Multistranded Structures Research Articles

Multi-strand Fibers with Hierarchical Helical Structures Driven by Water or Moisture for Soft Actuators.

Smart actuators that combine excellent mechanical properties and responsive actuating performance like biological muscles have attracted considerable attention. In this study, a water/humidity responsive actuator, consisting of multi-strand carboxyl methyl cellulose (CMC) fibers with helical structures, was prepared using wet-spinning and twisting methods. The results showed that owing to the multi-strand structure, the actuator consisted of one-, two-, three-, and four-strand helical fibers, thus achieving a combination of high strength (∼27 MPa), high toughness (>10.34 MJ/m3), and large load limit (>0.30 N), which enable the actuator to theoretically withstand a weight that is at least 20,000 times its weight. Meanwhile, owing to the excellent moisture-responsive ability of CMC, the actuator, with a 5 g load, could achieve untwisting motion. Additionally, its maximum speed was approximately 2158 ± 233 rpm/m under water stimulation, whereas the recovery speed could reach 804 ± 44 rpm/m. Moreover, this untwisting-recovery reversible process was cyclic, whereas the shape and the actuating speed of the actuator remained stable after more than 150 cycles. The actuator improved the load limit that the fiber could withstand when driving under stimulation, thereby enabling the actuator to lift or move heavy objects like human muscles when executing spontaneously under external stimuli. This result shows considerable potential applications in artificial muscles and biomimetic robots.

Observational Signatures of Coronal Heating in Magnetohydrodynamic Simulations without Radiation or a Lower Atmosphere

It is extremely difficult to simulate the details of coronal heating and also make meaningful predictions of the emitted radiation. Thus, testing realistic models with observations is a major challenge. Observational signatures of coronal heating depend crucially on radiation, thermal conduction, and the exchange of mass and energy with the transition region and chromosphere below. Many magnetohydrodynamic simulation studies do not include these effects, opting instead to devote computational resources to the magnetic aspects of the problem. We have developed a simple method of accounting approximately for the missing effects. It is applied to the simulation output ex post facto and therefore may be a valuable tool for many studies. We have used it to predict the emission from a model corona that is driven by vortical boundary motions meant to represent photospheric convection. We find that individual magnetic strands experience short-term brightenings, both scattered throughout the computational volume and in localized clusters. The former may explain the diffuse component of the observed corona, while the latter may explain bright coronal loops. Several observed properties of loops are reproduced reasonably well: width, lifetime, and quasi-circular cross section (aspect ratio not high). Our results lend support to the idea that loops are multistranded structures heated by “storms” of nanoflares.

Non-canonical DNA structures: Diversity and disease association.

A complete understanding of DNA double-helical structure discovered by James Watson and Francis Crick in 1953, unveil the importance and significance of DNA. For the last seven decades, this has been a leading light in the course of the development of modern biology and biomedical science. Apart from the predominant B-form, experimental shreds of evidence have revealed the existence of a sequence-dependent structural diversity, unusual non-canonical structures like hairpin, cruciform, Z-DNA, multistranded structures such as DNA triplex, G-quadruplex, i-motif forms, etc. The diversity in the DNA structure depends on various factors such as base sequence, ions, superhelical stress, and ligands. In response to these various factors, the polymorphism of DNA regulates various genes via different processes like replication, transcription, translation, and recombination. However, altered levels of gene expression are associated with many human genetic diseases including neurological disorders and cancer. These non-B-DNA structures are expected to play a key role in determining genetic stability, DNA damage and repair etc. The present review is a modest attempt to summarize the available literature, illustrating the occurrence of non-canonical structures at the molecular level in response to the environment and interaction with ligands and proteins. This would provide an insight to understand the biological functions of these unusual DNA structures and their recognition as potential therapeutic targets for diverse genetic diseases.

Physicochemical and Anti-UVB-Induced Skin Inflammatory Properties of Lacticaseibacillus paracasei Subsp. paracasei SS-01 Strain Exopolysaccharide

The exopolysaccharide secreted by Lacticaseibacillus paracasei subsp. paracasei SS-01 strain (LP-EPS) is isolated and purified from yogurt. It is a polysaccharide with a branched and multi-stranded structure, which exists in a smooth rod-like or cloud-like state, and possesses a good thermal stability and a molecular weight of 49.68 kDa (±4.436%). LP-EPS shows a high antioxidant capacity, anti-inflammatory and anti-sensitizing activity during in vitro experimental studies, with half clearance (IC50) rates of 0.449, 1.314, and 2.369 mg/mL for the ABTS, DPPH, and OH radicals, respectively, and a half inhibition rate (IC50) of hyaluronidase of 1.53 mg/mL. A cell-based assay, enzyme-linked immunosorbent assay (ELISA), and quantitative real-time fluorescence PCR (qRT-PCR) show that LP-EPS effectively treats or ameliorates the skin inflammatory responses triggered by UVB irradiation, as evidenced by a highly significant decrease in the secretion of inflammatory factors by human skin keratinocytes (HaCaT), and a highly significant downregulation of the mRNA expression of MAPK/AP-1 pathway cytokines.

Structural characterization of phosphorylated Pleurotus ostreatus polysaccharide and its hepatoprotective effect on carbon tetrachloride-induced liver injury in mice

This study aimed to explore the basic structural features of phosphorylated Pleurotus ostreatus polysaccharide (PPOP) and study the protective effect of PPOP on liver injury induced by carbon tetrachloride in male Kunming mice. The phosphorylated polysaccharide was prepared from the natural polysaccharide extracted from Pleurotus ostreatus (POP). The structures of PPOP and POP were characterized by FT-IR, ESEM spectroscopy, and Congo red test. Chemical composition analysis revealed that PPOP was mainly composed of rhamnose, galacturonic acid, and xylose in a molar ratio of 0.10: 1.98: 1.00. Structural analysis indicated that PPOP had multi-strand structure and the absorption peaks of PO and P-O-C. Furthermore, animal experiments showed that the hepatoprotective effect of PPOP against liver injury was reflected by decreasing the levels of alanine aminotransferase, aspartate aminotransferase, alkaline phosphatase, total cholesterol, trilaurin, and low-density lipoprotein cholesterol in the serum, increasing the content of high-density lipoprotein cholesterol and albumin in blood, reducing the content of malondialdehyde and promoting the activity of antioxidant enzymes in liver. PPOP exhibited stronger hepatoprotective effect and antioxidant activity in vivo than POP. The final results indicated that PPOP could be used in the treatment of chemical-induced hepatotoxicity based on the above biological research.

Theoretical study of hydrogen bonds and electronic properties in hexagonal arrangements composed of self-assembled DNA analogues

Aminopyrimidine, triaminopyrimidine, and cyanuric acid as artificial bases can be used to design and prepare multi-stranded DNA structures. In this manuscript, we theoretically investigate the stability of hydrogen-bonded hexagonal structures arising from the self-assembly of aminopyrimidine and cyanuric acid, as well as triaminopyrimidine and cyanuric acid in gas phase and in water. The influence of hydrogen bonds on the stability of hexagonal arrangements is examined by atoms in molecules and natural bond orbital analyses. Moreover, the mutual effects of hydrogen bonds on each other are also evaluated in the hexagonal structures using cooperative energy. Whereas the self-assembly of the hydrogen-bonded hexagonal arrangements may be considered to form new nanostructures, metal sensors, and ion channels, some electronic properties such as band gap, first ionization energy, electron affinity, electronegativity, electronic chemical potential, electrophilicity index, global chemical hardness, and chemical softness are theoretically estimated at M06-2X/6-311++G(d,p) level.

Exploring potential of i-motif DNA formed in the promoter region of GRIN1 gene for nanotechnological applications

The tandem repeat stretches of cytosine residues fold to construct intercalated motif (i-motif) structures over an acidic pH range. Several advancements have been made to understand the biological role of i-motif DNA. Potential i-motif structures found in gene promoters are involved in positive transcriptional regulation and sequence specific DNA binding. Herein, a 27-mer C-rich DNA stretch present at the promoter region of (Glutamate Ionotropic Receptor NMDA type subunit 1) GRIN1 gene, named GRIN1c was studied to investigate its structural status using biophysical and biochemical techniques. The gel electrophoresis, circular dichroism spectroscopy and pH-dependent monophasic UV-thermal melting data revealed that GRIN1c sequence adapts a bimolecular i-motif structure. Two alternative structural models are suggested for the bimolecular i-motif structure with typical C·C+ base pairing and significant Watson-Crick base pairing. Possibility of co-existence of non-Watson-Crick and Watson-Crick base pairing in the formation of i-motif expands the repertoire of various topologies of i-motif structures. Extensive applications of i-motif structure in the areas of nanotechnology, analytical and biomedical systems and many more have made it essential to explore these multistranded structures. Nanodevices-based on i-motif structure found applications as biosensors and drug release platform, and also serve as proof for their in vivo existence.

RiboSketch: versatile visualization of multi-stranded RNA and DNA secondary structure.

Creating clear, visually pleasing 2D depictions of RNA and DNA strands and their interactions is important to facilitate and communicate insights related to nucleic acid structure. Here we present RiboSketch, a secondary structure image production application that enables the visualization of multistranded structures via layout algorithms, comprehensive editing capabilities, and a multitude of simulation modes. These interactive features allow RiboSketch to create publication quality diagrams for structures with a wide range of composition, size and complexity. The program may be run in any web browser without the need for installation, or as a standalone Java application. https://rnastructure.cancer.gov/ribosketch.

Transverse Wave Induced Kelvin–Helmholtz Rolls in Spicules

Abstract In addition to their jet-like dynamic behavior, spicules usually exhibit strong transverse speeds, multi-stranded structure, and heating from chromospheric to transition region temperatures. In this work we first analyze Hinode and IRIS observations of spicules and find different behaviors in terms of their Doppler velocity evolution and collective motion of their sub-structure. Some have a Doppler shift sign change that is rather fixed along the spicule axis, and lack coherence in the oscillatory motion of strand-like structure, matching rotation models, or long-wavelength torsional Alfvén waves. Others exhibit a Doppler shift sign change at maximum displacement and coherent motion of their strands, suggesting a collective magnetohydrodynamic (MHD) wave. By comparing with an idealized 3D MHD simulation combined with radiative transfer modeling, we analyze the role of transverse MHD waves and associated instabilities in spicule-like features. We find that transverse wave induced Kelvin–Helmholtz (TWIKH) rolls lead to coherence of strand-like structure in imaging and spectral maps, as seen in some observations. The rapid transverse dynamics and the density and temperature gradients at the spicule boundary lead to ring-shaped Mg ii k and Ca ii H source functions in the transverse cross-section, potentially allowing IRIS to capture the Kelvin–Helmholtz instability dynamics. Twists and currents propagate along the spicule at Alfvénic speeds, and the temperature variations within TWIKH rolls, produce the sudden appearance/disappearance of strands seen in Doppler velocity and in Ca ii H intensity. However, only a mild intensity increase in higher-temperature lines is obtained, suggesting there is an additional heating mechanism at work in spicules.

High toughness of bio-inspired multistrand coiled carbon nanotube yarn

By impressive mechanical properties, strong and lightweight carbon nanotube yarns and fibers have been reported. Among the mechanical properties of yarn, toughness is considered an important mechanical property for the protection of damage caused by external force. Here, by bio-mimicking the human muscle structure, dramatically increased toughness of carbon nanotube yarn is reported. The bio-inspired tough carbon nanotube (CNT) yarn, which is simultaneously increased in strength and elongation, is made by poly(vinyl) alcohol (PVA) infiltration and coiled multi-strand structure. This bio-inspired tough yarn provides a high-energy absorption (357.2 J/g), which is higher than spider dragline silk (165 J/g), Kevlar (78 J/g), and previous spinnable carbon nanotube yarns. Also, not only demonstrates high tough performance in various conditions such as water and temperature but also shows an ease in shape change by using water. This sewable, wearable, and shape controllable tough yarn has a potential for various applications such as bulletproof material, wearable device, stretchable electrode, aerospace industry, and artificial muscle.

Spectra and Simulation of Model Beta-Sheets and Hairpins. Impact of Turn Sequences and Aromatic Contacts on Equilibria and Dynamics

β-sheets can be modeled as antiparallel strands coupled by turns. Isolated model systems are less stable, due to amide solvation), but can be induced to fold into sheet-like structures by restricting the conformational space of turn residues, such as with DPro-Gly sequences, or by inclusion of cross-strand hydrophobic interactions, especially aromatic contacts, which tend to desolvate parts of the structure. Expanding to multistrand structures can provide different characteristics for study of physical properties. Larger structures have strand differentiation, since the center strand of a three-strand sheet has a different H-bonds than the outer ones. In larger structures, integral hairpin-like sequences of either the first two or the second two strands might have different stabilities and folding mechanisms, if it is overall a multistate process. To explore these factors, we have prepared a series of three-stranded sheet structures that are stabilized by either DPro-Gly or by Aib-Gly turns and contain various Trp-Tyr side-chain contacts. To assess the impact of the third strand we also separately synthesized the component hairpins and studied their relative properties with temperature dependent circular dichroism, fluorescence, infrared (IR) and vibrational circular dichroism for evaluation of global thermodynamic equilibria as well as IR–detected temperature jump relaxation dynamics of selected components (strands, turns). Where possible, NMR structures were obtained for comparison and MD simulations of unfolding were used to aid interpretations. While all the sequences showed evidence of β-structure formation, the extent and stability varied markedly. In terms of equilibrium properties, the turn residues (especially DPro-Gly) seemed to have the strongest influence, while the aromatic contacts had minor effects. However in terms of dynamics, the aromatic effects differentiated the structures more, suggesting a variation in the folding mechanisms.

Structure-Specific Ligand Recognition of Multistranded DNA Structures.

Structural polymorphism is an extremely significant phenomenon of nucleic acids, in which DNA and RNA oligonucleotide sequences are able to adapt various canonical, alternative and multistranded structures. These alternative forms of DNA and RNA have an enormous potential of participating in various cellular processes by recognizing ligands such as proteins, drugs and metal ions in a sequence and structure-specific manner. Such DNA-ligand interactions prove to be highly beneficial when exploited for therapeutic purposes. Many of these DNA/ RNA structures recognizing drugs have already proved their potential as anticancer, antibacterial, anthelmintic and antiviral properties. Over the last 2-3 decades, many mechanisms of DNA-drug interactions have been documented, but still many other new mechanisms are being explored. Designing new drugs with improved efficacy and specificity is of prime concern for all researchers which not only deals with the experiments related to synthesizing drugs, but also takes care of searching novel routes or agents for administration or delivery of these therapeutic agents by increasing their nuclear and cellular uptake. This review aims at explaining the structural polymorphs/ multistranded DNA structures and their interactions with pharmaceutical drugs in a structure-specific manner, along with their modes of interactions and biological relevance. This detailed overview of multistranded DNA structures and interacting drugs might further facilitate our understanding about molecular targets and drug development in a more precise manner for the larger benefit of mankind.

Multistrand Structure Prediction of Nucleic Acid Assemblies and Design of RNA Switches.

RNA is an attractive material for the creation of molecular logic gates that release programmed functionalities only in the presence of specific molecular interaction partners. Here we present HyperFold, a multistrand RNA/DNA structure prediction approach for predicting nucleic acid complexes that can contain pseudoknots. We show that HyperFold also performs competitively compared to other published folding algorithms. We performed a large variety of RNA/DNA hybrid reassociation experiments for different concentrations, DNA toehold lengths, and G+C content and find that the observed tendencies for reassociation correspond well to computational predictions. Importantly, we apply this method to the design and experimental verification of a two-stranded RNA molecular switch that upon binding to a single-stranded RNA toehold disease-marker trigger mRNA changes its conformation releasing an shRNA-like Dicer substrate structure. To demonstrate the concept, connective tissue growth factor (CTGF) mRNA and enhanced green fluorescent protein (eGFP) mRNA were chosen as trigger and target sequences, respectively. In vitro experiments confirm the formation of an RNA switch and demonstrate that the functional unit is being released when the trigger RNA interacts with the switch toehold. The designed RNA switch is shown to be functional in MDA-MB-231 breast cancer cells. Several other switches were also designed and tested. We conclude that this approach has considerable potential because, in principle, it allows the release of an siRNA designed against a gene that differs from the gene that is utilized as a biomarker for a disease state.

Engineering Gold Nanoparticles with DNA Ligands for Selective Catalytic Oxidation of Chiral Substrates

Noble metal nanoparticles are promising materials for heterogeneous enantioselective catalysis because of their high surface-to-volume ratios, large concentrations of highly undercoordinated surface sites, and quantum confinement effects. In this work, we report on the use of DNA as an environment-responsive chiral ligand to engineer the selective catalytic behaviors of glucose oxidase-mimicking gold nanoparticles (AuNPs), with glucose enantiomers as the substrates. DNA can be stimulated externally to switch between random-coiled and multistranded structures (e.g., duplex, i-motif, or G-quadruplex). Random-coiled DNA-capped nanoparticles preferentially catalyze oxidation of l-glucose, and structured DNA-capped nanoparticles show higher activity toward d-glucose. pH-induced selectivity diminishment of DNA-treated AuNPs is also found, further demonstrating the chiral selector effect of DNA ligands. In the end, the selective catalysis of AuNPs allows control of the size enlargement of nanoparticles through s...

Advancement in the Structural Polymorphism of G-Quadruplexes

Genomes contain a large number of putative guanine-rich sequences, specifically on promoter regions, untranslated regions (UTR’s) and telomeres etc. that could form guanine-quadruplexes, and may serve as important structural and regulatory elements. They can also be the source of genomic instability which may lead to cancer, aging and human genetic diseases. Four guanines in the same plane, joined together with Hoogsteen hydrogen bonding, and stacked over one another resulting in guanine tetrads, give rise to an incredible class of G-quadruplexes. An extensive range of G-quadruplex structures is well documented, where they differ in number of strands (uni, bi, or tetramolecular), conformations (parallel, antiparallel or mixed), shapes (chair or basket form), or types of loops (edgewise/ lateral, diagonal, double chain reversal/ propeller, or V-shaped loops) etc. With advancements in the techniques, various new multistranded G-rich DNA structures are enriching the DNA structural databases. The most recent ones are (3+1) G-quadruplex, Tri-G-quadruplex, G-triplex DNA etc. which actually add to the diversity of G-quadruplex structures. Exploring their polymorphism with various biophysical and biochemical techniques has hence become extremely important. This review mainly focuses on the discussion of these unusual and comparatively new polymorphic G-quadruplex DNA structures. The robustness of these unique (3+1) G-quadruplex or G-triplex or tri-G-quadruplex structures actually can be exploited for providing a strong foundation for the designing of structure-specific drugs. Recently, G-quadruplex structures have been quantitatively visualized in human cells by engineering structure-specific antibody. Considering these developments, the mapping of G-quadruplex structures in genome may now be possible, with a goal of controlling the gene functions or other cellular processes, which might be involved in diseases like cancer.

Recent Developments in the Chemistry and Biology of G-Quadruplexes with Reference to the DNA Groove Binders

DNA is the chemotherapeutic target for treating diseases of genetic origin. Besides well-known double-helical structures (A, B, Z, parallel stranded-DNA etc.), DNA is capable of forming several multi-stranded structures (triplex, tetraplex, i-motif etc.) which have unique biological significance. The G-rich 3'-ends of chromosomes, called telomeres, are synthesized by telomerase, a ribonucleoprotein, and over-expression of telomerase is associated with cancer. The activity of telomerase is suppressed if the G-rich region is folded into the four stranded structures, called G-quadruplexes (G4-DNAs) using small synthetic ligands. Thus design and synthesis of new G4- DNA ligands is an attractive strategy to combat cancer. G4-DNA forming sequences are also prevalent in other genomic regions of biological significance including promoter regions of several oncogenes. Effective gene regulation may be achieved by inducing a G4-DNA structure within the G-rich promoter sequences. To date, several G4-DNA stabilizing ligands are known. DNA groove binders interact with the duplex B-DNA through the grooves (major and minor groove) in a sequence-specific manner. Some of the groove binders are known to stabilize the G4-DNA. However, this is a relatively under explored field of research. In this review, we focus on the recent advances in the understanding of the G4-DNA structures, particularly made from the human telomeric DNA stretches. We summarize the results of various investigations of the interaction of various organic ligands with the G4-DNA while highlighting the importance of groove binder-G4-DNA interactions.

Molecular basis of coiled-coil oligomerization-state specificity.

Coiled coils are extensively and successfully used nowadays to rationally design multistranded structures for applications, including basic research, biotechnology, nanotechnology, materials science, and medicine. The wide range of applications as well as the important functions these structures play in almost all biological processes highlight the need for a detailed understanding of the factors that control coiled-coil folding and oligomerization. Here, we address the important and unresolved question why the presence of particular oligomerization-state determinants within a coiled coil does frequently not correlate with its topology. We found an unexpected, general link between coiled-coil oligomerization-state specificity and trigger sequences, elements that are indispensable for coiled-coil formation. By using the archetype coiled-coil domain of the yeast transcriptional activator GCN4 as a model system, we show that well-established trimer-specific oligomerization-state determinants switch the peptide's topology from a dimer to a trimer only when inserted into the trigger sequence. We successfully confirmed our results in two other, unrelated coiled-coil dimers, ATF1 and cortexillin-1. We furthermore show that multiple topology determinants can coexist in the same trigger sequence, revealing a delicate balance of the resulting oligomerization state by position-dependent forces. Our experimental results should significantly improve the prediction of the oligomerization state of coiled coils. They therefore should have major implications for the rational design of coiled coils and consequently many applications using these popular oligomerization domains.

RNA FRABASE 2.0: an advanced web-accessible database with the capacity to search the three-dimensional fragments within RNA structures

BackgroundRecent discoveries concerning novel functions of RNA, such as RNA interference, have contributed towards the growing importance of the field. In this respect, a deeper knowledge of complex three-dimensional RNA structures is essential to understand their new biological functions. A number of bioinformatic tools have been proposed to explore two major structural databases (PDB, NDB) in order to analyze various aspects of RNA tertiary structures. One of these tools is RNA FRABASE 1.0, the first web-accessible database with an engine for automatic search of 3D fragments within PDB-derived RNA structures. This search is based upon the user-defined RNA secondary structure pattern. In this paper, we present and discuss RNA FRABASE 2.0. This second version of the system represents a major extension of this tool in terms of providing new data and a wide spectrum of novel functionalities. An intuitionally operated web server platform enables very fast user-tailored search of three-dimensional RNA fragments, their multi-parameter conformational analysis and visualization.DescriptionRNA FRABASE 2.0 has stored information on 1565 PDB-deposited RNA structures, including all NMR models. The RNA FRABASE 2.0 search engine algorithms operate on the database of the RNA sequences and the new library of RNA secondary structures, coded in the dot-bracket format extended to hold multi-stranded structures and to cover residues whose coordinates are missing in the PDB files. The library of RNA secondary structures (and their graphics) is made available. A high level of efficiency of the 3D search has been achieved by introducing novel tools to formulate advanced searching patterns and to screen highly populated tertiary structure elements. RNA FRABASE 2.0 also stores data and conformational parameters in order to provide "on the spot" structural filters to explore the three-dimensional RNA structures. An instant visualization of the 3D RNA structures is provided. RNA FRABASE 2.0 is freely available at http://rnafrabase.cs.put.poznan.pl.ConclusionsRNA FRABASE 2.0 provides a novel database and powerful search engine which is equipped with new data and functionalities that are unavailable elsewhere. Our intention is that this advanced version of the RNA FRABASE will be of interest to all researchers working in the RNA field.

Coronal Loops: Observations and Modeling of Confined Plasma

Coronal loops are the building blocks of the X-ray bright solar corona. They owe their brightness to the dense confined plasma, and this review focuses on loops mostly as structures confining plasma. After a brief historical overview, the review is divided into two separate but not independent sections: the first illustrates the observational framework, the second reviews the theoretical knowledge. Quiescent loops and their confined plasma are considered, and therefore topics such as loop oscillations and flaring loops (except for non-solar ones which provide information on stellar loops) are not specifically addressed here. The observational section discusses loop classification and populations, and then describes the morphology of coronal loops, its relationship with the magnetic field, and the concept of loops as multi-stranded structures. The following part of this section is devoted to the characteristics of the loop plasma, and of its thermal structure in particular, according to the classification into hot, warm and cool loops. Then, temporal analyses of loops and the observations of plasma dynamics and flows are illustrated. In the modeling section starts some basics of loop physics are provided, supplying some fundamental scaling laws and timescales, a useful tool for consultation. The concept of loop modeling is introduced, and models are distinguished between those treating loops as monolithic and static, and those resolving loops into thin and dynamic strands. Then more specific discussions address modeling the loop fine structure, and the plasma flowing along the loops. Special attention is devoted to the question of loop heating, with separate discussion of wave (AC) and impulsive (DC) heating. Finally, a brief discussion about stellar X-ray emitting structures related to coronal loops is included and followed by conclusions and open questions.

The determination of the structure of Saturn’s F ring by nearby moonlets

Saturn's narrow F ring exhibits several unusual features that vary on timescales of hours to years. These include transient clumps, a central core surrounded by a multistranded structure and a regular series of longitudinal channels associated with Prometheus, one of the ring's two 'shepherding' satellites. Several smaller moonlets and clumps have been detected in the ring's immediate vicinity, and a population of embedded objects has been inferred. Here we report direct evidence of moonlets embedded in the ring's bright core, and show that most of the F ring's morphology results from the continual gravitational and collisional effects of small satellites, often combined with the perturbing effect of Prometheus. The F-ring region is perhaps the only location in the Solar System where large-scale collisional processes are occurring on an almost daily basis.