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• W2078255086 abstract "Some spiders use glues while others deploy strands of fine filaments for fixing flies. Recent work has provided new insights into the mechanical properties of these nano-scale ropes. Some spiders use glues while others deploy strands of fine filaments for fixing flies. Recent work has provided new insights into the mechanical properties of these nano-scale ropes. A spider's web, typically in combination with a powerful poison [1Foelix R. Biology of Spiders. Oxford University Press, Oxford1996Google Scholar], is its principal tool in the struggle for survival. It is a highly tuned, light-weight net-structure that relies heavily on skilful engineering and the fitting deployment of a most versatile material: the spider's famous silk. A typical web spider makes up to seven different types of silk, some of which can be tuned to suit environmental circumstances by adjustment of the spider's spinning physiology [2Vollrath F. Knight D.P. Liquid crystal silk spinning in nature.Nature. 2001; 410: 541-548Crossref PubMed Scopus (1220) Google Scholar]. Individual silk threads, each around a micron in diameter, are actively drawn from microscopic spinning spigots perched on motile spinneret mounds, resembling miniature batteries of battle-ship gun-turrets. Some spiders — the Cribellata — have an extra structure, called a cribellum, which is a broad plate set firmly in the spider's abdominal cuticula, just in front of the spinnerets and covered with thousands of minuscule spigots [1Foelix R. Biology of Spiders. Oxford University Press, Oxford1996Google Scholar]. The performance of capture threads that incorporate the nanoscale silk produced from this plate is the focus of a recent study [3Blackledge T. Hayashi C. Unravelling the mechanical properties of composite silk threads spun by cribellate orb-weaving spiders.J. Exp. Biol. 2006; 209: 3131-3140Crossref PubMed Scopus (74) Google Scholar]. The cribellum plate produces swathes of the finest gossamer silk, which is drawn out — hackled — by combs on the spider's legs [1Foelix R. Biology of Spiders. Oxford University Press, Oxford1996Google Scholar, 4Peters H.M. Fine structure and function of capture threads.in: Nentwig W. Ecophysiology of Spiders. Springer, Berlin1987: 187-202Crossref Google Scholar]. This combing charges up the drying filaments, which repel each other and puff out to form a nanoscopic ‘wool’ yarn [4Peters H.M. Fine structure and function of capture threads.in: Nentwig W. Ecophysiology of Spiders. Springer, Berlin1987: 187-202Crossref Google Scholar]. The resulting filamentous tangle clings to, and often totally covers, a pair of much thicker support fibres issuing from spigots on the main spinnerets. This multiple-fibre rope can be further reinforced by crimped, spring-like fibres from yet another set of spigots, which pull up tight the by now rather complex, composite fibre (Figure 1A, inset). Hackling cribellum silk takes time, and a cribellate spider moving along and laying a thread in its web is slow (Figure 1A, red line). The manufacture of this complex fibre is not only time consuming, but also energetically costly, as is apparent from observation of the two hind-legs which rapidly comb away, not unlike a bassist fiddling busily — with a bow in each hand. The costs are well invested, it seems, as the cribellate capture-silk is rather sticky [5Opell B.D. The ability of spider cribellar prey capture thread to hold insects with different surface features.Funct. Ecol. 1994; 8: 145-150Crossref Scopus (38) Google Scholar, 6Opell D.B. Economics of spider orb-webs: the benefits of producing adhesive capture thread and of recycling silk.Funct. Ecol. 1998; 12: 613-624Crossref Scopus (89) Google Scholar] and is thought to bond with the prey's surface via Van der Waal's charges [7Hawthorn A.C. Opell B.D. Evolution of adhesive mechanisms in cribellar spider prey capture thread: evidence for van der Waals and hygroscopic forces.Biol. J. Linn. Soc. 2002; 77: 1-8Crossref Scopus (48) Google Scholar]. Clearly, both in form and function the hackled silk composite is a far cry from the simple, smooth double-filament strand of the typical spider silk. Hackled silk is complex even in comparison with the typical silkworm silk, with its multi-protein sheath embedding a pair of fibroin core fibres [8Shao Z. Vollrath F. The surprising strength of silkworm silk.Nature. 2002; 418: 741Crossref PubMed Scopus (716) Google Scholar]. This complexity of the hackled, cribellate silk raises interesting questions: How does the mixed multi-strand structure affect the mechanical properties of the fibre? How does this silk composite fit into the wider picture of silk function? Where does it stand in the evolution of web engineering? Blackledge and Hayashi [3Blackledge T. Hayashi C. Unravelling the mechanical properties of composite silk threads spun by cribellate orb-weaving spiders.J. Exp. Biol. 2006; 209: 3131-3140Crossref PubMed Scopus (74) Google Scholar] have recently addressed the first two of these questions: how do the cribellate capture threads compare between species with very different ecologies? How do they compare with each species' other, non-cribellate (structural) threads? And how do cribellate capture threads compare with capture threads made by spiders lacking a cribellum? These are relative newcomers, the Ecribellata, which, early in the Cretaceous [9Selden P.A. Orb-web weaving spiders in the early Cretaceous.Nature. 1989; 340: 711-713Crossref Scopus (78) Google Scholar], ‘gave up’ their cribellum and, instead of the dry, adhesive nano-filaments of the Cribellata, catch their flies by deploying a water based glyco-protein glue [10Vollrath F. Tillinghast E. Glycoprotein glue inside the spider web's aqueous coat.Naturwissenschaften. 1991; 78: 557-559Crossref Scopus (109) Google Scholar]. Structurally and functionally, the ecribellate capture thread is very different (Figure 1B, inset) and relies on its self-assembling water droplets [11Edmonds D. Vollrath F. The contribution of atmospheric water vapour to the effectiveness of a spider's web.Proc. Roy. Soc. Lond. B. 1992; 248: 145-148Crossref Scopus (48) Google Scholar] to soften the core fibre silk [12Bonthrone K.M. Vollrath F. Hunter B. Sanders J.K.M. The elasticity of spider's webs is due to water-induced mobility at a molecular level.Proc. Roy. Soc. Lond. B. 1992; 248: 141-144Crossref Scopus (45) Google Scholar] and render it rubbery [13Gosline J.M. Denny M.W. Demont M.E. Spider silk as rubber.Nature. 1984; 309: 551-552Crossref Scopus (318) Google Scholar]. Thus, in the wet capture silk of the ecribellate spiders, the functional mechanisms of extensibility, contraction and adhesion are uncoupled from the spider's production system, while in the dry cribellate silk the animal's actions directly determine both the material's structure and its function. Moreover, the two types of capture silk respond very differently to stretching [14Köhler T. Vollrath F. Thread biomechanics in the two orb weaving spiders Araneus diadematus (Araneae, Araneidae) and Uloborus walckenaerius (Araneae, Uloboridae).J. Exp. Zool. 1995; 271: 1-17Crossref Scopus (102) Google Scholar]. While the wet capture thread of the ecribellate orb extends in a smooth curve to its ultimate stretch of several times its original length, the dry cribellate silks are less stretchy and always have a rather ragged extension curve (Figure 1A, blue inset). Blackledge and Hayashi [3Blackledge T. Hayashi C. Unravelling the mechanical properties of composite silk threads spun by cribellate orb-weaving spiders.J. Exp. Biol. 2006; 209: 3131-3140Crossref PubMed Scopus (74) Google Scholar] measured the bulk properties of both types of capture thread, and argue that the mechanical response of the cribellate thread reflects the multi-strand properties with the thicker core fibres over-stretching and breaking well before the hackled filaments. The dramatic fall in the stress-strain curve would indicate the point when the core fibres break. This would then be followed by the thin cribellum threads stretching out ever longer until they, eventually, fail singly or in small groups, creating the ragged response curve. This hypothesis is interesting and could perhaps be further tested by examining the response curves to repeated cycles of stretching and relaxing these threads both below and above the breaking point of the core fibres. Fully understanding this silk is important as it opens the way to interpret the evolution of spiders in general and the orb weavers specifically [1Foelix R. Biology of Spiders. Oxford University Press, Oxford1996Google Scholar, 15Penney D. Does the fossil record of spiders track that of, their principal prey, the insects?.Trans. Roy. Soc. Edinburgh. 2004; 94: 275-281Google Scholar]. About 130 million years ago [9Selden P.A. Orb-web weaving spiders in the early Cretaceous.Nature. 1989; 340: 711-713Crossref Scopus (78) Google Scholar, 15Penney D. Does the fossil record of spiders track that of, their principal prey, the insects?.Trans. Roy. Soc. Edinburgh. 2004; 94: 275-281Google Scholar] the nano-filamentous ‘rope’ structure of the cribellate capture thread seems to have been first challenged and then largely superseded by another capture-thread mechanism: a micro-scale windlass (Figure 1B, inset). This mechanism operates on a very different size scale and uses a very different mechanism to the hackled threads. It is set up by the spider coating the paired core fibres as they emerge from their spigots [4Peters H.M. Fine structure and function of capture threads.in: Nentwig W. Ecophysiology of Spiders. Springer, Berlin1987: 187-202Crossref Google Scholar] with a fine layer of an aqueous solution containing a surprising (for a silk) mix of compounds [16Vollrath F. Fairbrother W.J. Williams R.J.P. Tillinghast E.K. Bernstein D.T. Gallager K.S. Townley M.A. Compounds in the droplets of the orb spider's viscid spiral.Nature. 1990; 345: 526-528Crossref Scopus (160) Google Scholar]. Neurotransmitter molecules in the solution render it strongly hygroscopic and the thin layer, immediately after extrusion, rapidly takes up water from the atmosphere, swells, becomes unstable and forms evenly spaced Raleigh droplets [11Edmonds D. Vollrath F. The contribution of atmospheric water vapour to the effectiveness of a spider's web.Proc. Roy. Soc. Lond. B. 1992; 248: 145-148Crossref Scopus (48) Google Scholar], resembling pearls on a string. Inside each droplet a little torus of glyco-protein segregates into a sticky dollop of glue straddling the pair of core fibres [10Vollrath F. Tillinghast E. Glycoprotein glue inside the spider web's aqueous coat.Naturwissenschaften. 1991; 78: 557-559Crossref Scopus (109) Google Scholar]. The formation of this silk composite needs neither hackling nor any other action by the spider. The whole, complex system thus is fully self-assembling. It is also very effective as an elastomer composite [17Vollrath F. Porter D. Spider silk as archetypal protein elastomer.Softmatter. 2006; 2: 377-385Google Scholar]. The water plasticises the core fibres making them soft and extensible well beyond their dry range [18Vollrath F. Edmonds D. Modulation of normal spider silk by coating with water.Nature. 1989; 340: 305-307Crossref Scopus (215) Google Scholar]. Stretching the fibre composite deforms the water droplets, which are interconnected by water sleeves [11Edmonds D. Vollrath F. The contribution of atmospheric water vapour to the effectiveness of a spider's web.Proc. Roy. Soc. Lond. B. 1992; 248: 145-148Crossref Scopus (48) Google Scholar]. Relaxing the thread allows the water droplets to minimize surface tension by re-assuming their more rounded shape in the process drawing-in the core fibres, which curl up inside each droplet into a little ball [18Vollrath F. Edmonds D. Modulation of normal spider silk by coating with water.Nature. 1989; 340: 305-307Crossref Scopus (215) Google Scholar]. The effortless self-assembling process gives the ecribellate spiders a competitive edge. The hackled threads may (or may not, the judgment is still out) be stickier for longer or on more surfaces [6Opell D.B. Economics of spider orb-webs: the benefits of producing adhesive capture thread and of recycling silk.Funct. Ecol. 1998; 12: 613-624Crossref Scopus (89) Google Scholar], but they certainly are much costlier in both assembly time and metabolism as they cannot do without the combing process. But the windlass system also wins in mechanical performance, allowing for very many cycles of very large extensions (>400%) with the threads always maintaining thread tension. Hence we should not be surprised to find that the ecribellate spiders with their gluey threads outnumber the cribellate spiders [1Foelix R. Biology of Spiders. Oxford University Press, Oxford1996Google Scholar]. This superiority is most obvious among the orb-weavers, with thousands of ecribellate species and only a handful of cribellate ones. A study analysing silk gene sequences [19Garb J.E. DiMauro T. Vo V. Hayashi C.Y. Silk genes support the single origin of orb-webs.Science. 2006; 312: 1762Crossref PubMed Scopus (79) Google Scholar] suggests that, probably over 130 million years ago, a tribe of cribellate orb-weavers might have evolved the windlass system. This ‘invention’ would have enabled a rapid and thorough adaptive radiation of the web-building behaviour that could make use of this new capture mechanism thus giving rise to the huge abundance as well as diversity of the aerial webs using glue instead of nano-filaments to fasten flies. While the mechanical behaviour of the cribellum fibres is interesting for the arachnologist it also may have some important lessons for the materials engineer. The diameter of the smaller filaments range between 10 and 100 nanometers, there are thousands of them in a bundle, and the individal filaments are all further processed post-extrusion by the spider's hackling to give specific properties (most of which we do not yet understand). Anyone interested in nano-scale materials should take note, not least the rapidly growing community of materials scientists working on electro-spinning with the aim of making bundles and mats of electrically charged nanofibres. After all, the cribellate spiders have been doing this kind of thing for millions of years. The cribellate silk and capture threads are presumably well tried and tested — like any material or structure that develops in an ‘arms-race’ as intense as the contest between the fly and the spider." @default.
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• W2078255086 title "Spider Silk: Thousands of Nano-Filaments and Dollops of Sticky Glue" @default.
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