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Arthropod adhesion
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<h2 id="smooth-adhesive-pads-for-locomotion">Smooth adhesive pads for locomotion</h2> <p>Smooth adhesion has evolved in many families of organisms independently, which creates structures that appear unrelated to each other but generate the same function.<a class="footnote-ref" id="fnref:32" href="#fn:32"><sup>32</sup></a> <a href="/facts/Phylogenetic/bw4rsovt">Phylogenetic</a> analyses indicate that adhesive structures of arthropods evolved several times.<a class="footnote-ref" id="fnref:33" href="#fn:33"><sup>33</sup></a> Organisms such as ants, bees, cockroaches and grasshoppers use smooth adhesive pads.<a class="footnote-ref" id="fnref:34" href="#fn:34"><sup>34</sup></a> There are different types of smooth adhesive pads in these organisms such as the arolia, pulvilli, and euplantulae, all of which have a cuticle that is extremely soft and deformable.<a class="footnote-ref" id="fnref:35" href="#fn:35"><sup>35</sup></a><a class="footnote-ref" id="fnref:36" href="#fn:36"><sup>36</sup></a> The arolia of some ant species has been observed to be fluid-filled and is extended and contracted to provide adhesive force.<a class="footnote-ref" id="fnref:37" href="#fn:37"><sup>37</sup></a> The euplantulae in crickets have a <a href="/facts/Hexagonal/DPzuCzWE">hexagonal</a> microstructure which is similar to toe pads in tree frogs.<a class="footnote-ref" id="fnref:38" href="#fn:38"><sup>38</sup></a> Generally, insects are able to adhere to surfaces through the contact between the insect adhesive organs and substrates that are mediated by nanometre-thin films of adhesive fluid.<a class="footnote-ref" id="fnref:39" href="#fn:39"><sup>39</sup></a> Some functional principles of smooth pads (adaptability, <a href="/facts/Viscoelasticity/pDhHgx2T">viscoelasticity</a>, pressure sensitivity) are similar to those known from industrial pressure-sensitive adhesion.<a class="footnote-ref" id="fnref:40" href="#fn:40"><sup>40</sup></a> Smooth adhesive organs are ‘‘pillowlike,’’ which refers to the soft and fluid-filled, cuticular sac that moulds to the surface increasing the contact area on rough surfaces.<a class="footnote-ref" id="fnref:41" href="#fn:41"><sup>41</sup></a> It appears that the fluid in smooth adhesive systems mainly serves to maximize contact on rough substrates.<a class="footnote-ref" id="fnref:42" href="#fn:42"><sup>42</sup></a> The internal fibrous structure of smooth pads might be vital to their ability to deform, for shear-induced lateral increase in contact area, or for efficient transfer of tensile forces, yet at this point its specific function is unknown.<a class="footnote-ref" id="fnref:43" href="#fn:43"><sup>43</sup></a> </p> <h2 id="hairy-adhesive-pads-for-locomotion">Hairy adhesive pads for locomotion</h2> <p>Both hairy and smooth pads in arthropods act to maximize the amount of contact with a surface.<a class="footnote-ref" id="fnref:44" href="#fn:44"><sup>44</sup></a> The foot pads of flies are densely covered with flexible hair-like structures called <a href="/facts/Setae/jUmts6C2">setae</a>, and some <a href="/facts/Lizards/VPCpCwjT">lizards</a> and <a href="/facts/Spiders/aqpU7eg3">spiders</a> use similar hairy pads to create adhesive effects.<a class="footnote-ref" id="fnref:45" href="#fn:45"><sup>45</sup></a> This indicates a favourable design for hairy pad adhesion.<a class="footnote-ref" id="fnref:46" href="#fn:46"><sup>46</sup></a> Hairy pads can be classified as pulvillus, fossula spongiosa, and tenent hairs.<a class="footnote-ref" id="fnref:47" href="#fn:47"><sup>47</sup></a> Hairy attachment pads employed few other features, such as flaw tolerance, lower sensitivity to contamination, and roughness.<a class="footnote-ref" id="fnref:48" href="#fn:48"><sup>48</sup></a> Hairy attachment systems are typical for evolutionary younger and successful insect groups, such as <a href="/facts/Beetle/hv33fxsL">Coleoptera</a> and <a href="/facts/Fly/ugzGBFKG">Diptera</a>.<a class="footnote-ref" id="fnref:49" href="#fn:49"><sup>49</sup></a> The density of hairs increases with increasing body weight.<a class="footnote-ref" id="fnref:50" href="#fn:50"><sup>50</sup></a> An increase of the attachment strength in hairy systems is realized by increasing the number of single contact points.<a class="footnote-ref" id="fnref:51" href="#fn:51"><sup>51</sup></a> Protuberances on the hairy pads of Coleoptera, <a href="/facts/Earwig/JHSLqPEB">Dermaptera</a>, and Diptera belong to different types. Representatives of the first two lineages have socketed setae on their pads.<a class="footnote-ref" id="fnref:52" href="#fn:52"><sup>52</sup></a> Setae can range in length from a few micrometers to several millimeters.<a class="footnote-ref" id="fnref:53" href="#fn:53"><sup>53</sup></a> <a href="/facts/Dipteran/ugzGBFKG">Dipteran</a> outgrowths are acanthae, which are single <a href="/facts/Sclerite/KWTShcvT">sclerotized</a> protuberances originating from a single cell.<a class="footnote-ref" id="fnref:54" href="#fn:54"><sup>54</sup></a> The acanthae are hollow inside, and some have pores under the terminal plate, which presumably deliver an adhesive secretion directly to the contact area.<a class="footnote-ref" id="fnref:55" href="#fn:55"><sup>55</sup></a> Hairy attachment pads of reduviid bugs,<a class="footnote-ref" id="fnref:56" href="#fn:56"><sup>56</sup></a> flies <a class="footnote-ref" id="fnref:57" href="#fn:57"><sup>57</sup></a> and <a href="/facts/Beetles/hv33fxsL">beetles</a><a class="footnote-ref" id="fnref:58" href="#fn:58"><sup>58</sup></a> secrete fluid into the contact area. The secretion contains non-volatile, lipid-like substances, but in some species it is two-phasic emulsion presumably containing water-soluble and lipid-soluble fractions.<a class="footnote-ref" id="fnref:59" href="#fn:59"><sup>59</sup></a> Adhesion strongly decreases as the volume of the secretion decreases, which indicates that a layer of pad secretion that covers the terminal plates is crucial for generating a strong attractive force.<a class="footnote-ref" id="fnref:60" href="#fn:60"><sup>60</sup></a> Data suggests that besides van der Waals and <a href="/facts/Coulomb_force/V0lR5NKv">Coulomb forces</a> flies rely on attractive capillary forces mediated by the pad secretion.<a class="footnote-ref" id="fnref:61" href="#fn:61"><sup>61</sup></a> At low humidity, adhesion strongly depends on the amount of liquid deposited on the surface, and therefore contact duration.<a class="footnote-ref" id="fnref:62" href="#fn:62"><sup>62</sup></a> </p><p>The hairy pad design has been argued to have a number of advantages over the smooth design, such as superior performance on rough substrates, effortless detachment, self-cleaning properties, and increased adhesion due to contact splitting.<a class="footnote-ref" id="fnref:63" href="#fn:63"><sup>63</sup></a> </p><p>Unlike frog and lizard adhesive pads which are often dry, insects tend to have an associated fluid for adhesion. The fluid that is secreted has a special property of being composed of an <a href="/facts/Miscibility/HwQaolbQ">immiscible</a> mixture of <a href="/facts/Hydrophile/k8p95WYu">hydrophilic</a> and <a href="/facts/Hydrophobe/Ifya0CXQ">hydrophobic</a> material.<a class="footnote-ref" id="fnref:64" href="#fn:64"><sup>64</sup></a> </p><p>Adhesive <a href="/facts/Foot/gnX2RTz7">foot</a> pads only stick when pulled toward the body, but unstick when moved away from it, which allows for effortless and rapid detachment. Insects can do this actively through their claw flexor muscle, but in most cases, the foot is able to attach and detach passively, without the help of any nerves and muscles. (Bullock, Drechsler, & Federle, 2008) </p> <h2 id="other-uses-for-adhesives-in-insects">Other uses for adhesives in insects</h2> <p>Adhesive chemical secretions are also used for predation defence, mating, holding substrates, anchoring eggs, building retreats, prey capture, and self grooming. Structures for use in repelling attackers or temporarily or permanently adhering to a substratum or a mating partner have been found in the developmental stages of the egg, larvae, pupae, and adult. Some species have developed adhesives for prey capture and some use adhesive glue for cocoon building. Adhesive glands of the head can involve mouthparts, antennae, the labial salivary glands, or species specific glands. A variety of glands, often located in the abdomen, can be used for defensive adhesion mechanisms.<a class="footnote-ref" id="fnref:65" href="#fn:65"><sup>65</sup></a> </p> <h2 id="insect-adhesive-glands">Insect adhesive glands</h2> <p><a href="/facts/Epidermal/JvRABubP">Epidermal</a> <a href="/facts/Glands/gfq2of7k">glands</a> and their secretions are highly diverse and vary in their function for: protection from adverse environmental conditions and <a href="/facts/Microbial/pysJYcE2">microbial</a> <a href="/facts/Contamination/RC0L7KRy">contamination</a>, regulation of <a href="/facts/Water/0lkXnJPK">water</a> balance, communication with <a href="/facts/Pheromone/ymEDICw3">pheromones</a> and alelochemicals, defense from <a href="/facts/Predator/JEGdrX3f">predators</a> and <a href="/facts/Parasites/HWPcgky7">parasites</a>, construction and making food accessible.<a class="footnote-ref" id="fnref:66" href="#fn:66"><sup>66</sup></a> </p><p>Class 1 epidermal cells are the predominant glandular <a href="/facts/Cell_(biology)/bLM9UQvy">cell</a> type for adhesive gland systems in insects with features that indicate either <a href="/facts/Lipid/TdQR87HX">lipid</a> or <a href="/facts/Protein/Jxmagu3E">protein</a> secretion. In class 1 cells for locomotion lipoidal secretion is most common, although the secretions are often mixtures of lipids with proteins and <a href="/facts/Carbohydrates/Geuuzxsf">carbohydrates</a>. Class 1 cells that are used for more permanent body or egg anchorage and for retreat building make use of protein-based secretions.<a class="footnote-ref" id="fnref:67" href="#fn:67"><sup>67</sup></a> </p><p>Class 2 epidermal adhesive gland cells have only been found in the defence systems of <a href="/facts/Aphid/kbvClSLl">Aphidoidea</a> and <a href="/facts/Tingidae/s9qLcKn2">Tingidae</a>. Defensive adhesive secretions function mechanically and also develop a chemical <a href="/facts/Irritation/vGtMyXUR">irritant</a> function caused by reactive substances of low <a href="/facts/Molecular/N9ljSfFq">molecular</a> weight which combines within the sticky secretion to produce <a href="/facts/Toxic/qa96rs60">toxic</a> glue.<a class="footnote-ref" id="fnref:68" href="#fn:68"><sup>68</sup></a> </p><p>Class 3 epidermal adhesive glands are usually bicellular and consist of a terminal secretarily active cell and an adjacent canal cell that surrounds the cuticular conducting duct.<a class="footnote-ref" id="fnref:69" href="#fn:69"><sup>69</sup></a> </p><p>Hundreds of gland cells and glandular units are contained in class 1 or 3 and might aggregate to form whole gland organs so as to discharge large amounts of a secretion. Adhesive cells used for locomotion are all class 1 epidermal adhesive cells. Class 3 epidermal adhesive cells may play a role in some hairy adhesive pads, but this has not yet been confirmed. Some adhesive glands that are used for locomotion are also used for capturing or holding on to prey (Fac, 2010). The secretion of some class 1 cells and class 3 cells are mixed in the subcuticular or intracuticular spaces. They may also be mixed in the larger glandular reservoirs before being discharged, which allows the formation of complex structural mixtures as well as chemical reactions between the components of the mixture. Gland cells used by female insects for gluing eggs to a substratum during oviposition have not been well studied. Glands used for sticking eggs to surfaces have been observed to be of the class 1 type. Adhesive glands are involved in the production of silks, which are produced by a variety of dermal glands for building shelters, cocoons, and supporting sperm.<a class="footnote-ref" id="fnref:70" href="#fn:70"><sup>70</sup></a> Class 1 cells are often applied for this purpose. </p> <h2 id="adhesive-secretions">Adhesive secretions</h2> <p>Most bioadhesives use <a href="/facts/Polymers/eF32E50K">polymers</a> (carbohydrates and proteins) to create the adhesive and cohesive strength.<a class="footnote-ref" id="fnref:71" href="#fn:71"><sup>71</sup></a> Natural adhesives used by both plants and animals are composed of only a few basic components, such as proteins, <a href="/facts/Polysaccharides/lqXq2cJU">polysaccharides</a>, <a href="/facts/Polyphenols/H6Q7QIQH">polyphenols</a>, and lipids that are mixed in various combinations.<a class="footnote-ref" id="fnref:72" href="#fn:72"><sup>72</sup></a> Natural adhesive chemical and micromechanical functions are often not well understood.<a class="footnote-ref" id="fnref:73" href="#fn:73"><sup>73</sup></a> Adhesives that are for mechanical work are often composed of high-molecular compounds containing proteins, <a href="/facts/Resins/r1uVThQ8">resins</a>, mixtures of long-chain hydrocarbons and mucopolysaccharides, or waxes.<a class="footnote-ref" id="fnref:74" href="#fn:74"><sup>74</sup></a> Defensive adhesive secretions often combine their mechanical effect with a low molecular weight chemical irritant to deter predators.<a class="footnote-ref" id="fnref:75" href="#fn:75"><sup>75</sup></a> </p><p>There is a great diversity of <a href="/facts/Aliphatic/9usN3neo">aliphatic</a> compounds in insect adhesive secretions. Aliphatic compounds are a major constituent of secretions for some locomotion organs in insects, and they are also involved in the formation of defence secretions. Adhesives of this type contain only limited amounts or no polar components such as fatty acids, esters, and alcohols. Often these compounds are temperature sensitive.<a class="footnote-ref" id="fnref:76" href="#fn:76"><sup>76</sup></a> Very little research has been done on classifying and identifying <a href="/facts/Carbohydrate/Geuuzxsf">carbohydrates</a> within insect adhesive secretions. So far, <a href="/facts/Glucose/oUSkoLCD">glucose</a>, <a href="/facts/Trehalose/1N3XTfxV">trehalose</a> and <a href="/facts/Mucopolysaccharide/fo22ktAQ">mucopolysaccharides</a> that contain glucose, <a href="/facts/Galactose/sh2hEFpC">galactose</a>, <a href="/facts/Mannose/U05bJ7p8">mannose</a>, beta-<a href="/facts/Glucopyranose/oUSkoLCD">glucopyranose</a>, and/or (N-acetyl-beta-) <a href="/facts/Glucosamine/HKCp9hoR">glucosamine</a> have been identified as components of insect adhesives. Carbohydrates have been found in defense secretions as well as for sticking eggs together.<a class="footnote-ref" id="fnref:77" href="#fn:77"><sup>77</sup></a> <a href="/facts/Aromatic_compound/3EXq8Tgf">Aromatic compounds</a> have been identified in the adhesive defence secretion of <a href="/facts/Termites/KRFbDumW">termites</a> and ants. It is also thought to be used by <a href="/facts/Butterflies/qlSgwlRx">butterflies</a> to secure eggs.<a class="footnote-ref" id="fnref:78" href="#fn:78"><sup>78</sup></a> Insect adhesives contain a broad spectrum of <a href="/facts/Isoprenoid/1GvG7W7v">isoprenoids</a>. These compounds have been found in defense mechanisms in some species such as termites.<a class="footnote-ref" id="fnref:79" href="#fn:79"><sup>79</sup></a> <a href="/facts/Amino_acid/WDxYwBny">Amino acids</a>, peptides, and proteins are nearly always found in insects' adhesive secretions. They are employed for adhesion across many functions such as defense, locomotion and cocoon building.<a class="footnote-ref" id="fnref:80" href="#fn:80"><sup>80</sup></a> </p> <h2 id="arachnid-adhesion">Arachnid adhesion</h2> <p>Spiders have independently evolved hairy adhesive pads. Their pads do not use an associated fluid and are much similar to many lizards, not like the hairy pads that are used by insects.<a class="footnote-ref" id="fnref:81" href="#fn:81"><sup>81</sup></a> </p> <h2 id="convergence-with-tetrapods">Convergence with tetrapods</h2> <p>Smooth adhesive pads are an example of <a href="/facts/Convergent_evolution/dd0L8z4R">convergent evolution</a> between <a href="/facts/Amphibia/r2BleD2W">amphibians</a> (<a href="/facts/Gecko/13Hu5c1w">geckos</a> and frogs), arthropods, and mammals (<a href="/facts/Phalangeriformes/76UruwPo">possum</a>).<a class="footnote-ref" id="fnref:82" href="#fn:82"><sup>82</sup></a> The mechanisms involved even appear to be similar.<a class="footnote-ref" id="fnref:83" href="#fn:83"><sup>83</sup></a> This could indicate that this method of locomotion has found its optimal form in many species of animals.<a class="footnote-ref" id="fnref:84" href="#fn:84"><sup>84</sup></a> Hairy attachment systems of the gekkonid lizards and spiders do not produce fluids; these organisms rely on <a href="/facts/Van_der_Waals_interactions/jyGU8Vh9">van der Waals interactions</a> for the generation of strong attractive forces.<a class="footnote-ref" id="fnref:85" href="#fn:85"><sup>85</sup></a><a class="footnote-ref" id="fnref:86" href="#fn:86"><sup>86</sup></a> <a href="/facts/Tree_frog/D7LhXSTX">Tree frog</a> toe-pads are made of columnar epithelial cells that are separated from each other at the apices.<a class="footnote-ref" id="fnref:87" href="#fn:87"><sup>87</sup></a> Pores for mucous glands open into the channels that are between the cells which create a toe pad epithelium that has an array of flat topped cells with mucus-filled grooves between them.<a class="footnote-ref" id="fnref:88" href="#fn:88"><sup>88</sup></a> The purpose of having cells separated at the tip is to allow the toe to conform to the structure it will adhere to.<a class="footnote-ref" id="fnref:89" href="#fn:89"><sup>89</sup></a> The hexagonal design around the outside of the cells (similar to the crickets) is likely to allow for the mucus to spread evenly over the cell.<a class="footnote-ref" id="fnref:90" href="#fn:90"><sup>90</sup></a> Smooth adhesive pads are found in arboreal possums, which are <a href="/facts/Marsupials/zN5s4bMq">marsupials</a> that glide between trees.<a class="footnote-ref" id="fnref:91" href="#fn:91"><sup>91</sup></a> The possum is also capable of using smooth adhesive pads to climb vertically, making use of large toe pads.<a class="footnote-ref" id="fnref:92" href="#fn:92"><sup>92</sup></a> The pads consist of an epidermal layer of <a href="/facts/Stratified_squamous_epithelium/RLlp5jnr">stratified squamous epithelium</a>, with the outer most layer's cells being flattened.<a class="footnote-ref" id="fnref:93" href="#fn:93"><sup>93</sup></a> The pad has alternating ridges and grooves with <a href="/facts/Sweat_glands/T9nLT1V4">sweat glands</a> emptying into the grooves providing fluid for wet adhesion.<a class="footnote-ref" id="fnref:94" href="#fn:94"><sup>94</sup></a> <a href="/facts/Bat/CJudzNt1">Bats</a> have also evolved adhesive pads separately. Some bats make use of an adhesive appendage, while others have <a href="/facts/Suction/IATXs4b3">suctioning</a> adhesive organs.<a class="footnote-ref" id="fnref:95" href="#fn:95"><sup>95</sup></a> </p> <h2 id="importance-to-humans">Importance to humans</h2> <p>Some researchers propose using the advanced locomotive mechanisms seen in arthropods for modelling robotic movement to create maximally efficient movement.<a class="footnote-ref" id="fnref:96" href="#fn:96"><sup>96</sup></a><a class="footnote-ref" id="fnref:97" href="#fn:97"><sup>97</sup></a><a class="footnote-ref" id="fnref:98" href="#fn:98"><sup>98</sup></a> Currently insect adhesive pads still outperform most artificial adhesives with respect to rapid controllability.<a class="footnote-ref" id="fnref:99" href="#fn:99"><sup>99</sup></a> Some researchers also suggest using arthropod-based adhesive mechanisms for more effective tape and binding tools.<a class="footnote-ref" id="fnref:100" href="#fn:100"><sup>100</sup></a><a class="footnote-ref" id="fnref:101" href="#fn:101"><sup>101</sup></a> Additionally, some research indicates that the wrinkling effect that occurs in human fingers when submerged in water acts to increase grip on wet objects.<a class="footnote-ref" id="fnref:102" href="#fn:102"><sup>102</sup></a> The mechanism is unknown but it may be due to changes in adhesion properties of the finger pads. By examining the properties of bioadhesion, finger pad adhesion can be better understood. However, this study on increased finger pad dexterity from wrinkling has been heavily disputed.<a class="footnote-ref" id="fnref:103" href="#fn:103"><sup>103</sup></a> Despite this, it can be argued that a better understanding of insect adhesion mechanisms can help guide the development of better adhesives for human mobility and technology, as well as inform a better understanding of human finger function. </p> <h2 id="see-also">See also</h2> <ul><li><a href="/facts/Gecko_feet/DSykeLLM">Gecko feet</a></li> <li><a href="/facts/Synthetic_setae/7eGQ0sBo">Synthetic setae</a></li></ul> <h2 id="references">References</h2> <ol> <li id="fn:1"><p>Barnes, W. Jon. P. (2011). "Functional Morphology and Design Constraints of Smooth Adhesive Pads". MRS Bulletin. 32 (6): 479–485. doi:10.1557/mrs2007.81. <a href="/wiki/Doi_(identifier)" target="_blank">/wiki/Doi_(identifier)</a> <a href="#fnref:1" class="footnote-back-ref">↩</a></p></li> <li id="fn:2"><p>Dirks, Jan-Henning; Federle, Walter (2011). "Fluid-based adhesion in insects – principles and challenges". Soft Matter. 7 (23): 11047. Bibcode:2011SMat....711047D. doi:10.1039/C1SM06269G. <a href="/wiki/Bibcode_(identifier)" target="_blank">/wiki/Bibcode_(identifier)</a> <a href="#fnref:2" class="footnote-back-ref">↩</a></p></li> <li id="fn:3"><p>Bullock, J. M.; Drechsler, P.; Federle, W. (2008). "Comparison of smooth and hairy attachment pads in insects: friction, adhesion and mechanisms for direction-dependence". The Journal of Experimental Biology. 211 (20): 3333–3343. doi:10.1242/jeb.020941. PMID 18840668. <a href="https://doi.org/10.1242%2Fjeb.020941" target="_blank">https://doi.org/10.1242%2Fjeb.020941</a> <a href="#fnref:3" class="footnote-back-ref">↩</a></p></li> <li id="fn:4"><p>Bullock, J. M.; Drechsler, P.; Federle, W. (2008). "Comparison of smooth and hairy attachment pads in insects: friction, adhesion and mechanisms for direction-dependence". The Journal of Experimental Biology. 211 (20): 3333–3343. doi:10.1242/jeb.020941. PMID 18840668. <a href="https://doi.org/10.1242%2Fjeb.020941" target="_blank">https://doi.org/10.1242%2Fjeb.020941</a> <a href="#fnref:4" class="footnote-back-ref">↩</a></p></li> <li id="fn:5"><p>von Byern, Janek; Grunwald, Ingo, eds. (2010). Biological Adhesive Systems: From Nature to Technical and Medical Application (1st ed.). Wien: Springer. ISBN 978-3-7091-0141-4. <a href="978-3-7091-0141-4" target="_blank">978-3-7091-0141-4</a> <a href="#fnref:5" class="footnote-back-ref">↩</a></p></li> <li id="fn:6"><p>Zhou, Y.; Robinson, A.; Steiner, U.; Federle, W. (2014). "Insect adhesion on rough surfaces: analysis of adhesive contact of smooth and hairy pads on transparent microstructured substrates". J. R. Soc. Interface. 11 (98): 20140499. doi:10.1098/rsif.2014.0499. PMC 4233698. PMID 24990289. <a href="http://classic.rsif.royalsocietypublishing.org/content/11/98/20140499.full" target="_blank">http://classic.rsif.royalsocietypublishing.org/content/11/98/20140499.full</a> <a href="#fnref:6" class="footnote-back-ref">↩</a></p></li> <li id="fn:7"><p>von Byern, Janek; Grunwald, Ingo, eds. (2010). Biological Adhesive Systems: From Nature to Technical and Medical Application (1st ed.). Wien: Springer. ISBN 978-3-7091-0141-4. <a href="978-3-7091-0141-4" target="_blank">978-3-7091-0141-4</a> <a href="#fnref:7" class="footnote-back-ref">↩</a></p></li> <li id="fn:8"><p>Bullock, J. M.; Drechsler, P.; Federle, W. (2008). "Comparison of smooth and hairy attachment pads in insects: friction, adhesion and mechanisms for direction-dependence". The Journal of Experimental Biology. 211 (20): 3333–3343. doi:10.1242/jeb.020941. PMID 18840668. <a href="https://doi.org/10.1242%2Fjeb.020941" target="_blank">https://doi.org/10.1242%2Fjeb.020941</a> <a href="#fnref:8" class="footnote-back-ref">↩</a></p></li> <li id="fn:9"><p>Gorb, Stanislav N; Sinha, Mitali; Peressadko, Andrei; Daltorio, Kathryn A; Quinn, Roger D (2007). "Insects did it first: a micropatterned adhesive tape for robotic applications". Bioinspiration & Biomimetics. 2 (4): S117 – S125. Bibcode:2007BiBi....2..117G. doi:10.1088/1748-3182/2/4/S01. PMID 18037721. S2CID 6709383. <a href="/wiki/Bibcode_(identifier)" target="_blank">/wiki/Bibcode_(identifier)</a> <a href="#fnref:9" class="footnote-back-ref">↩</a></p></li> <li id="fn:10"><p>Bullock, J. M.; Drechsler, P.; Federle, W. (2008). "Comparison of smooth and hairy attachment pads in insects: friction, adhesion and mechanisms for direction-dependence". The Journal of Experimental Biology. 211 (20): 3333–3343. doi:10.1242/jeb.020941. PMID 18840668. <a href="https://doi.org/10.1242%2Fjeb.020941" target="_blank">https://doi.org/10.1242%2Fjeb.020941</a> <a href="#fnref:10" class="footnote-back-ref">↩</a></p></li> <li id="fn:11"><p>Barnes, W. Jon. P. (2011). "Functional Morphology and Design Constraints of Smooth Adhesive Pads". MRS Bulletin. 32 (6): 479–485. doi:10.1557/mrs2007.81. <a href="/wiki/Doi_(identifier)" target="_blank">/wiki/Doi_(identifier)</a> <a href="#fnref:11" class="footnote-back-ref">↩</a></p></li> <li id="fn:12"><p>Dirks, Jan-Henning; Federle, Walter (2011). "Fluid-based adhesion in insects – principles and challenges". Soft Matter. 7 (23): 11047. Bibcode:2011SMat....711047D. doi:10.1039/C1SM06269G. <a href="/wiki/Bibcode_(identifier)" target="_blank">/wiki/Bibcode_(identifier)</a> <a href="#fnref:12" class="footnote-back-ref">↩</a></p></li> <li id="fn:13"><p>Ritzmann, Roy E.; Quinn, Roger D.; Fischer, Martin S. (2004). "Convergent evolution and locomotion through complex terrain by insects, vertebrates and robots". Arthropod Structure & Development. 33 (3): 361–379. CiteSeerX 10.1.1.94.4692. doi:10.1016/j.asd.2004.05.001. PMID 18089044. <a href="/wiki/CiteSeerX_(identifier)" target="_blank">/wiki/CiteSeerX_(identifier)</a> <a href="#fnref:13" class="footnote-back-ref">↩</a></p></li> <li id="fn:14"><p>Dirks, Jan-Henning; Federle, Walter (2011). "Fluid-based adhesion in insects – principles and challenges". Soft Matter. 7 (23): 11047. Bibcode:2011SMat....711047D. doi:10.1039/C1SM06269G. <a href="/wiki/Bibcode_(identifier)" target="_blank">/wiki/Bibcode_(identifier)</a> <a href="#fnref:14" class="footnote-back-ref">↩</a></p></li> <li id="fn:15"><p>Dirks, Jan-Henning; Federle, Walter (2011). "Fluid-based adhesion in insects – principles and challenges". Soft Matter. 7 (23): 11047. Bibcode:2011SMat....711047D. doi:10.1039/C1SM06269G. <a href="/wiki/Bibcode_(identifier)" target="_blank">/wiki/Bibcode_(identifier)</a> <a href="#fnref:15" class="footnote-back-ref">↩</a></p></li> <li id="fn:16"><p>Bullock, J. M.; Drechsler, P.; Federle, W. (2008). "Comparison of smooth and hairy attachment pads in insects: friction, adhesion and mechanisms for direction-dependence". The Journal of Experimental Biology. 211 (20): 3333–3343. doi:10.1242/jeb.020941. PMID 18840668. <a href="https://doi.org/10.1242%2Fjeb.020941" target="_blank">https://doi.org/10.1242%2Fjeb.020941</a> <a href="#fnref:16" class="footnote-back-ref">↩</a></p></li> <li id="fn:17"><p>Ritzmann, Roy E.; Quinn, Roger D.; Fischer, Martin S. (2004). "Convergent evolution and locomotion through complex terrain by insects, vertebrates and robots". Arthropod Structure & Development. 33 (3): 361–379. CiteSeerX 10.1.1.94.4692. doi:10.1016/j.asd.2004.05.001. PMID 18089044. <a href="/wiki/CiteSeerX_(identifier)" target="_blank">/wiki/CiteSeerX_(identifier)</a> <a href="#fnref:17" class="footnote-back-ref">↩</a></p></li> <li id="fn:18"><p>Gorb, Stanislav N; Sinha, Mitali; Peressadko, Andrei; Daltorio, Kathryn A; Quinn, Roger D (2007). "Insects did it first: a micropatterned adhesive tape for robotic applications". Bioinspiration & Biomimetics. 2 (4): S117 – S125. Bibcode:2007BiBi....2..117G. doi:10.1088/1748-3182/2/4/S01. PMID 18037721. S2CID 6709383. <a href="/wiki/Bibcode_(identifier)" target="_blank">/wiki/Bibcode_(identifier)</a> <a href="#fnref:18" class="footnote-back-ref">↩</a></p></li> <li id="fn:19"><p>Gorb, Stanislav N; Sinha, Mitali; Peressadko, Andrei; Daltorio, Kathryn A; Quinn, Roger D (2007). "Insects did it first: a micropatterned adhesive tape for robotic applications". Bioinspiration & Biomimetics. 2 (4): S117 – S125. Bibcode:2007BiBi....2..117G. doi:10.1088/1748-3182/2/4/S01. PMID 18037721. S2CID 6709383. <a href="/wiki/Bibcode_(identifier)" target="_blank">/wiki/Bibcode_(identifier)</a> <a href="#fnref:19" class="footnote-back-ref">↩</a></p></li> <li id="fn:20"><p>Gorb, Stanislav N; Sinha, Mitali; Peressadko, Andrei; Daltorio, Kathryn A; Quinn, Roger D (2007). "Insects did it first: a micropatterned adhesive tape for robotic applications". Bioinspiration & Biomimetics. 2 (4): S117 – S125. Bibcode:2007BiBi....2..117G. doi:10.1088/1748-3182/2/4/S01. PMID 18037721. S2CID 6709383. <a href="/wiki/Bibcode_(identifier)" target="_blank">/wiki/Bibcode_(identifier)</a> <a href="#fnref:20" class="footnote-back-ref">↩</a></p></li> <li id="fn:21"><p>Bullock, J. M.; Drechsler, P.; Federle, W. (2008). "Comparison of smooth and hairy attachment pads in insects: friction, adhesion and mechanisms for direction-dependence". The Journal of Experimental Biology. 211 (20): 3333–3343. doi:10.1242/jeb.020941. PMID 18840668. <a href="https://doi.org/10.1242%2Fjeb.020941" target="_blank">https://doi.org/10.1242%2Fjeb.020941</a> <a href="#fnref:21" class="footnote-back-ref">↩</a></p></li> <li id="fn:22"><p>Bullock, J. M.; Drechsler, P.; Federle, W. (2008). "Comparison of smooth and hairy attachment pads in insects: friction, adhesion and mechanisms for direction-dependence". The Journal of Experimental Biology. 211 (20): 3333–3343. doi:10.1242/jeb.020941. PMID 18840668. <a href="https://doi.org/10.1242%2Fjeb.020941" target="_blank">https://doi.org/10.1242%2Fjeb.020941</a> <a href="#fnref:22" class="footnote-back-ref">↩</a></p></li> <li id="fn:23"><p>Bullock, J. M.; Drechsler, P.; Federle, W. (2008). "Comparison of smooth and hairy attachment pads in insects: friction, adhesion and mechanisms for direction-dependence". The Journal of Experimental Biology. 211 (20): 3333–3343. doi:10.1242/jeb.020941. PMID 18840668. <a href="https://doi.org/10.1242%2Fjeb.020941" target="_blank">https://doi.org/10.1242%2Fjeb.020941</a> <a href="#fnref:23" class="footnote-back-ref">↩</a></p></li> <li id="fn:24"><p>Bullock, J. M.; Drechsler, P.; Federle, W. (2008). "Comparison of smooth and hairy attachment pads in insects: friction, adhesion and mechanisms for direction-dependence". The Journal of Experimental Biology. 211 (20): 3333–3343. doi:10.1242/jeb.020941. PMID 18840668. <a href="https://doi.org/10.1242%2Fjeb.020941" target="_blank">https://doi.org/10.1242%2Fjeb.020941</a> <a href="#fnref:24" class="footnote-back-ref">↩</a></p></li> <li id="fn:25"><p>Bullock, J. M.; Drechsler, P.; Federle, W. (2008). "Comparison of smooth and hairy attachment pads in insects: friction, adhesion and mechanisms for direction-dependence". The Journal of Experimental Biology. 211 (20): 3333–3343. doi:10.1242/jeb.020941. PMID 18840668. <a href="https://doi.org/10.1242%2Fjeb.020941" target="_blank">https://doi.org/10.1242%2Fjeb.020941</a> <a href="#fnref:25" class="footnote-back-ref">↩</a></p></li> <li id="fn:26"><p>Bullock, J. M.; Drechsler, P.; Federle, W. (2008). "Comparison of smooth and hairy attachment pads in insects: friction, adhesion and mechanisms for direction-dependence". The Journal of Experimental Biology. 211 (20): 3333–3343. doi:10.1242/jeb.020941. PMID 18840668. <a href="https://doi.org/10.1242%2Fjeb.020941" target="_blank">https://doi.org/10.1242%2Fjeb.020941</a> <a href="#fnref:26" class="footnote-back-ref">↩</a></p></li> <li id="fn:27"><p>Bullock, J. M.; Drechsler, P.; Federle, W. (2008). "Comparison of smooth and hairy attachment pads in insects: friction, adhesion and mechanisms for direction-dependence". The Journal of Experimental Biology. 211 (20): 3333–3343. doi:10.1242/jeb.020941. PMID 18840668. <a href="https://doi.org/10.1242%2Fjeb.020941" target="_blank">https://doi.org/10.1242%2Fjeb.020941</a> <a href="#fnref:27" class="footnote-back-ref">↩</a></p></li> <li id="fn:28"><p>Bullock, J. M.; Drechsler, P.; Federle, W. (2008). "Comparison of smooth and hairy attachment pads in insects: friction, adhesion and mechanisms for direction-dependence". The Journal of Experimental Biology. 211 (20): 3333–3343. doi:10.1242/jeb.020941. PMID 18840668. <a href="https://doi.org/10.1242%2Fjeb.020941" target="_blank">https://doi.org/10.1242%2Fjeb.020941</a> <a href="#fnref:28" class="footnote-back-ref">↩</a></p></li> <li id="fn:29"><p>Bullock, J. M.; Drechsler, P.; Federle, W. (2008). "Comparison of smooth and hairy attachment pads in insects: friction, adhesion and mechanisms for direction-dependence". The Journal of Experimental Biology. 211 (20): 3333–3343. doi:10.1242/jeb.020941. PMID 18840668. <a href="https://doi.org/10.1242%2Fjeb.020941" target="_blank">https://doi.org/10.1242%2Fjeb.020941</a> <a href="#fnref:29" class="footnote-back-ref">↩</a></p></li> <li id="fn:30"><p>Zhou, Y.; Robinson, A.; Steiner, U.; Federle, W. (2014). "Insect adhesion on rough surfaces: analysis of adhesive contact of smooth and hairy pads on transparent microstructured substrates". J. R. Soc. Interface. 11 (98): 20140499. doi:10.1098/rsif.2014.0499. PMC 4233698. PMID 24990289. <a href="http://classic.rsif.royalsocietypublishing.org/content/11/98/20140499.full" target="_blank">http://classic.rsif.royalsocietypublishing.org/content/11/98/20140499.full</a> <a href="#fnref:30" class="footnote-back-ref">↩</a></p></li> <li id="fn:31"><p>von Byern, Janek; Grunwald, Ingo, eds. (2010). Biological Adhesive Systems: From Nature to Technical and Medical Application (1st ed.). Wien: Springer. ISBN 978-3-7091-0141-4. <a href="978-3-7091-0141-4" target="_blank">978-3-7091-0141-4</a> <a href="#fnref:31" class="footnote-back-ref">↩</a></p></li> <li id="fn:32"><p>Dirks, Jan-Henning; Federle, Walter (2011). "Fluid-based adhesion in insects – principles and challenges". Soft Matter. 7 (23): 11047. Bibcode:2011SMat....711047D. doi:10.1039/C1SM06269G. <a href="/wiki/Bibcode_(identifier)" target="_blank">/wiki/Bibcode_(identifier)</a> <a href="#fnref:32" class="footnote-back-ref">↩</a></p></li> <li id="fn:33"><p>Barnes, W. Jon. P. (2011). 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PMID 24990289. <a href="http://classic.rsif.royalsocietypublishing.org/content/11/98/20140499.full" target="_blank">http://classic.rsif.royalsocietypublishing.org/content/11/98/20140499.full</a> <a href="#fnref:43" class="footnote-back-ref">↩</a></p></li> <li id="fn:44"><p>Dirks, Jan-Henning; Federle, Walter (2011). "Fluid-based adhesion in insects – principles and challenges". Soft Matter. 7 (23): 11047. Bibcode:2011SMat....711047D. doi:10.1039/C1SM06269G. <a href="/wiki/Bibcode_(identifier)" target="_blank">/wiki/Bibcode_(identifier)</a> <a href="#fnref:44" class="footnote-back-ref">↩</a></p></li> <li id="fn:45"><p>Dirks, Jan-Henning; Federle, Walter (2011). "Fluid-based adhesion in insects – principles and challenges". Soft Matter. 7 (23): 11047. 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