Inspiration from Verse 41 of Surah Al-Ankabut in Science and Technology

Abstract

The fragility of the spider’s web is one of the deepest and most beautiful divine allegories, employed by God in Verse 41 of Surah Al-Ankabut to illustrate the inherent weakness of structures that, despite their seemingly powerful and stable appearance, are fragile before the majesty and will of the Divine. This allegory reflects the Quranic miracle in expressing truths that are simultaneously scientific, mystical, and social. Through the phrases “if only they knew” (law kānū yaʿlamūn) and “none comprehend it except the knowledgeable” (mā yaʿqiluhā illā al-ʿālimūn) in this verse and Verse 43, God emphasizes that understanding this analogy is impossible without knowledge and awareness, as reflecting upon it requires expertise, and only the learned can grasp its depths.

Though spider silk appears delicate and fragile, it is composed of one of the most advanced biological metamaterials (Bio-Metamaterials) in the world. This study, through extensive analysis of existing research on this biological metamaterial and its impact on scientific and technological advancement, presents its findings in four sections and identifies six pioneering research areas for future studies.

Keywords: Biological Metamaterials, Spider Silk, Spider Web.

1. Introduction

The fragility of the spider’s web is one of the most profound and exquisite divine allegories employed by God in Verse 41 of Surah Al-Ankabut to illustrate the inherent weakness of structures that, despite their outwardly powerful and stable appearance, are fragile before the majesty and divine will. This allegory exemplifies the Quranic miracle in conveying truths that are simultaneously scientific, mystical, and social. Through the phrases “if only they knew” (law kānū yaʿlamūn) and “none comprehend it except the knowledgeable” (mā yaʿqiluhā illā al-ʿālimūn) in this verse and Verse 43, God emphasizes that understanding this analogy is impossible without knowledge and awareness, as reflecting upon it requires expertise, and only the learned can grasp its depths.

Though spider silk appears delicate and fragile, it is composed of one of the most advanced biological metamaterials (Bio-Metamaterials) in the world. These threads, formed from specialized proteins, exhibit tensile strength surpassing steel and unparalleled lightness and flexibility. Had the natural world been defined solely by the scale of flies and bees, the spider—with its web of such wondrous materials and sophisticated techniques—would reign as the ultimate superpower. Yet, in reality, no structure, despite its advanced material, is as easily destroyed as the spider’s web. This paradox between strength and frailty carries a profound message: humans, with all their capabilities and achievements, remain dwarfed by the spider’s knowledge. If so, how can humanity withstand natural calamities or divine punishment? What can they rely upon to escape worldly and hereafter torment, unless they seek aid from God alone?

The spider’s web, with all its complexity and marvels, symbolizes the civilizations and socio-political structures humans build. These civilizations, though seemingly intricate and formidable, are as vulnerable and defenseless before divine power as the spider’s thread. Through this verse, God warns humanity to place their trust not in their limited power but in Him, the sole protector from peril and punishment. The verse presents the spider’s web as a metaphor for civilizations, structures, and individuals—like the ʿĀd, who built towering palaces, or the Thamūd, who carved fortresses from mountains; or figures such as Qārūn (symbolizing boundless wealth), Pharaoh (embodying absolute power), and Hāmān (representing deceit and political machinations). All are reflections of the spider’s web. Yet these civilizations and individuals, though appearing intricate and mighty to their peers, were annihilated by divine tempests. This analogy serves as a warning: whatever humans construct, if not founded upon reliance on God, is ultimately doomed to weakness and oblivion.

Alongside this warning, the verse invites intellectual pursuit and contemplation. By revealing the spider’s web-spinning mechanism and the complex structure of its silk, God urges us to ponder the unparalleled knowledge embedded in this tiny creature. This insect, with its home crafted from nature’s strongest known material, challenges our scholars and retains many unsolved mysteries. Yet this very web, despite its advanced technology, cannot endure wind or rain. God intends for us to learn from this intricate science while recognizing the structure’s fragility—so we may understand that no power but His can shield us from His omnipotence.

These verses deliver a dual message: they invite us to reflect and acquire knowledge, drawing lessons from creation’s complex mechanisms, while also warning of the instability of anything not rooted in reliance on God. How can humans, still incapable of fully comprehending the spider’s web, hope to build civilizations that withstand divine power? This allegory jolts the human mind, dispelling arrogance and false confidence in limited abilities, and directing it toward faith and trust in the Infinite Creator. “None comprehend it except the knowledgeable” (mā yaʿqiluhā illā al-ʿālimūn).

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2. Research Methodology

2-1. Theological and Quranic Research Methodology

This study draws upon Verse 31 of Surah Al-Māʾidah and cites authoritative exegeses (tafsīr), including: Tafsīr al-Mīzān fī Tafsīr al-Qurʾān, Tafsīr al-Namūnah, Al-Amthal fī Tafsīr Kitāb Allāh al-Munzal, Al-Tibyān fī Tafsīr al-Qurʾān, Majmaʿ al-Bayān, Tafsīr Jawāmiʿ al-Jāmiʿ, Bayān al-Saʿādah fī Maqāmāt al-ʿIbādah, Jāmiʿ al-Bayān ʿan Taʾwīl Āy al-Qurʾān (Tafsīr al-Ṭabarī), Al-Durr al-Manthūr fī al-Tafsīr bi-l-Maʾthūr, Al-Tafsīr al-Kabīr (Mafātīḥ al-Ghayb), Al-Kashshāf, Al-Kashf wa-l-Bayān fī Tafsīr al-Qurʾān (Tafsīr al-Thaʿlabī), Tafsīr Rūḥ al-Bayān, Al-Taḥrīr wa-l-Tanwīr min al-Tafsīr (Ibn ʿĀshūr), Tafsīr Kabīr Minhaj al-Ṣādiqīn fī Ilzām al-Mukhālifīn, Tafsīr al-Jalālayn, Tafsīr Qurʾān-e Mehr, Tafsīr al-Qummī, Tafsīr Nūr al-Thaqalayn, Tafsīr al-Ṣāfī, Tafsīr Kanz al-Daqāʾiq wa Baḥr al-Gharāʾib, among others.

2-2. Scientific Research Methodology

Three key insights derived from Verse 41 of Surah Al-Ankabut served as scientific leads, investigated through comprehensive searches in high-impact journals (Nature, Science, and specialized journals with Impact Factors >5), which publish cutting-edge global research. Keywords included Spider, Spider-web, Spider Silk, and Technology. Outputs were categorized based on concepts extracted from the verse, and the most relevant articles were selected for analysis.

3. Results and Discussion

In the natural world, certain materials challenge the boundaries of human imagination. One such natural masterpiece is spider silk. This extraordinary material, with properties difficult to replicate synthetically, combines strength, elasticity, lightness, and toughness in a way rarely seen in engineered materials. Remarkably, spider silk is five times stronger than steel and twice as elastic as Kevlar, while being significantly lighter [1, 2]. These unique properties stem from its specialized protein structure, which consists of alternating crystalline β-sheet regions and amorphous domains, creating an unparalleled balance between strength and flexibility [3, 4].

The spider’s web, meticulously and intricately woven, serves not only as a shelter and hunting tool but also as a blueprint for engineering designs and smart materials. Its radial and spiral architecture is optimized to withstand extreme forces even with minor structural defects [2, 5]. Additionally, the web’s adhesive properties and vibrational sensing capabilities allow spiders to detect the subtle movements of prey from a distance. Scientific studies on spider silk have a long history, beginning with early observations by naturalists. In the 19th century, systematic investigations into the mechanical properties of spider silk revealed its exceptional strength and elasticity [1]. With advancements in materials science during the 20th century, spectroscopic and electron microscopy techniques enabled researchers to uncover its molecular structure, including crystalline β-sheets and amorphous regions [4, 5].

However, this was only the beginning. Since the 1990s, efforts to synthesize artificial spider silk have intensified, with genetic engineering emerging as a key tool for reproducing spidroins [3]. In recent decades, research has shifted toward nanostructural analysis, vibrational properties, and bioinspired applications in advanced technologies [6, 7]. These studies, inspired by the biological structures and complex behaviors of spiders, have opened new frontiers in materials science and engineering. Spider silk has inspired innovations in biomedical engineering, defense technology, and even 4D printing [8, 9]. The spider’s ability to convert a protein solution into solid fibers at ambient temperature and pressure remains one of materials science’s greatest puzzles, driving extensive efforts to replicate it industrially [4, 10].

This article provides a detailed exploration of spider silk’s marvels—from its molecular structure to its technological applications—offering fresh insights into one of nature’s most advanced engineered systems. Through their silent, miniature world, spiders compel us to consider how nature’s precision and ingenuity might address humanity’s greatest challenges.

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3-1. Section 1: Structure and Composition of Spider Silk

This section examines the molecular structure, mechanical properties, and environmental influences on spider silk, as well as the silk production process and the transformation of protein solution into solid fibers.

3-1-1. Silk Production Process

The production of spider silk is a biological marvel, beginning with the synthesis of spidroin proteins in specialized glands. These high-molecular-weight proteins form a solution that, due to its unique composition, can transition into a solid, high-strength fiber. The spidroin solution is stored in glands under controlled pH and ionic conditions to prevent premature structuring [4, 11].

During spinning, the solution passes through a narrow duct called the spinneret, where key chemical and mechanical changes occur. A drop in pH and mechanical shear forces induce polypeptide chain rearrangement, forming crystalline β-sheet structures. This phase transition converts the solution into a solid fiber [3, 5].

Spiders precisely regulate fiber thickness and tension by adjusting the spinneret’s extrusion speed and force, enabling them to produce silk with varying properties. Post-spinning, the fiber is exposed to air, where drying stabilizes its crystalline and amorphous regions, achieving an optimal balance of strength and flexibility. The mechanical spinning process—central to silk formation—ensures a harmonious distribution of rigid crystalline domains and elastic amorphous regions, granting the silk its unmatched mechanical properties [2, 7]. This energy-efficient process, occurring at ambient conditions, has inspired sustainable advanced material production [3, 12].

3-1-2. Silk Recycling and Regeneration

Spider silk is among the few biological materials capable of full recycling. This unique feature stems from the biophysical and chemical properties of spidroins, which allow controlled degradation, recovery, and reuse of damaged webs. When a spider decides to rebuild its web, it ingests old silk, subjecting it to enzymatic breakdown into amino acids and small oligomers. These components are then transported back to the silk glands, where they are reincorporated into new spidroins [13, 14].

The solubility and structural adaptability of spidroins facilitate this process. While amorphous regions dissolve easily under enzymatic or environmental triggers, crystalline β-sheets degrade in a controlled manner. This dual mechanism allows spiders to reprocess old silk without compromising the mechanical integrity of new fibers [4].

Biologically, silk recycling offers critical advantages: it conserves energy in resource-scarce environments (since de novo silk production is energetically costly), minimizes ecological waste, and enables adaptive web reconstruction. Orb-weaver spiders, for instance, routinely dismantle and recycle webs, particularly in food-limited habitats—a survival strategy highlighting silk’s sustainability [13].

This recyclability has inspired synthetic materials designed for closed-loop production in fields like medicine and green engineering, where energy efficiency and waste reduction are paramount [14].

3-1-3. Molecular Structure and Spidroin Proteins

Spider silk consists of specialized spidroin proteins, whose unique molecular architecture underpins its exceptional properties. Spidroins comprise two primary domains: crystalline (β-sheet) and amorphous regions, which together confer strength, elasticity, and toughness [3, 4].

• Crystalline (β-sheet) regions act as mechanical anchors. These tightly packed, hydrogen-bonded structures provide tensile strength and fracture resistance. Though smaller than steel or Kevlar crystals, their nanoscale density ensures remarkable strength-to-weight ratios [1]. Distributed throughout the fiber, β-sheets maintain stability under environmental stress (e.g., temperature/humidity shifts) and form during spinning via pH-driven self-assembly [11].
• Amorphous regions impart flexibility. Composed of disordered glycine- and alanine-rich chains, they absorb mechanical energy and distribute stress, preventing localized failures. These regions also mediate humidity-responsive behavior, softening when hydrated—a trait exploited in biomimetic smart materials [7].

The synergy between these domains enables spider silk to endure extreme stresses while dissipating energy. Their optimized distribution creates an unparalleled equilibrium of stiffness and elasticity [3, 4].

3-1-4. Mechanical Properties of Spider Silk

Spider silk’s mechanical prowess arises from its molecular interplay, as evidenced by recent studies:

• High tensile strength: β-sheet networks allow some silks to rival steel [1, 13].
• Superelasticity: Amorphous regions enable ~30% elongation without rupture [4, 14].
• Toughness: Energy absorption is facilitated by stress distribution between domains [2, 16].
• Supercontraction: Humidity-induced shrinkage (up to 50%) optimizes web tension [12, 15].
• Dynamic responsiveness: Hydration softens amorphous zones, enhancing adaptability [7, 15].
• Fatigue resistance: Withstands cyclic loading, ideal for durable applications [2, 14].
• Composite-like hierarchy: Some silks feature micro-/submicro-fibrils for added strength [16].

3-1-5. Environmental and Mechanical Influences

Spider silk’s properties adapt to external conditions:

• Humidity: Hydration increases elasticity but may reduce strength via amorphous-phase softening [7, 15]. Supercontraction aids web tension regulation [12].
• Temperature: Crystalline regions stabilize silk in cold, while heat may compromise amorphous integrity [14].
• Strain rate: Rapid loading engages β-sheets (high strength); slow loading exploits amorphous plasticity (high toughness) [7].

Despite its adaptability, spider silk has limits. While optimized for distributed forces (e.g., wind, prey impact), concentrated stresses (e.g., human contact) cause rapid failure—a trade-off that facilitates energy-efficient web recycling but underscores the bounds of its resilience.

3-2. Section 2: Behavioral Complexity and Web Architecture

The spider web, particularly in Orb-weaver species, represents a masterpiece of biological design refined through millions of years of evolution. The construction, adjustment, and reconstruction of these intricate structures demonstrate spiders’ remarkable adaptability to environmental changes and biological needs. Beyond their primary role in prey capture, webs function as self-regulating systems capable of autonomously optimizing their architecture and material properties.

3-2-1. Web Construction Process

Web construction follows a defined sequence of behaviors: establishing a primary frame, adding radial threads, and completing adhesive spiral patterns. Research confirms that these stages follow consistent behavioral patterns across species, though environmental factors like wind or energy constraints may alter them. For instance, under resource limitations, spiders may use less silk or reduce radial threads [17, 18].

3-2-2. Web Architecture and Patterns

The architecture of spider webs is optimized for stress distribution and hunting efficiency. Radial threads form the structural backbone, connected to spiral junctions—key points with high mechanical strength that enhance stability. Studies show that variations in web geometry (e.g., spiral spacing or radial thickness) directly influence hunting success, with spiders continuously adjusting designs to environmental demands [19, 20].

3-2-3. Adaptive Web Design Behaviors

Spiders exhibit remarkable plasticity in web construction:

• High winds: Shorter radials and tighter spirals improve stability [16, 17].
• Food scarcity: Smaller web size or reduced adhesive density conserves energy [17].
• Cave-dwelling species: Simplify web structure while enhancing stickiness for low-light environments [21].

3-2-4. Web Repair and Recycling

Spiders efficiently repair damaged webs by replacing radials or reconfiguring spirals, restoring functionality without full reconstruction [22, 23]. Damaged silk is often recycled, showcasing biological resource efficiency. Even partial damage leaves functional segments for prey detection, highlighting spiders’ adaptive intelligence [19].

3-2-5. External Force Dynamics

Environmental factors shape web mechanics:

• Wind: Alters stress distribution, prompting denser architectures [20].
• Humidity: Modifies silk elasticity and adhesion via water absorption [15].

3-2-6. Bioinspired Applications

Web adaptations inspire engineering breakthroughs:

• Smart grids: Stress-distribution models enhance resilient network designs [24].
• Self-healing materials: Mimicking web repair mechanisms [25].

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3-3. Section 3: Acoustic and Vibrational Properties

Spider webs are precision instruments for vibrational communication, enabling prey detection, localization, and mating signals. Recent studies reveal how web geometry, silk material properties, and tension tuning govern these functions.

3-3-1. Signal Transmission and Reception

Webs transmit vibrations (prey movements, wind, or mating cues) via radial threads to the hub, where specialized leg sensilla detect minute amplitude/frequency changes. Key features:

• Radial threads: High-tension pathways for rapid longitudinal wave transmission [26, 27].
• Spiral threads: Dissipate vibrations laterally, minimizing energy loss [28].
• Tuning: Adjusting thread tension filters noise (e.g., wind) or amplifies prey signals [29].

Spiders exploit time-delay differences between vibration arrival points to locate prey with mammalian-level precision [27, 30]. This system inspires metamaterials for acoustic filtering and advanced sensors [31].

3-3-2. Prey Identification and Localization

Vibrational signatures encode critical data:

• Amplitude: Correlates with prey size/weight [28].
• Frequency: Indicates movement type (e.g., mosquito wingbeats vs. struggling) [30].
• Spatial mapping: Time-delay analysis between radial threads pinpoints prey location [27].

Tension modulation allows dynamic adaptation—increasing sensitivity to small prey or dampening environmental noise [29]. Post-detection, spiders execute targeted strikes guided by vibrational triangulation [26, 28].

This natural signal-processing system, refined by evolutionary pressures, underpins biomimetic technologies in vibroacoustics and precision sensing [29, 31].

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3-4. Section 4: Bioinspiration and Technological Applications

3-4-1. Advanced Material Design and Mechanical Properties

Spider silk’s remarkable combination of strength and flexibility has inspired the development of advanced materials. These properties have been replicated in synthetic fibers for engineering applications, conductive and flexible fibers for mechanical sensors, and impact-resistant materials. Additionally, biodegradable materials inspired by spider silk have emerged as promising solutions for reducing environmental pollution. Notable examples include reinforced polymer fibers and conductive fibers for smart technologies [33, 34].

3-4-2. Printing and Manufacturing Technologies

Three-dimensional and four-dimensional printing technologies, inspired by spider web architectures, enable the production of structures capable of shape-shifting or responsive behavior to external stimuli. Techniques such as Optical Force Brush have enhanced the speed and precision of manufacturing complex structures. Applications include artificial webs for biological networks and robotics, as well as rapid prototyping of 4D structures. These methods leverage high-speed, high-precision fabrication for developing intelligent systems [35, 36].

3-4-3. Sensors and Smart Technologies

The spider’s sensory system has inspired highly sensitive vibration and microstrain detection sensors. These sensors find particular application in wearable technologies and medical devices requiring environmental adaptability. Temperature- and humidity-responsive materials have been used to develop mechanical sensors and adaptive tools. Furthermore, naturally inspired designs have led to high-precision mechanical sensors [37, 38].

3-4-4. Biomedical Applications and Tissue Engineering

Spider web structures have inspired tissue engineering technologies for bone, cartilage, and muscle regeneration. Artificial silk fibers have been employed in drug delivery systems and tissue regeneration, including intervertebral disc repair and neural systems. The biocompatibility and antimicrobial properties of these materials have been utilized in medical and hygienic applications, particularly in biological coatings and tissue scaffolds [39, 40].

3-4-5. Environmental Applications

Spider webs have inspired water-harvesting structures for arid regions. These technologies include designs for collecting atmospheric moisture from fog and air, optimized using ecological and renewable techniques. Additionally, bioinspired, eco-friendly materials derived from spider silk have been proposed as sustainable solutions for reducing environmental impact [33, 41].

3-4-6. Robotics and Defense Applications

Spider webs have served as models for designing safety nets and adaptive soft robots capable of environmental responsiveness. Impact-resistant networks mimicking spider web structures have been developed for military and engineering applications, with potential uses in protective systems and industrial/military applications [34, 42].

Spiders and their webs have inspired advanced technologies across multiple domains, including materials science, bioengineering, nanotechnology, and environmental science. These studies have introduced innovative concepts for smart materials, eco-adaptive systems, and biomedical technologies, demonstrating nature’s potential to address scientific and industrial challenges while improving quality of life.

“Spider silk, despite its strength surpassing steel and extraordinary flexibility, symbolizes the fragility of any seemingly robust structure built without reliance on divine will. This research, while uncovering the scientific secrets of this biological metamaterial, echoes the Quranic warning about the instability of civilizations dependent solely on material power.”

4. Cutting-Edge Research and Future Research Directions

Advanced research methodologies in spider silk and related behavioral studies have employed novel technologies and techniques to investigate the structure, function, and biological properties of silk and spider behaviors. Below we present the key methods and instruments utilized in these investigations:

4-1. Single-Cell and Molecular Analysis

The application of single-cell RNA sequencing (scRNA-seq) and 10x Genomics technology has enabled comprehensive gene expression analysis in spider brain neurons. In one landmark study, over 30,000 cells were classified, leading to the identification of various neuronal and non-neuronal cell types. This approach has facilitated the discovery of genes associated with learning and memory, providing unprecedented insights into the neurobiology of web-building spiders [43].

4-2. Advanced Imaging and Microscopy

Scanning electron microscopy (SEM) and transmission electron microscopy (TEM) have been instrumental in characterizing the micro- and nano-structures of spider silk and attachment discs. These techniques revealed that the diving bell spider (Argyroneta aquatica) employs superhydrophobic layers in its underwater attachment discs, offering new understanding about silk thread adhesion mechanisms in aquatic environments [44].

4-3. Genomics and Bioinformatics

Cutting-edge sequencing technologies including Oxford Nanopore and Illumina platforms have been applied to spider genome sequencing. A comprehensive study on Trichonephila clavata identified 28 complete silk-related genes and elucidated their chromosomal distribution. These genomic investigations have significantly advanced our understanding of silk production evolution and informed synthetic material design [45].

4-4. Mechanical and Dynamic Testing

Sophisticated mechanical testing systems such as Instron mechanical testers and laser Doppler vibrometers (LDV) have been employed to evaluate the tensile properties and vibrational responses of both natural and artificial spider silks. These studies demonstrated the critical influence of spiral spacing on vibrational transmission and force distribution within web structures [46].

4-5. Water-Material Interaction Studies

Investigations of water-responsive behavior in recombinant spider silk (RSF) have combined structural analysis of secondary conformations with quantitative measurements of free-to-bound water ratios. This research established that a critical ratio of bound to free water governs the mechanical behavior of silk under varying humidity conditions [47].

4-6. Interdisciplinary Approaches for Practical Applications

Bioinspired engineering approaches have successfully translated spider silk principles into practical applications, including the development of underwater adhesives and humidity-responsive materials. These interdisciplinary efforts, combining biological insights with engineering innovations, have accelerated the development of smart, silk-inspired materials [44,45].

References:

1.         Becker, N., et al., Molecular nanosprings in spider capture-silk threads. Nature materials, 2003. 2(4): p. 278-283.

2.         Cranford, S.W., et al., Nonlinear material behaviour of spider silk yields robust webs. Nature, 2012. 482(7383): p. 72-76.

3.         Chan, N.J.-A., et al., Spider-silk inspired polymeric networks by harnessing the mechanical potential of β-sheets through network guided assembly. Nature communications, 2020. 11(1): p. 1630.

4.         Wang, Q., et al., Protein secondary structure in spider silk nanofibrils. Nature Communications, 2022. 13(1): p. 4329.

5.         Nova, A., et al., Molecular and nanostructural mechanisms of deformation, strength and toughness of spider silk fibrils. Nature Precedings, 2010: p. 1-1.

6.         Grove, L., Nanofibrils study successfully measures

strength of spider web ‘super fibers’. Advanced Functional Materials, 2024.

7.         Yazawa, K., et al., Simultaneous effect of strain rate and humidity on the structure and mechanical behavior of spider silk. Communications Materials, 2020. 1(1): p. 10.

8.         Alencastre, J., C. Mago, and R. Rivera, Determination of energy dissipation of a spider silk structure under impulsive loading. Frontiers of Mechanical Engineering, 2015. 10: p. 306-310.

9.         Okumura, K., Strength and toughness of bio-fusion materials. Polymer Journal, 2015. 47(2): p. 99-105.

10.       Rech, L.P.S.E.L., Unravelling the biodiversity of nanoscale signatures

of spider silk fibres. nature communications, 2013.

11.       Jin, H.-J. and D.L. Kaplan, Mechanism of silk processing in insects and spiders. Nature, 2003. 424(6952): p. 1057-1061.

12.       Liu, Y., Z. Shao, and F. Vollrath, Relationships between supercontraction and mechanical properties of spider silk. Nature Materials, 2005. 4(12): p. 901-905.

13.       Kluge, J.A., et al., Spider silks and their applications. Trends in biotechnology, 2008. 26(5): p. 244-251.

14.       Yarger, J.L., B.R. Cherry, and A. Van Der Vaart, Uncovering the structure–function relationship in spider silk. Nature Reviews Materials, 2018. 3(3): p. 1-11.

15.       Podbevšek, D., et al., The role of water mobility on water-responsive actuation of silk. Nature Communications, 2024. 15(1): p. 8287.

16.       Haynl, C., et al., Free-standing spider silk webs of the thomisid Saccodomus formivorus are made of composites comprising micro-and submicron fibers. Scientific Reports, 2020. 10(1): p. 17624.

17.       Blamires, S.J., et al., Spider web and silk performance landscapes across nutrient space. Scientific reports, 2016. 6(1): p. 26383.

18.       Corver, A., et al., Distinct movement patterns generate stages of spider web building. Current Biology, 2021. 31(22): p. 4983-4997. e5.

19.       Greco, G., et al., Imaging and mechanical characterization of different junctions in spider orb webs. Scientific Reports, 2019. 9(1): p. 5776.

20.       Tew, N. and T. Hesselberg, The effect of wind exposure on the web characteristics of a tetragnathid orb spider. Journal of insect behavior, 2017. 30: p. 273-286.

21.       Hesselberg, T., D. Simonsen, and C. Juan, Do cave orb spiders show unique behavioural adaptations to subterranean life? A review of the evidence. Behaviour, 2019. 156(10): p. 969-996.

22.       Vollrath, F. and D.P. Knight, Liquid crystalline spinning of spider silk. Nature, 2001. 410(6828): p. 541-548.

23.       Tew, E.R., A. Adamson, and T. Hesselberg, The web repair behaviour of an orb spider. Animal Behaviour, 2015. 103: p. 137-146.

24.       Lu, W., N.A. Lee, and M.J. Buehler, Modeling and design of heterogeneous hierarchical bioinspired spider web structures using deep learning and additive manufacturing. Proceedings of the National Academy of Sciences, 2023. 120(31): p. e2305273120.

25.       Rising, A. and J. Johansson, Toward spinning artificial spider silk. Nature chemical biology, 2015. 11(5): p. 309-315.

26.       Wu, J., et al., Spider dynamics under vertical vibration and its implications for biological vibration sensing. Journal of the Royal Society Interface, 2023. 20(206): p. 20230365.

27.       Zaera, R., et al., Eco-localization of a prey in a spider orb web. Journal of Vibration and Control, 2022. 28(11-12): p. 1229-1238.

28.       Guo, C., et al., 3D-printed spider-web structures for highly efficient water collection. Heliyon, 2022. 8(8).

29.       Mortimer, B., et al., Remote monitoring of vibrational information in spider webs. The Science of Nature, 2018. 105: p. 1-9.

30.       Lott, M., et al., Prey localization in spider orb webs using modal vibration analysis. Scientific Reports, 2022. 12(1): p. 19045.

31.       Miniaci, M., et al., Spider web-inspired acoustic metamaterials. Applied Physics Letters, 2016. 109(7).

32.       Yavuz, K., et al., Effect of spider’s weight on signal transmittance in vertical orb webs. Royal Society Open Science, 2024. 11(10): p. 240986.

33.       Zheng, Y., et al., Directional water collection on wetted spider silk. Nature, 2010. 463(7281): p. 640-643.

34.       Zou, S., D. Therriault, and F.P. Gosselin, Spiderweb-inspired, transparent, impact-absorbing composite. Cell Reports Physical Science, 2020. 1(11).

35.       Li, G., et al., Bio-inspired 4D printing of dynamic spider silks. Polymers, 2022. 14(10): p. 2069.

36.       Yi, C., et al., Optical force brush enabled free-space painting of 4D functional structures. Science Advances, 2023. 9(38): p. eadg0300.

37.       Kang, D., et al., Ultrasensitive mechanical crack-based sensor inspired by the spider sensory system. Nature, 2014. 516(7530): p. 222-226.

38.       Fan, L., et al., Bioactive hierarchical silk fibers created by bioinspired self-assembly. Nature Communications, 2021. 12(1): p. 2375.

39.       Kumari, S., et al., Engineered spider silk-based 2D and 3D materials prevent microbial infestation. Materials Today, 2020. 41: p. 21-33.

40.       Salehi, S., K. Koeck, and T. Scheibel, Spider silk for tissue engineering applications. Molecules, 2020. 25(3): p. 737.

41.       Vaghasiya, J.V. and M. Pumera, The rise of 3D/4D-printed water harvesting materials. Materials Today, 2024.

42.       Lee, Y., et al., Ionic spiderwebs. Science robotics, 2020. 5(44): p. eaaz5405.

43.       Jin, P., et al., Single-cell transcriptomics reveals the brain evolution of web-building spiders. Nature Ecology & Evolution, 2023. 7(12): p. 2125-2142.

44.       Schaber, C.F., I. Grawe, and S.N. Gorb, Attachment discs of the diving bell spider Argyroneta aquatica. Communications Biology, 2023. 6(1): p. 1232.

45.       Hu, W., et al., A molecular atlas reveals the tri-sectional spinning mechanism of spider dragline silk. Nature Communications, 2023. 14(1): p. 837.

46.       Jyoti, J., et al., Structural and Vibrational Response of Artificial Spider Webs with Different Spacing. Journal of Vibration Engineering & Technologies, 2022. 10(8): p. 3101-3117.

47.       He, W., et al., Establishing superfine nanofibrils for robust polyelectrolyte artificial spider silk and powerful artificial muscles. Nature Communications, 2024. 15(1): p. 3485.