion that there is no direct link between overhanging surface structures and superhydrophobicity in Collembola.
doi:10.1371/journal.pone.0086783
PMCID: PMC3911920  PMID: 24498281
13.  Preparation and characterization of superhydrophobic surfaces based on hexamethyldisilazane-modified nanoporous alumina 
Nanoscale Research Letters  2011;6(1):487.
Superhydrophobic nanoporous anodic aluminum oxide (alumina) surfaces were prepared using treatment with vapor-phase hexamethyldisilazane (HMDS). Nanoporous alumina substrates were first made using a two-step anodization process. Subsequently, a repeated modification procedure was employed for efficient incorporation of the terminal methyl groups of HMDS to the alumina surface. Morphology of the surfaces was characterized by scanning electron microscopy, showing hexagonally ordered circular nanopores with approximately 250 nm in diameter and 300 nm of interpore distances. Fourier transform infrared spectroscopy-attenuated total reflectance analysis showed the presence of chemically bound methyl groups on the HMDS-modified nanoporous alumina surfaces. Wetting properties of these surfaces were characterized by measurements of the water contact angle which was found to reach 153.2 ± 2°. The contact angle values on HMDS-modified nanoporous alumina surfaces were found to be significantly larger than the average water contact angle of 82.9 ± 3° on smooth thin film alumina surfaces that underwent the same HMDS modification steps. The difference between the two cases was explained by the Cassie-Baxter theory of rough surface wetting.
doi:10.1186/1556-276X-6-487
PMCID: PMC3212001  PMID: 21827683
superhydrophobic surfaces; surface modification; hexamethyldisilazane; nanoporous alumina
14.  Wettability Switching Techniques on Superhydrophobic Surfaces 
Nanoscale Research Letters  2007;2(12):577-596.
The wetting properties of superhydrophobic surfaces have generated worldwide research interest. A water drop on these surfaces forms a nearly perfect spherical pearl. Superhydrophobic materials hold considerable promise for potential applications ranging from self cleaning surfaces, completely water impermeable textiles to low cost energy displacement of liquids in lab-on-chip devices. However, the dynamic modification of the liquid droplets behavior and in particular of their wetting properties on these surfaces is still a challenging issue. In this review, after a brief overview on superhydrophobic states definition, the techniques leading to the modification of wettability behavior on superhydrophobic surfaces under specific conditions: optical, magnetic, mechanical, chemical, thermal are discussed. Finally, a focus on electrowetting is made from historical phenomenon pointed out some decades ago on classical planar hydrophobic surfaces to recent breakthrough obtained on superhydrophobic surfaces.
doi:10.1007/s11671-007-9102-4
PMCID: PMC3246619
Microfluidic; Superhydrophobic surfaces; Wettability switching; Electrowetting
15.  Superhydrophobicity in perfection: the outstanding properties of the lotus leaf 
Summary
Lotus leaves have become an icon for superhydrophobicity and self-cleaning surfaces, and have led to the concept of the ‘Lotus effect’. Although many other plants have superhydrophobic surfaces with almost similar contact angles, the lotus shows better stability and perfection of its water repellency. Here, we compare the relevant properties such as the micro- and nano-structure, the chemical composition of the waxes and the mechanical properties of lotus with its competitors. It soon becomes obvious that the upper epidermis of the lotus leaf has developed some unrivaled optimizations. The extraordinary shape and the density of the papillae are the basis for the extremely reduced contact area between surface and water drops. The exceptional dense layer of very small epicuticular wax tubules is a result of their unique chemical composition. The mechanical robustness of the papillae and the wax tubules reduce damage and are the basis for the perfection and durability of the water repellency. A reason for the optimization, particularly of the upper side of the lotus leaf, can be deduced from the fact that the stomata are located in the upper epidermis. Here, the impact of rain and contamination is higher than on the lower epidermis. The lotus plant has successfully developed an excellent protection for this delicate epistomatic surface of its leaves.
doi:10.3762/bjnano.2.19
PMCID: PMC3148040  PMID: 21977427
epicuticular wax; leaf surface; Lotus effect; papillae; water repellency
16.  Self-similarity of contact line depinning from textured surfaces 
Nature Communications  2013;4:1492-.
The mobility of drops on surfaces is important in many biological and industrial processes, but the phenomena governing their adhesion, which is dictated by the morphology of the three-phase contact line, remain unclear. Here we describe a technique for measuring the dynamic behaviour of the three-phase contact line at micron length scales using environmental scanning electron microscopy. We examine a superhydrophobic surface on which a drop’s adhesion is governed by capillary bridges at the receding contact line. We measure the microscale receding contact angle of each bridge and show that the Gibbs criterion is satisfied at the microscale. We reveal a hitherto unknown self-similar depinning mechanism that shows how some hierarchical textures such as lotus leaves lead to reduced pinning, and counter-intuitively, how some lead to increased pinning. We develop a model to predict adhesion force and experimentally verify the model’s broad applicability on both synthetic and natural textured surfaces.
The motion of liquid drops on surfaces is governed by adhesion forces, but the microscopic mechanism is unclear. Paxson et al. image the dynamic distortion of the edge of a droplet as it moves across a surface, allowing them to predict the wetting ability of different hierarchically textured surfaces.
doi:10.1038/ncomms2482
PMCID: PMC3586717  PMID: 23422660
17.  Biomimetic Transferable Surface for a Real Time Control over Wettability and Photoerasable Writing with Water Drop Lens 
Scientific Reports  2014;4:7407.
We demonstrate a transferable device that can turn wettability of surfaces to sticky or slippy, as per requirement. It is composed of polymeric yarn with a fibrous structure, which can be lifted and placed on any surface to render it the unique wettability properties. We introduce Polyvinylidenefluoride (PVDF) random fiber as biomimetic rose petal surface. When it is decorated with PVDF nanofibers yarns, the random mesh transform from rose petal sticky state into grass leaf slippy state. When it is placed on sticky, hydrophilic metal coin, it converts the surface of the coin to super hydrophobic. Adjustments in the yarn system, like interyarn spacing, can be done in real time to influence its wettability, which is a unique feature. Next, we load the polymer with a photochromic compound for chemical restructuring. It affects the sliding angle of water drop and makes the fibers optically active. We also demonstrate a “water droplets lens” concept that enables erasable writing on photochromic rose petal sticky fibrous surface. The droplet on a highly hydrophobic surface acts as a ball lens to concentrate light onto a hot spot; thereby we demonstrate UV light writing with water lenses and visible light erasing.
doi:10.1038/srep07407
PMCID: PMC4261171  PMID: 25491016
18.  Micro to nano: Surface size scale and superhydrophobicity 
Summary
This work looks at the fundamental question of how the surface mobility of drops in the composite state is related to the size scale of the roughness features of the surface. To this end, relevant literature is first reviewed and the important terms are clarified. We then describe and discuss contact and roll-off angle measurements on a set of hydrophobicized silicon post surfaces for which all parameters except for the surface size scale were held constant. It was found that a critical transition from “sticky superhydrophobic” (composite state with large contact angle hysteresis) to “truly superhydrophobic” (composite state with low hysteresis) takes place as the size of the surface features reaches 1 μm.
doi:10.3762/bjnano.2.38
PMCID: PMC3148034  PMID: 21977446
contact angle; hysteresis; superhydrophobic; wetting
19.  Ability of Bumblebees to Discriminate Differences in the Shape of Artificial Flowers of Primula sieboldii (Primulaceae) 
Annals of Botany  2007;99(6):1175-1182.
Background and Aims
Flower shapes are important visual cues for pollinators. However, the ability of pollinators to discriminate between flower shapes under natural conditions is poorly understood. This study focused on the diversity of flower shape in Primula sieboldii and investigated the ability of bumblebees to discriminate between flowers by combining computer graphics with a traditional behavioural experiment.
Methods
Elliptic Fourier descriptors described shapes by transforming coordinate information for the contours into coefficients, and principal components analysis summarized these coefficients. Using these methods, artificial flowers were created based on the natural diversity of petal shape in P. sieboldii. Dual-choice tests were then performed to investigate the ability of the bumblebees to detect differences in the aspect ratio of petals and the depth of their head notch.
Key Results
The insects showed no significant ability to detect differences in the aspect ratio of the petals under natural conditions unless the morphological distance increased to an unrealistic level. These results suggest the existence of a perception threshold for distances in this parameter. The bumblebees showed a significant preference for narrow petals even after training using flowers with wide petals. The bumblebees showed a significant ability to discriminate based on the depth of the petal head notch after training using artificial flowers with a deep head notch. However, they showed no discrimination in tests with training using extreme distances between flowers in this parameter.
Conclusions
A new type of behavioural experiment was demonstrated using real variation in flower corolla shape in P. sieboldii. If the range in aspect ratios of petals expands much further, bumblebees may learn to exhibit selective behaviour. However, because discrimination by bumblebees under natural conditions was low, there may be no strong selective behaviour based on innate or learned preferences under natural conditions.
doi:10.1093/aob/mcm059
PMCID: PMC3243577  PMID: 17553825
Bumblebee; computer graphics; elliptic Fourier descriptors; flower corolla shape; Primula sieboldii; principal components analysis; visual cue
20.  Determining the Contribution of Epidermal Cell Shape to Petal Wettability Using Isogenic Antirrhinum Lines 
PLoS ONE  2011;6(3):e17576.
The petal epidermis acts not only as a barrier to the outside world but also as a point of interaction between the flower and potential pollinators. The presence of conical petal epidermal cells has previously been shown to influence the attractiveness of the flower to pollinating insects. Using Antirrhinum isogenic lines differing only in the presence of a single epidermal structure, conical cells, we were able to investigate how the structure of the epidermis influences petal wettability by measuring the surface contact angle of water drops. Conical cells have a significant impact on how water is retained on the flower surface, which may have indirect consequences for pollinator behaviour. We discuss how the petal epidermis is a highly multifunctional one and how a battery of methods, including the use of isogenic lines, is required to untangle the impacts of specific epidermal properties in an ecological context.
doi:10.1371/journal.pone.0017576
PMCID: PMC3053357  PMID: 21423738
21.  Antiobesity and hypolipidemic effects of lotus leaf hot water extract with taurine supplementation in rats fed a high fat diet 
Journal of Biomedical Science  2010;17(Suppl 1):S42.
Background
Lotus (Nelumbo nucifera) leaf has been used to treat obesity. The purpose of this study was to investigate the antiobesity and hypolipidemic effects of lotus leaf hot water extract with taurine supplementation in high fat diet-induced obese rats.
Methods
Four week-old male Sprague-Dawley rats were randomly divided into four groups with 8 rats in each group for a period of 6 weeks (normal diet, N group; high fat diet, HF group; high fat diet + lotus leaf hot water extract, HFL group; high fat diet + lotus leaf hot water extract + taurine, HFLT group). Lotus leaf hot water extract was orally administrated to HFL and HFLT groups and the same amount of distilled water was orally administered (400 mg/kg/day) to N and HF groups. Taurine was supplemented by dissolving in feed water (3% w/v).
Results
The body weight gain and relative weights of epididymal and retroperitoneal adipose tissues were significantly lower in N, HFL and HFLT groups compared to HF group. HFL and HFLT groups showed lower concentrations of total cholesterol, triglyceride and low density lipoprotein cholesterol in serum. HFLT group showed higher the ratio of high density lipoprotein cholesterol/total cholesterol compared to HFL group. HFLT group showed better blood lipid profiles compared to HFL group.
Conclusions
Lotus leaf hot water extract with taurine supplementation showed antiobesity and hypolipidemic effects in high fat diet-induced obese rats, which was more effective than lotus leaf hot water extract alone.
doi:10.1186/1423-0127-17-S1-S42
PMCID: PMC2994410  PMID: 20804619
22.  Transcriptome sequencing of a chimaera reveals coordinated expression of anthocyanin biosynthetic genes mediating yellow formation in herbaceous peony (Paeonia lactiflora Pall.) 
BMC Genomics  2014;15(1):689.
Background
Herbaceous peony (Paeonia lactiflora Pall.) is a traditional flower in China and a wedding attractive flower in worldwide. In its flower colour, yellow is the rarest which is ten times the price of the other colours. However, the breeding of new yellow P. lactiflora varieties using genetic engineering is severely limited due to the little-known biochemical and molecular mechanisms underlying its characteristic formation.
Results
In this study, two cDNA libraries generated from P. lactiflora chimaera with red outer-petal and yellow inner-petal were sequenced using an Illumina HiSeq™ 2000 platform. 66,179,398 and 65,481,444 total raw reads from red outer-petal and yellow inner-petal cDNA libraries were generated, which were assembled into 61,431 and 70,359 Unigenes with an average length of 628 and 617 nt, respectively. Moreover, 61,408 non-redundant All-unigenes were obtained, with 37,511 All-unigenes (61.08%) annotated in public databases. In addition, 6,345 All-unigenes were differentially expressed between the red outer-petal and yellow inner-petal, with 3,899 up-regulated and 2,446 down-regulated All-unigenes, and the flavonoid metabolic pathway related to colour development was identified using the Kyoto encyclopedia of genes and genomes database (KEGG). Subsequently, the expression patterns of 10 candidate differentially expressed genes (DEGs) involved in the flavonoid metabolic pathway were examined, and flavonoids were qualitatively and quantitatively analysed. Numerous anthoxanthins (flavone and flavonol) and a few anthocyanins were detected in the yellow inner-petal, which were all lower than those in the red outer-petal due to the low expression levels of the phenylalanine ammonialyase gene (PlPAL), flavonol synthase gene (PlFLS), dihydroflavonol 4-reductase gene (PlDFR), anthocyanidin synthase gene (PlANS), anthocyanidin 3-O-glucosyltransferase gene (Pl3GT) and anthocyanidin 5-O-glucosyltransferase gene (Pl5GT).
Conclusion
Transcriptome sequencing (RNA-Seq) analysis based on the high throughput sequencing technology was an efficient approach to identify critical genes in P. lactiflora and other non-model plants. The flavonoid metabolic pathway and glucide metabolic pathway were identified as relatived yellow formation in P. lactiflora, PlPAL, PlFLS, PlDFR, PlANS, Pl3GT and Pl5GT were selected as potential candidates involved in flavonoid metabolic pathway, which inducing inhibition of anthocyanin biosynthesis mediated yellow formation in P. lactiflora. This study could lay a theoretical foundation for breeding new yellow P. lactiflora varieties.
Electronic supplementary material
The online version of this article (doi:10.1186/1471-2164-15-689) contains supplementary material, which is available to authorized users.
doi:10.1186/1471-2164-15-689
PMCID: PMC4159507  PMID: 25134523
Anthocyanins; Flower colour; Flavonoids; Herbaceous peony; Transcriptome; Yellow
23.  A double-flowered variety of lesser periwinkle (Vinca minor fl. pl.) that has persisted in the wild for more than 160 years 
Annals of Botany  2011;107(9):1445-1452.
Background and Aims
Homeotic transitions are usually dismissed by population geneticists as credible modes of evolution due to their assumed negative impact on fitness. However, several lines of evidence suggest that such changes in organ identity have played an important role during the origin and subsequent evolution of the angiosperm flower. Better understanding of the performance of wild populations of floral homeotic varieties should help to clarify the evolutionary potential of homeotic mutants. Wild populations of plants with changes in floral symmetry, or with reproductive organs replacing perianth organs or sepals replacing petals have already been documented. However, although double-flowered varieties are quite popular as ornamental and garden plants, they are rarely found in the wild and, if they are, usually occur only as rare mutant individuals, probably because of their low fitness relative to the wild-type. We therefore investigated a double-flowered variety of lesser periwinkle, Vinca minor flore pleno (fl. pl.), that is reported to have existed in the wild for at least 160 years. To assess the merits of this plant as a new model system for investigations on the evolutionary potential of double-flowered varieties we explored the morphological details and distribution of the mutant phenotype.
Methods
The floral morphology of the double-flowered variety and of a nearby population of wild-type plants was investigated by means of visual inspection and light microscopy of flowers, the latter involving dissected or sectioned floral organs.
Key Results
The double-flowered variety was found in several patches covering dozens of square metres in a forest within the city limits of Jena (Germany). It appears to produce fewer flowers than the wild-type, and its flowers are purple rather than blue. Most sepals in the first floral whorl resemble those in the wild-type, although occasionally one sepal is broadened and twisted. The structure of second-whorl petals is very similar to that of the wild-type, but their number per flower is more variable. The double-flowered character is due to partial or complete transformation of stamens in the third whorl into petaloid organs. Occasionally, ‘flowers within flowers’ also develop on elongated pedicels in the double-flowered variety.
Conclusions
The flowers of V. minor fl. pl. show meristic as well as homeotic changes, and occasionally other developmental abnormalities such as mis-shaped sepals or loss of floral determinacy. V. minor fl. pl. thus adds to a growing list of natural floral homeotic varieties that have established persistent populations in the wild. Our case study documents that even mutant varieties that have reproductive organs partially transformed into perianth organs can persist in the wild for centuries. This finding makes it at least conceivable that even double-flowered varieties have the potential to establish new evolutionary lineages, and hence may contribute to macroevolutionary transitions and cladogenesis.
doi:10.1093/aob/mcr090
PMCID: PMC3108809  PMID: 21527418
Double-flowered variety; homeosis; lesser periwinkle; macroevolution; Vinca minor fl. pl
24.  'Who's who' in two different flower types of Calluna vulgaris (Ericaceae): morphological and molecular analyses of flower organ identity 
BMC Plant Biology  2009;9:148.
Background
The ornamental crop Calluna vulgaris is of increasing importance to the horticultural industry in the northern hemisphere due to a flower organ mutation: the flowers of the 'bud-flowering' phenotype remain closed i.e. as buds throughout the total flowering period and thereby maintain more colorful flowers for a longer period of time than the wild-type. This feature is accompanied and presumably caused by the complete lack of stamens. Descriptions of this botanical particularity are inconsistent and partially conflicting. In order to clarify basic questions of flower organ identity in general and stamen loss in detail, a study of the wild-type and the 'bud-flowering' flower type of C. vulgaris was initiated.
Results
Flowers were examined by macro- and microscopic techniques. Organ development was investigated comparatively in both the wild-type and the 'bud-flowering' type by histological analyses. Analysis of epidermal cell surface structure of vegetative tissues and perianth organs using scanning electron microscopy revealed that in wild-type flowers the outer whorls of colored organs may be identified as sepals, while the inner ones may be identified as petals. In the 'bud-flowering' type, two whorls of sepals are directly followed by the gynoecium. Both, petals and stamens, are completely missing in this flower type. The uppermost whorl of green leaves represents bracts in both flower types.
In addition, two MADS-box genes (homologs of AP3/DEF and SEP1/2) were identified in C. vulgaris using RACE-PCR. Expression analysis by qRT-PCR was conducted for both genes in leaves, bracts, sepals and petals. These experiments revealed an expression pattern supporting the organ classification based on morphological characteristics.
Conclusions
Organ identity in both wild-type and 'bud-flowering' C. vulgaris was clarified using a combination of microscopic and molecular methods. Our results for bract, sepal and petal organ identity are supported by the 'ABCDE model'. However, loss of stamens in the 'bud-flowering' phenotype is an exceptional flower organ modification that cannot be explained by modified spatial expression of known organ identity genes.
doi:10.1186/1471-2229-9-148
PMCID: PMC2803492  PMID: 20003430
25.  Physiological temperature regulation by flowers of the sacred lotus 
Flowers of the sacred lotus, Nelumbo nucifera Gaertn. (Nelumbonaceae) are thermogenic and physiologically thermoregulatory. The 42 g flowers remain between 30-36°C during a 2 to 4-day period despite fluctuations in environmental temperatures between about 10-45°C. As the ambient temperature drops, the flowers increase heat production in proportion. Temperature regulation apparently occurs at a cellular level, by a steep, reversible thermal inhibition of respiration at flower temperatures above 30°C. There was a marked time lag between change in flower temperature and compensatory response, suggesting regulation through a biochemical feedback mechanism rather than structural changes in enzymes or membranes. By oxidizing carbohydrate, the flowers produce up to 1 W, with about half of the heat coming from the 8.5 g carpellary receptacle. The period of temperature regulation begins before petal opening and continues through the period of stigma receptivity. Temperature regulation may reward insect pollinators with a warm, equable environment, or it possibly enhances and coordinates flower development.
doi:10.1098/rstb.1998.0258
PMCID: PMC1692294

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