Spider silk
Spider silk is a
In some cases, spiders may use silk as a food source.[1] While methods have been developed to collect silk from a spider by force,[2] gathering silk from many spiders is more difficult than from silk-spinning organisms such as silkworms.
All spiders produce silk, although some spiders do not make webs. Silk is tied to courtship and mating. Silk produced by females provides a transmission channel for male vibratory courtship signals, while webs and draglines provide a substrate for female sex pheromones. Observations of male spiders producing silk during sexual interactions are common across widespread taxa. The function of male-produced silk in mating has received little study.[3]
Properties
Structural
Silks have a hierarchical structure. The
Termonia introduced this first basic model of silk in 1994.
The fibres' microstructural information and macroscopic mechanical properties are related.[14] Ordered regions (i) mainly reorient by deformation for low-stretched fibres and (ii) the fraction of ordered regions increases progressively for higher fibre stretching.
-
Schematic of the spider's orb web, structural modules, and spider silk structure.[15] On the left is shown a schematic drawing of an orb web. The red lines represent the dragline, radial line, and frame lines. The blue lines represent the spiral line, and the centre of the orb web is called the "hub". Sticky balls drawn in blue are made at equal intervals on the spiral line with viscous material secreted from the aggregate gland. Attachment cement secreted from the piriform gland is used to connect and fix different lines. Microscopically, the spider silk secondary structure is formed of spidroin with the structure shown on the right side. In the dragline and radial line, a crystalline β-sheet and an amorphous helical structure are interwoven. The large amount of β-spiral structure gives elastic properties to the capture part of the orb web. In the structural modules diagram, a microscopic structure of dragline and radial lines is shown, composed mainly of two proteins of MaSp1 and MaSp2, as shown in the upper central part. The spiral line has no crystalline β-sheet region.
Mechanical
Each spider and each type of silk has a set of mechanical properties optimised for their biological function.
Most silks, in particular dragline silk, have exceptional mechanical properties. They exhibit a unique combination of high
Strength and toughness are distinct quantities. Weight for weight, silk is stronger than steel, but not as strong as Kevlar. Spider silk is, however, tougher than both.
The variability of spider silk fibre mechanical properties is related to their degree of molecular alignment.[16] Mechanical properties also depend on ambient conditions, i.e. humidity and temperature.[17]
Strength
A dragline silk's
Density
Consisting of mainly protein, silks are about a sixth of the density of steel (1.3 g/cm3). As a result, a strand long enough to circle the Earth would weigh about 2 kilograms (4.4 lb). (Spider dragline silk has a tensile strength of roughly 1.3 GPa. The tensile strength listed for steel might be slightly higher – e.g. 1.65 GPa,[21][22] but spider silk is a much less dense material, so that a given weight of spider silk is five times as strong as the same weight of steel.)
Energy density
The energy density of dragline spider silk is roughly 1.2×108 J/m3.[23]
Ductility
Silks are
Toughness
The combination of strength and ductility gives dragline silks a high
Temperature
While unlikely to be relevant in nature, dragline silks can hold their strength below -40 °C (-40 °F) and up to 220 °C (428 °F).
Supercontraction
When exposed to water, dragline silks undergo supercontraction, shrinking up to 50% in length and behaving like a weak rubber under tension.[17] Many hypotheses have attempted to explain its use in nature, most popularly to re-tension webs built in the night using the morning dew.[citation needed]
Highest-performance
The toughest known spider silk is produced by the species Darwin's bark spider (Caerostris darwini): "The toughness of forcibly silked fibers averages 350 MJ/m3, with some samples reaching 520 MJ/m3. Thus, C. darwini silk is more than twice as tough as any previously described silk and over 10 times tougher than Kevlar".[27]
Adhesive
Silk fibre is a two-compound
Uses
All spiders produce silks, and a single spider can produce up to seven different types of silk for different uses.[30] This is in contrast to insect silks, where an individual usually only produces a single type.[31] Spiders use silks in many ways, in accord with the silk's properties. As spiders have evolved, so has their silks' complexity and uses, for example from primitive tube webs 300–400 million years ago to complex orb webs 110 million years ago.[32]
Use | Example | Reference |
---|---|---|
Prey capture | Orb webs produced by the Araneidae (typical orb-weavers); tube webs; tangle webs; sheet webs; lace webs, dome webs; single thread used by the Bolas spiders for "fishing". | [30][32] |
Prey immobilisation | "Swathing bands" to envelop prey. Often combined with immobilising prey using a venom. In species of Scytodes the silk is combined with venom and squirted from the chelicerae. | [30] |
Reproduction | Male spiders may produce sperm webs; spider eggs are covered in silk cocoons. | [30][33] |
Dispersal | "Ballooning" or "kiting" used by smaller spiders to float through the air, for instance for dispersal. | [34] |
Food | The kleptoparasitic Argyrodes eats the silk of host spider webs. Some daily weavers of temporary webs eat their own unused silk, thus mitigating an otherwise heavy metabolic expense.
|
[1][35] |
Nest lining and nest construction | Tube webs used by "primitive" spiders such as the European tube web spider ( Argyroneta aquatica forms a silk diving bell .
|
[32] |
Guide lines | Some spiders that venture from shelter leave a silk trail by which to find their way home again. | [35] |
Drop lines and anchor lines | Spiders such as the Salticidae venture from shelter and leave a trail of silk, use that as an emergency line in case of falling from inverted or vertical surfaces. Others, even web dwellers, deliberately drop from a web when alarmed, using a silken thread as a drop line by which they can return in due course. Some, such as species of Paramystaria, hang from a drop line while feeding.
|
[35] |
Alarm lines | Some spiders that do not spin actual trap build alarm webs that the feet of their prey (such as ants) can disturb, cueing the spider to pounce on prey or flee a formidable intruder. | [35] |
Pheromonal trails | Some wandering spiders leave a largely continuous trail of silk impregnated with pheromones that the opposite sex can follow to find a mate. | [35] |
Silk types
Meeting the specification for all these ecological uses requires different types of silk presenting different properties, as either a fibre, a structure of fibres, or a globule. These types include glues and fibres. Some types of fibres are used for structural support, others for protective structures. Some can absorb energy effectively, whereas others transmit vibration efficiently. These silk types are produced in different glands; so the silk from a particular gland can be linked to its use.
Gland | Silk Use |
---|---|
Ampullate (major) | Dragline silk – used for the web's outer rim and spokes, also for lifeline and for ballooning |
Ampullate (minor) | Used for temporary scaffolding during web construction |
Flagelliform | Capture-spiral silk – used for the capturing lines of the web |
Tubuliform | Egg cocoon silk – used for egg sacs |
Aciniform | Used to wrap and secure prey; used in male sperm webs; used in stabilimenta |
Aggregate | Sticky globules |
Piriform | Bonds between separate threads for attachment point. |
Many species have different glands to produce silk with different properties for different purposes, including housing,
Silk | Use |
---|---|
Major-ampullate (dragline) silk | The web's outer rim and spokes and the lifeline. Can be as strong per unit weight as steel, but much tougher. |
Capture-spiral (flagelliform) silk | Capturing lines. Sticky, stretchy, and tough. The capture spiral is sticky due to droplets of aggregate (a spider glue) that are placed on the spiral. The elasticity of flagelliform allows enough time for the aggregate to adhere to the aerial prey flying into the web. |
Tubiliform (a.k.a. cylindriform) silk | Protective egg sacs. Stiffest silk. |
Aciniform silk | Wrap and secure prey. Two to three times as tough as the other silks, including dragline. |
Minor-ampullate silk | Temporary scaffolding during web construction. |
Synthesis and fibre spinning
Silk production differs in an important aspect from that of most other fibrous biomaterials. It is pulled on demand from a precursor out of specialised glands,[38] rather than continuously grown like plant cell walls.[23]
The spinning process occurs when a fibre is pulled away from the body of a spider, whether by the spider's legs, by the spider's falling under its own weight, or by any other method. The term "spinning" is misleading because no rotation occurs. It comes from analogy to the textile spinning wheels. Silk production is a pultrusion,[39] similar to extrusion, with the subtlety that the force is induced by pulling at the finished fibre rather than squeezing it out of a reservoir. The fibre is pulled through (possibly multiple) silk glands of multiple types.[38]
Silk gland
The gland's visible, or external, part is termed the
Behind each spinneret on the surface of the spider lies a gland, a generalised form of which is shown in the figure.
- Gland characteristics
- The leftmost section s the secretory or tail section. The walls of this section are lined with cells that secrete proteins Spidroin I and Spidroin II, the main components of this spider's dragline. These proteins are found in the form of droplets that gradually elongate to form long channels along the length of the final fibre, hypothesised to assist in preventing crack formation or self-healing.[56]
- The ampulla (storage sac) is next. This stores and maintains the gel-like unspun silk dope. In addition, it secretes proteins that coat the surface of the final fibre.[24]
- The funnel rapidly reduces the large diameter of the storage sac to the small diameter of the tapering duct.
- The final length is the tapering duct, the site of most of the fibre formation. This consists of a tapering tube with several tight sharp turns, a valve near the end includes a spigot from which the solid silk fibre emerges. The tube tapers hyperbolically, therefore the unspun silk is under constant elongational shear stress, an important factor in fibre formation. This section is lined with cells that exchange ions, reduce the dope pH from neutral to acidic, and remove water from the fibre.[57] Collectively, the shear stress and the ion and pH changes induce the liquid silk dope to undergo a phase transition and condense into a solid protein fibre with high molecular organisation. The spigot at the end has lips that clamp around the fibre, controlling fibre diameter and further retaining water.
- Almost at the end is a valve. Though discovered some time ago, its precise purpose is still under discussion. It is believed to assist in restarting and rejoining broken fibres, The similarity of the silk worm's silk press and the roles each of these valves play in the silk production in these two organisms are under discussion.
Throughout the process the silk appears to have a nematic texture,[60] in a manner similar to a liquid crystal, arising in part due to the high protein concentration of silk dope (around 30% in terms of weight per volume).[61] This allows the silk to flow through the duct as a liquid while maintaining molecular order.
As an example of a complex spinning field, the spinneret apparatus of an adult Araneus diadematus (garden cross spider) consists of many glands shown below.[9] A similar gland architecture appears in the black widow spider.[62]
- 500 pyriform glands for attachment points
- 4 ampullate glands for the web frame
- 300 aciniform glands for the outer lining of egg sacs, and for ensnaring prey
- 4 tubuliform glands for egg sac silk
- 4 aggregate glands for adhesive functions
- 2 coronate glands for the thread of adhesion lines
Artificial synthesis
To artificially synthesise spider silk into fibres, two broad tasks are required. These are synthesis of the feedstock (the unspun silk dope in spiders), and synthesis of the production conditions (the funnel, valve, tapering duct, and spigot). Few strategies have produced silk that can efficiently be synthesised into fibres.
Feedstock
The molecular structure of unspun silk is both complex and long. Though this endows the fibres with desirable properties, it also complicates replication. Various organisms have been used as a basis for attempts to replicate necessary protein components. These proteins must then be extracted, purified, and then spun before their properties can be tested.
Organism | Details | Average Maximum breaking stress (MPa) | Average Strain (%)
|
Reference |
---|---|---|---|---|
Darwin's bark spider (Caerostris darwini) | Malagasy spider famed for making 25 m long strands. "C. darwini silk is more than twice as tough as any previously described silk"[citation needed] | 1850 ±350 | 33 ±0.08 | [27] |
Nephila clavipes | Typical golden orb weaving spider | 710–1200 | 18–27 | [63][64] |
Bombyx mori Silkworms | Silkworms genetically altered to express spider proteins and fibres.[65] | 660 | 18.5 | [66] |
Escherichia coli | Synthesising a large and repetitive molecule (~300 kDa ) is complex, but required for the strongest silk. Here E. coli was engineered to produce a 556 kDa protein. Fibers spun from these synthetic spidroins are the first to fully replicate the mechanical performance of natural spider silk by all common metrics. |
1030 ±110 | 18 ±6 | [67] |
Goats | Genetically modified to secrete silk proteins in their milk. | 285–250 | 30–40 | [68] |
Tobacco & potato plants | Genetically modified to produce silk proteins. Patents were granted in 2010,[69] but no fibres have yet been described. | n/a | n/a | [70] |
Geometry
Spider silks with comparatively simple molecular structure need complex ducts to be able to form an effective fibre. Approaches:
Syringe and needle
Feedstock is forced through a hollow needle using a syringe.[71][72]
Although cheap and easy to produce, gland shape and conditions are loosely approximated. Fibres created using this method may need encouragement to solidify by removing water from the fibre with chemicals such as (environmentally undesirable) methanol[73] or acetone,[72] and also may require later stretching of the fibre to achieve desirable properties.[74][71]
Superhydrophobic surfaces
Placing a solution of spider silk on a superhydrophobic surface can generate sheets, particles, and nanowires of spider silk.[75][76]
Sheets
Self-assembly of silk at standing liquid-gas interphases of a solution tough and strong sheets. These sheets are now explored for mimicking the basal membrane in tissue modeling.[77][78]
Microfluidics
Microfluidics have the advantage of being controllable and able to test spin small volumes of unspun fibre,[79][80] but setup and development costs are high. A patent has been granted and continuously spun fibres have achieved commercial use.[81]
Electrospinning
Electrospinning is an old technique whereby a fluid is held in a container such that it flows out through capillary action. A conducting substrate is positioned below, and a difference in electrical potential is applied between the fluid and the substrate. The fluid is attracted to the substrate, and tiny fibres jump from their point of emission, the Taylor cone, to the substrate, drying as they travel. This method creates nano-scale fibres from silk dissected from organisms and regenerated silk fibroin.[citation needed]
Other shapes
Silk can be formed into other shapes and sizes such as spherical capsules for drug delivery, cell scaffolds and wound healing, textiles, cosmetics, coatings, and many others.[82][83] Spider silk proteins can self-assemble on superhydrophobic surfaces into nanowires, as well as micron-sized circular sheets.[83] Recombinant spider silk proteins can self-assemble at the liquid-air interface of a standing solution to form protein-permeable, strong and flexible nanomembranes that support cell proliferation. Potential applications include skin transplants, and supportive membranes in organ-on-a-chip.[84] These nanomembranes have been used to create a static in-vitro model of a blood vessel.[85]
Synthetic spider silk
Replicating the complex conditions required to produce comparable fibres has challenged research and early-stage manufacturing. Through genetic engineering, E. coli bacteria, yeasts, plants, silkworms, and animals other than silkworms have been used to produce spider silk-like proteins, which have different characteristics than those from a spider.[86] Extrusion of protein fibres in an aqueous environment is known as "wet-spinning". This process has produced silk fibres of diameters ranging from 10 to 60 μm, compared to diameters of 2.5–4 μm for natural spider silk. Artificial spider silks have fewer and simpler proteins than natural dragline silk, and consequently offer half the diameter, strength, and flexibility of natural dragline silk.[86]
Research
- In March 2010, researchers from the Nephila clavipes. This approach eliminates the need to "milk" spiders.[87]
- A 556 kDa spider silk protein was manufactured from 192 repeat motifs of the N. clavipes dragline spidroin, having similar mechanical characteristics as their natural counterparts, i.e.,
- AMSilk developed spidroin using bacteria.[86][88]
- Bolt Threads produced a recombinant spidroin using yeast, for use in apparel fibers and personal care. They produced the first commercial apparel products made of recombinant spider silk, trademarked Microsilk, demonstrated in ties and beanies.[89][90]
- Kraig Biocraft Laboratories used research from the Universities of Wyoming and Notre Dame to create silkworms genetically altered to produce spider silk.[91][92]
- Defunct Canadian
- Spiber produced a synthetic spider silk (Q/QMONOS). In partnership with Goldwin, a ski parka made from this was in testing in 2016.[95][96]
- Researchers from Japan's RIKEN Center constructed an artificial gland that reproduced spider silk's molecular structure. Precise microfluidic mechanisms directed proteins to self-assemble into functional fibers. The process used negative pressure to pull (rather than push) a spidroin solution through the device. The resulting fibers matched the hierarchical structure of natural fiber.[97]
Research
Area of contribution | Year | Main researchers | Title | Contribution to the field |
---|---|---|---|---|
Chemical Basis | 1960 | Fischer, F. & Brander, J.[98] | "Eine Analyse der Gespinste der Kreuzspinne" (Amino acid composition analysis of spider silk) | |
1960 | Lucas, F. et al.[99][100] | "The Composition of Arthropod Silk Fibroins; Comparative studies of fibroins" | ||
Gene Sequence | 1990 | Xu, M. & Lewis, R. V.[101] | "Structure of a Protein Superfiber − Spider Dragline Silk" | |
Mechanical Properties | 1964 | Lucas, F.[102] | "Spiders and their silks" | First time mechanical properties of spider silk compared with other materials in a scientific paper. |
1989 | Vollrath, F. & Edmonds, D. T.[103] | "Modulation of the Mechanical Properties of Spider Silk by Coating with Water" | First important paper suggesting the water interplay with spider silk fibroin modulating silk properties. | |
2001 | Vollrath, F. & Shao, Z.Z.[104] | "The effect of spinning conditions on the mechanics of a spider's dragline silk" | ||
2006 | Plaza, G.R., Guinea, G.V., Pérez-Rigueiro, J. & Elices, M.[17] | "Thermo-hygro-mechanical behavior of spider dragline silk: Glassy and rubbery states" | Combined effect of humidity and temperature on mechanical properties. Glass-transition temperature dependence on humidity. | |
Structural Characterisation | 1992 | Hinman, M.B. & Lewis, R. V[4] | "Isolation of a clone encoding a second dragline silk fibroin. Nephila clavipes dragline silk is a two-protein fiber" | |
1994 | Simmons, A. et al.[105] | "Solid-State C-13 Nmr of Nephila-Clavipes Dragline Silk Establishes Structure and Identity of Crystalline Regions" | First NMR study of spider silk. | |
1999 | Shao, Z., Vollrath, F. et al.[106] | "Analysis of spider silk in native and supercontracted states using Raman spectroscopy" | First Raman study of spider silk. | |
1999 | Riekel, C., Muller, M.et al.[107] | "Aspects of X-ray diffraction on single spider fibers" | First X-ray on single spider silk fibres. | |
2000 | Knight, D.P., Vollrath, F. et al.[108] | "Beta transition and stress-induced phase separation in the spinning of spider dragline silk" | Secondary structural transition confirmation during spinning. | |
2001 | Riekel, C. & Vollrath, F.[109] | "Spider silk fibre extrusion: combined wide- and small-angle X- ray microdiffraction experiments" | First X-ray on spider silk dope. | |
2002 | Van Beek, J. D. et al.[6] | "The molecular structure of spider dragline silk: Folding and orientation of the protein backbone" | ||
Structure-Property Relationship | 1986 | Gosline, G.M. et al.[110] | "The structure and properties of spider silk" | First attempt to link structure with properties of spider silk |
1994 | Termonia, Y[10] | "Molecular Modeling of Spider Silk Elasticity" | X-ray evidence presented in this paper; simple model of crystallites embedded in amorphous regions. | |
1996 | Simmons, A. et al.[5] | "Molecular orientation and two-component nature of the crystalline fraction of spider dragline silk" | Two types of alanine-rich crystalline regions were defined. | |
2006 | Vollrath, F. & Porter, D.[111] | "Spider silk as an archetypal protein elastomer" | New insight and model to spider silk based on Group Interaction Modelling. | |
Native Spinning | 1991 | Kerkam, K., Kaplan, D. et al.[112] | "Liquid Crystallinity of Natural Silk Secretions" | |
1999 | Knight, D.P. & Vollrath, F.[113] | "Liquid crystals and flow elongation in a spider's silk production line" | ||
2001 | Vollrath, F. & Knight, D.P.[23] | "Liquid crystalline spinning of spider silk" | Most cited spider silk paper | |
2005 | Guinea, G.V., Elices, M., Pérez-Rigueiro, J. & Plaza, G.R.[16] | "Stretching of supercontracted fibers: a link between spinning and the variability of spider silk" | Explanation of the variability of mechanical properties. | |
Reconstituted /Synthetic Spider Silk and Artificial Spinning | 1995 | Prince, J. T., Kaplan, D. L. et al.[114] | "Construction, Cloning, and Expression of Synthetic Genes Encoding Spider Dragline Silk" | First successful synthesis of Spider silk by E. coli. |
1998 | Arcidiacono, S., Kaplan, D.L. et al.[115] | "Purification and characterization of recombinant spider silk expressed in Escherichia coli" | ||
1998 | Seidel, A., Jelinski, L.W. et al.[116] | "Artificial Spinning of Spider Silk" | First controlled wet-spinning of reconstituted spider silk. |
Human uses
The earliest recorded attempt to weave fabric from spider silk was in 1709 by
The development of methods to
Medicine
Peasants in the southern
Science and technology
Spider silk has been used as a thread for
Silk has been used to suspend inertial confinement fusion targets during laser ignition, as it remains considerably elastic and has a high energy to break at temperatures as low as 10–20 K. In addition, it is made from "light" atomic number elements that emit no x-rays during irradiation that could preheat the target, limiting the pressure differential required for fusion.[129]
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External links
- "The Silk Spinners", a BBC program about silk-producing animals
- Meadows, Robin (5 August 2014). "How Spiders Spin Silk". PLOS Biology. 12 (8): e1001922. PMID 25093404.
- Rejcek, Peter (11 April 2019). "The Tangled Web of Turning Spider Silk into a Super Material". Singularity Hub. Retrieved 24 April 2019.
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- "Synthetic spider silk stronger and tougher than the real thing". New Atlas. 21 July 2021. Retrieved 21 July 2021.