Hierarchical looping results in extreme extensibility of silk fibre composites produced by Southern house spiders (Kukulcania hibernalis).

Spider silk is a tough and versatile biological material combining high tensile strength and extensibility through nanocomposite structure and its nonlinear elastic behaviour. Notably, spiders rarely use single silk fibres in isolation, but instead process them into more complex composites, such as silk fibre bundles, sheets and anchorages, involving a combination of spinneret, leg and body movements. While the material properties of single silk fibres have been extensively studied, the mechanical properties of silk composites and meta-structures are poorly understood and exhibit a hereto largely untapped potential for the bio-inspired design of novel fabrics with outstanding mechanical properties. In this study, we report on the tensile mechanics of the adhesive capture threads of the Southern house spider (Kukulcania hibernalis), which exhibit extreme extensibility, surpassing that of the viscid capture threads of orb weavers by up to tenfold. By combining high-resolution mechanical testing, microscopy and in silico experiments based on a hierarchical modified version of the Fibre Bundle Model, we demonstrate that extreme extensibility is based on a hierarchical loops-on-loops structure combining linear and coiled elements. The stepwise unravelling of the loops leads to the repeated fracture of the connected linear fibres, delaying terminal failure and enhancing energy absorption. This principle could be used to achieve tailored fabrics and materials that are able to sustain high deformation without failure.

Change of mechanical characteristics in spider silk capture threads after contact with prey.

Most spiders rely on specialized capture threads to subdue prey. Cribellate spiders use capture threads, whose adhesion is based on thousands of nanofibers instead of specialized glue. The nanofibers adhere due to van der Waals and hygroscopic forces, but the adhesion is strengthened by an interaction with the cuticular hydrocarbons (CHCs) covering almost all insects. The interaction between CHCs and cribellate threads becomes visible through migration of the CHCs into the thread even far beyond the point of contact. In this study, we were able to show that the migrated CHCs not only influence adhesion but also change the mechanical characteristics of the thread. While adhesion, extensibility and total energy decreased in threads treated with CHCs from different insects, we observed an increasing force required to break threads. Such mechanical changes could be beneficial for the spider: Upon the first impact of the insect in the web, it is important to absorb all the energy without breaking. Afterwards, a reduction in extensibility could cause the insect to stay closer to the web and thus become additionally entangled in neighboring threads. An increased tensile force would additionally ensure that for insects already in the web, it is even harder to free themselves. Taken together, all these changes make it unlikely that cribellate spiders reuse their capture threads, if not reacting rapidly and removing the prey insect before the CHCs can spread across the thread. STATEMENT OF SIGNIFICANCE: Cribellate spiders use capture threads that, unlike other spiders, consist of nanofibers and do not rely glue. Instead, prey adheres mainly because their surface compounds, so-called cuticular hydrocarbons (CHCs), interact with the thread, this way generating strong adhesion forces. Previous studies on biomechanics and adhesion of cribellate threads only dealt with artificial surfaces, neglecting any interaction with surface compounds. This study examines the dramatical mechanical changes of a cribellate thread after interaction with prey CHCs, showing modifications of the thread's extensibility, tensile force and total energy. Our results highlight the importance of studying mechanical properties of silk not only in an artificial context, but also in real life.

Why do ants differ in acclimatory ability? Biophysical mechanisms behind cuticular hydrocarbon acclimation across species.

Maintaining water balance is vital for terrestrial organisms. Insects protect themselves against desiccation via cuticular hydrocarbons (CHCs). CHC layers are complex mixtures of solid and liquid hydrocarbons, with a surprisingly diverse composition across species. This variation may translate into differential phase behaviour, and hence varying waterproofing capacity. This is especially relevant when temperatures change, which requires acclimatory CHC changes to maintain waterproofing. Nevertheless, the physical consequences of CHC variation are still little understood. We studied acclimatory responses and their consequences for CHC composition, phase behaviour and drought survival in three congeneric ant species. Colony sub-groups were kept under cool, warm and fluctuating temperature regimes. Lasius niger and Lasius platythorax, both of which are rich in methyl-branched alkanes, showed largely predictable acclimatory changes of the CHC profile. In both species, warm acclimation increased drought resistance. Warm acclimation increased the proportion of solid compounds in L. niger but not in L. platythorax. In both species, the CHC layer formed a liquid matrix of constantly low viscosity, which contained highly viscous and solid parts. This phase heterogeneity may be adaptive, increasing robustness to temperature fluctuations. In Lasius brunneus, which is rich in unsaturated hydrocarbons, acclimatory CHC changes were less predictable, and warm acclimation did not enhance drought survival. The CHC layer was more homogeneous, but matrix viscosity changed with acclimation. We showed that ant species use different physical mechanisms to enhance waterproofing during acclimation. Hence, the ability to acclimate, and thus climatic niche breadth, may strongly depend on species-specific CHC profile.

Insect cuticular hydrocarbon composition influences their interaction with spider capture threads.

Insects represent the main prey of spiders, and spiders and insects co-diversified in evolutionary history. One of the main features characterizing spiders is their web as a trap to capture prey. Phylogenetically, the cribellate thread is one of the earliest thread types that was specialized to capture prey. In contrast to other capture threads, it lacks adhesive glue and consists of nanofibres, which do not only adhere to insects via van der Waals forces but also interact with the insects' cuticular hydrocarbon (CHC) layer, thus enhancing adhesion. The CHC layer consists of multiple hydrocarbon types and is highly diverse between species. In this study, we show that CHC interaction with cribellate capture threads is affected by CHC composition of the insect. We studied the interaction in detail for four insect species with different CHC profiles and observed a differential migration of CHCs into the thread. The migration depends on the molecular structure of the hydrocarbon types as well as their viscosity, influenced by the ambient temperature during the interaction. As a consequence, adhesion forces to CHC layers differ depending on their chemical composition. Our results match predictions based on biophysical properties of hydrocarbons, and show that cribellate spiders can exert selection pressure on the CHC composition of their insect prey.