Tension Generation Research Articles

Yeast Kinesin-5 Motor Protein CIN8 Promotes Accurate Chromosome Segregation.

Accurate chromosome segregation depends on bipolar chromosome–microtubule attachment and tension generation on chromosomes. Incorrect chromosome attachment results in chromosome missegregation, which contributes to genome instability. The kinetochore is a protein complex that localizes at the centromere region of a chromosome and mediates chromosome–microtubule interaction. Incorrect chromosome attachment leads to checkpoint activation to prevent anaphase onset. Kinetochore detachment activates the spindle assembly checkpoint (SAC), while tensionless kinetochore attachment relies on both the SAC and tension checkpoint. In budding yeast Saccharomyces cerevisiae, kinesin-5 motor proteins Cin8 and Kip1 are needed to separate spindle pole bodies for spindle assembly, and deletion of CIN8 causes lethality in the absence of SAC. To study the function of Cin8 and Kip1 in chromosome segregation, we constructed an auxin-inducible degron (AID) mutant, cin8-AID. With this conditional mutant, we first confirmed that cin8-AID kip1∆ double mutants were lethal when Cin8 is depleted in the presence of auxin. These cells arrested in metaphase with unseparated spindle pole bodies and kinetochores. We further showed that the absence of either the SAC or tension checkpoint was sufficient to abolish the cell-cycle delay in cin8-AID mutants, causing chromosome missegregation and viability loss. The tension checkpoint-dependent phenotype in cells with depleted Cin8 suggests the presence of tensionless chromosome attachment. We speculate that the failed spindle pole body separation in cin8 mutants could increase the chance of tensionless syntelic chromosome attachments, which depends on functional tension checkpoint for survival.

Achieving Combined Tension-Torsion Split Hopkinson Bar test based on electromagnetic loading

This paper presents a Combined Tension-Torsion (or Compression-Torsion) Split Hopkinson Bar (T-TorSHB or C-TorSHB) system which is able to load a specimen by and torsion stress pulse synchronously under high strain rates. To achieve the precision control of synchronous loading, electromagnetic energy conversion technique is adopted. A new Electromagnetic-release Split Hopkinson Torsion Bar system (E-SHTorB) is proposed to generate the torsion stress pulse, while the tension stress pulse is generated by the technique we previously proposed. A trapezoidal torsion stress pulse with rising time of 200μs is acquired and repeated tests show very good consistency in generating time and pulse shape. A synchronous system is designed to control the trigger time of the tension and torsion stress pulse generator. Repetitive experiments are performed and the results prove that the time difference of stress pulses arriving at the specimen between tension and torsion pulses is less than 5μs. The proposed T-TorSHB system is verified by tests on specimens made by T2 copper.

Genotype-Driven Pathogenesis of Atrial Fibrillation in Hypertrophic Cardiomyopathy: The Case of Different TNNT2 Mutations.

Atrial dilation and atrial fibrillation (AF) are common in Hypertrophic CardioMyopathy (HCM) patients and associated with a worsening of prognosis. The pathogenesis of atrial myopathy in HCM remains poorly investigated and no specific association with genotype has been identified. By re-analysis of our cohort of thin-filament HCM patients (Coppini et al. 2014) AF was identified in 10% of patients with sporadic mutations in the cardiac Troponin T gene (TNNT2), while AF occurrence was much higher (25–75%) in patients carrying specific “hot-spot” TNNT2 mutations. To determine the molecular basis of arrhythmia occurrence, two HCM mouse models expressing human TNNT2 variants (a “hot-spot” one, R92Q, and a “sporadic” one, E163R) were selected according to the different pathophysiological pathways previously demonstrated in ventricular tissue. Echocardiography studies showed a significant left atrial dilation in both models, but more pronounced in the R92Q. In E163R atrial trabeculae, in line with what previously observed in ventricular preparations, the energy cost of tension generation was markedly increased. However, no changes of twitch amplitude and kinetics were observed, and there was no atrial arrhythmic propensity. R92Q atrial trabeculae, instead, displayed normal ATP consumption but markedly increased myofilament calcium sensitivity, as previously observed in ventricular preparations. This was associated with reduced inotropic reserve and slower kinetics of twitch contractions and, importantly, with an increased occurrence of spontaneous beats and triggered contractions that represent an intrinsic arrhythmogenic mechanism promoting AF. The association of specific TNNT2 mutations with AF occurrence depends on the mutation-driven pathomechanism (i.e., increased atrial myofilament calcium sensitivity rather than increased myofilament tension cost) and may influence the individual response to treatment.

Brane world creation from flat or almost flat space in dynamical tension string theories

There is great interest in the construction of brane worlds, where matter and gravity are forced to be effective only in a lower dimensional surface, the brane . How these could appear as a consequence of string theory is a crucial question and this has been widely discussed. Here we will examine a distinct scenario that appears in dynamical tension theories and where string tension is positive between two surfaces separated by a short distance and at the two surfaces themselves the string tensions become infinite, therefore producing an effective confinement of the strings and therefore of all matter and gravity to the space between these to surfaces, which is in fact a new type of stringy brane world scenario. The basic formulation for obtaining this scenario consist of assuming two types of strings characterized by a different constant of integration related to the spontaneous string tension generation. These string tension multiplied by the embedding metric define conformally related metrics that both satisfy Einsteins equation. The braneworlds appear very naturally when these two metrics are both flat spaces related by a special conformal transformation. The two types of string tensions are determined and they blow up at two close expanding surfaces. A puzzling aspect appears then: the construction is based on flat spaces, but then there are also strings with very large tension near the boundaries of the braneworld,so can the back reaction from the infinite tension strings destroy the flat space background? Fortunatelly that can be resolved using the mechanism Universe creation from almost flat (or empty) spaces, which incorporates a gas of very large string tensions in a membrane, studied before in 1+1 membranes in a 2+1 embedding space and now is generalized for a 1+(D-2) membrane moving in a 1+(D-1) space.

Functional and structural differences between skinned and intact muscle preparations.

Myofilaments and their associated proteins, which together constitute the sarcomeres, provide the molecular-level basis for contractile function in all muscle types. In intact muscle, sarcomere-level contraction is strongly coupled to other cellular subsystems, in particular the sarcolemmal membrane. Skinned muscle preparations (where the sarcolemma has been removed or permeabilized) are an experimental system designed to probe contractile mechanisms independently of the sarcolemma. Over the last few decades, experiments performed using permeabilized preparations have been invaluable for clarifying the understanding of contractile mechanisms in both skeletal and cardiac muscle. Today, the technique is increasingly harnessed for preclinical and/or pharmacological studies that seek to understand how interventions will impact intact muscle contraction. In this context, intrinsic functional and structural differences between skinned and intact muscle pose a major interpretational challenge. This review first surveys measurements that highlight these differences in terms of the sarcomere structure, passive and active tension generation, and calcium dependence. We then highlight the main practical challenges and caveats faced by experimentalists seeking to emulate the physiological conditions of intact muscle. Gaining an awareness of these complexities is essential for putting experiments in due perspective.

Multimedia Recognition of Piano Music Based on the Hidden Markov Model

Piano performance is an art with rich artistic elements and unpredictable performance skills. It is an important carrier for playing beautiful piano sounds. The generation of musical tension and expression of piano performance is a vivid display of piano performance skills. In piano performance, we should pay attention to the cultivation and flexible application of performance skills. In order to ensure the richness and artistry of piano performance, it is fully based on the artistic characteristics of piano performance. Through in-depth analysis of the principle of the hidden Markov model, it is applied to the multimedia recognition process of piano playing music. In the process of obtaining the template, the fundamental frequency of the piano playing music differs greatly, and the piano playing music appears during the performance process. For the problem of low recognition rate, this paper proposes a multimedia recognition method for piano music. Finally, the analysis of experimental results shows that the method proposed in this paper has a 16% higher recognition rate than the traditional method, and it has a certain value in the multimedia recognition of piano music.

Abstract 13285: Sensitivity to Beta-Adrenergic Stimulation in Pediatric Failing Single Ventricle Myocardium

Introduction: Children with heart failure (HF) in the context of palliated single ventricle congenital heart disease (SV) are treated with various pharmacotherapies, all with limited efficacy. Specifically, outcomes with β-blockade for HF in SV children are disparate from those in adult HF, even with similar downregulation of β 1 -adrenergic receptors (AR) in both populations. Hypothesis: Downregulation of β 1 -AR in pediatric SV subjects results in a unique contractile phenotype. Methods: Hearts were obtained from SV children who underwent transplant between 2010-2020. Non-failing SV (SV-NF) subjects underwent primary transplantation and had no signs of HF at the time of transplant. Failing SV (SV-F) subjects required transplantation due to end-stage HF (including refractory protein losing enteropathy or plastic bronchitis). At the time of transplant, trabeculae are isolated from the right ventricle of the explanted SV heart and mounted in a multi-chamber muscle bath. Ex vivo contractile response of isolated trabeculae is measured in response to the non-selective β-agonist isoproterenol; only trabeculae with an appropriate isoproterenol response are included in analyses. Results: Although β 1 -AR density is decreased in both SV-NF and SV-F myocardium, trabecular contractility assessment demonstrates increased sensitivity to isoproterenol in SV-F compared to SV-NF. Maximal tension generation with isoproterenol is similar between SV-NF and SV-F. Isoproterenol sensitivity in SV-F is comparable to that of non-failing adult hearts (age <40, n=4). Conclusions: Pediatric SV-F trabeculae demonstrate increased sensitivity to β-adrenergic stimulation when compared to SV-NF, but is similar to that of non-failing adult hearts. The preservation of adrenergic responsiveness may suggest that chronic, adverse β 1 -AR signaling is not part of the pathophysiology of failing SV, which may limit the effectiveness of β-blockade for HF indications in this population.

Myopalladin knockout mice develop cardiac dilation and show a maladaptive response to mechanical pressure overload.

Myopalladin (MYPN) is a striated muscle-specific immunoglobulin domain-containing protein located in the sarcomeric Z-line and I-band. MYPN gene mutations are causative for dilated (DCM), hypertrophic, and restrictive cardiomyopathy. In a yeast two-hybrid screening, MYPN was found to bind to titin in the Z-line, which was confirmed by microscale thermophoresis. Cardiac analyses of MYPN knockout (MKO) mice showed the development of mild cardiac dilation and systolic dysfunction, associated with decreased myofibrillar isometric tension generation and increased resting tension at longer sarcomere lengths. MKO mice exhibited a normal hypertrophic response to transaortic constriction (TAC), but rapidly developed severe cardiac dilation and systolic dysfunction, associated with fibrosis, increased fetal gene expression, higher intercalated disc fold amplitude, decreased calsequestrin-2 protein levels, and increased desmoplakin and SORBS2 protein levels. Cardiomyocyte analyses showed delayed Ca2+ release and reuptake in unstressed MKO mice as well as reduced Ca2+ spark amplitude post-TAC, suggesting that altered Ca2+ handling may contribute to the development of DCM in MKO mice.

Microarray profiling of gene expression in C2C12 myotubes trained by electric pulse stimulation

Electric pulse-stimulated C2C12 myotubes are gaining interest in the field of muscle physiology and biotechnology because electric pulse stimulation (EPS) enhances sarcomere structure development and active tension generation capability. Recently, we found that termination of EPS results in the rapid loss of active tension generation accompanied by disassembly of the sarcomere structure, which may represent an invitro muscle atrophy model. To elucidate the molecular mechanism underlying this rapid loss of active tension generation and sarcomere structure disassembly after termination of EPS, we performed transcriptomic analysis using microarray. After termination of EPS, 74 genes were upregulated and 120 genes were downregulated after 30min; however, atrophy-related genes were not found among these genes. To further assess the effect of EPS on gene expression, we re-applied EPS after its termination for 8h and searched for genes whose expression was reversed. Four genes were upregulated by termination of EPS and downregulated by the re-application of EPS, whereas two genes were downregulated by termination of EPS and upregulated by the re-application of EPS. Although none of these genes were atrophy- or hypertrophy-related, the results presented in this study will contribute to the understanding of gene expression changes that mediate rapid loss of active tension generation and sarcomere structure disassembly following termination of EPS in C2C12 myotubes.

Electro-Mechanical Whole-Heart Digital Twins: A Fully Coupled Multi-Physics Approach

Mathematical models of the human heart are evolving to become a cornerstone of precision medicine and support clinical decision making by providing a powerful tool to understand the mechanisms underlying pathophysiological conditions. In this study, we present a detailed mathematical description of a fully coupled multi-scale model of the human heart, including electrophysiology, mechanics, and a closed-loop model of circulation. State-of-the-art models based on human physiology are used to describe membrane kinetics, excitation-contraction coupling and active tension generation in the atria and the ventricles. Furthermore, we highlight ways to adapt this framework to patient specific measurements to build digital twins. The validity of the model is demonstrated through simulations on a personalized whole heart geometry based on magnetic resonance imaging data of a healthy volunteer. Additionally, the fully coupled model was employed to evaluate the effects of a typical atrial ablation scar on the cardiovascular system. With this work, we provide an adaptable multi-scale model that allows a comprehensive personalization from ion channels to the organ level enabling digital twin modeling.

Mechanical plasticity of collagen directs branch elongation in human mammary gland organoids

Epithelial branch elongation is a central developmental process during branching morphogenesis in diverse organs. This fundamental growth process into large arborized epithelial networks is accompanied by structural reorganization of the surrounding extracellular matrix (ECM), well beyond its mechanical linear response regime. Here, we report that epithelial ductal elongation within human mammary organoid branches relies on the non-linear and plastic mechanical response of the surrounding collagen. Specifically, we demonstrate that collective back-and-forth motion of cells within the branches generates tension that is strong enough to induce a plastic reorganization of the surrounding collagen network which results in the formation of mechanically stable collagen cages. Such matrix encasing in turn directs further tension generation, branch outgrowth and plastic deformation of the matrix. The identified mechanical tension equilibrium sets a framework to understand how mechanical cues can direct ductal branch elongation.

Tension management in cable transmission systems for remote manipulators

This paper presents the design of a novel kinematic tensioning system to remedy the situation where the input-output relation in the cable transmission for a remotely operated multi-link end-effector is compromised due to the onset of slack. The actuation of multi-link end-effectors in remote manipulators via cable transmission exhibits loss of tension and slack generation on the non-driving side of the cable transmission. This paper outlines an analytical formulation of the kinematic component of slack resulting from the geometry of the end-effector links that are serially connected. If left unaddressed, cable slack adversely impacts transmission stiffness and produces backlash at the end-effector wherein for a fixed input, the output can move freely over a range of motion (i.e. dead-band). In case of articulating instruments for minimally invasive surgery, this limits the ability to position and orient the end-effector precisely, the ability to effectively transmit forces via the end-effectors to the surrounding tissues, and the tactile feedback to the user operating the instrument. To overcome these limitations, this paper presents a simple solution that effectively mitigates slack on the non-driving side of a cable transmission between a driving pulley and a multi-link end-effector with minimal cost or complexity. The proposed design uses a driving pulley modified with tensioner extensions to alter the transmission path, which provides a kinematic or geometric solution in contrast to the various spring based solutions that have several drawbacks. Based on a derivation of slack in the system, a design optimization is performed to generate the key dimensions of the tensioning system that minimizes slack without causing over-constraint. An experimental setup is designed and fabricated to validate the slack predictions in the cable transmission system for a multi-link end-effector, without and with the proposed tensioner. The experimental measurements demonstrate the efficacy of the proposed solution.

Mechanosignaling pathways alter muscle structure and function by post-translational modification of existing sarcomeric proteins to optimize energy usage.

A transduced mechanical signal arriving at its destination in muscle alters sarcomeric structure and function. A major question addressed is how muscle mass and tension generation are optimized to match actual performance demands so that little energy is wasted. Three cases for improved energy efficiency are examined: the troponin complex for tuning force production, control of the myosin heads in a resting state, and the Z-disc proteins for sarcomere assembly. On arrival, the regulation of protein complexes is often controlled by post-translational modification (PTM), of which the most common are phosphorylation by kinases, deacetylation by histone deacetylases and ubiquitination by E3 ligases. Another branch of signals acts not through peptide covalent bonding but via ligand interactions (e.g. Ca2+ and phosphoinositide binding). The myosin head and the regulation of its binding to actin by the troponin complex is the best and earliest example of signal destinations that modify myofibrillar contractility. PTMs in the troponin complex regulate both the efficiency of the contractile function to match physiologic demand for work, and muscle mass via protein degradation. The regulation of sarcomere assembly by integration of incoming signaling pathways causing the same PTMs or ligand binding are discussed in response to mechanical loading and unloading by the Z-disc proteins CapZ, α-actinin, telethonin, titin N-termini, and others. Many human mutations that lead to cardiomyopathy and heart disease occur in the proteins discussed above, which often occur at their PTM or ligand binding sites.

Measuring Single-Cell Platelet Forces via Microcontact-Printed, Reference-Free Traction Force Microscopy Reveals Relationships between Cell Shape, F-Actin Localization, and Force

Platelets bind, spread, and contract to stop bleeding and reinforce hemostatic plugs. Platelet contraction occurs when interactions between actin and myosin create cytoskeletal tension and force generation. In this study, single-cell platelet forces were measured using a novel microcontact-printed, reference-free traction force microscopy approach. This approach facilitates single-cell force measurement on a contiguous topology that enables unrestricted platelet spreading without necessitating platelet removal to calculate force. Microcontact printing is used to deposit a grid of fluorescent bovine serum albumin onto a flexible polydimethylsiloxane substrate. Platelets from healthy donors were washed and seeded onto von Willebrand Factor-coated substrates. Platelets were fixed, stained, and imaged via confocal microscopy to simultaneously visualize fluorescently labeled platelets and grid displacement (used to calculate single-cell force.) Because microcontact-printing facilitates printing the grid on large, scalable surfaces, platelet forces were able to be measured on >100 single-cells per donor (N = 6 donors). Donor-to-donor platelet force variability was examined, and some donors produce significantly more force per platelet than other donors (p<0.01). Even within each donor, ∼15-fold variability in single platelet forces exists. A well-established area-force relationship (more spread cells produce more force) accounts for some variability in force generation, but for cells spread equally, there is still ∼5-fold differences in force generation. After examining the relationship between cell shape and force, a relationship was observed wherein platelets that are more circular produce more force. Additionally, amongst spread platelets, platelets with more spread F-actin produce more force. This versatile method that facilitates measurement of single-cell forces over large areas without restricting cell spreading or requiring cell removal reveals donor variability, cell shape, and F-actin localization as contributors to the large variability of single-platelet forces.

Computational modelling of membrane gating in capsule translocation through microchannel with variable section

As an effective physical method, cell squeezing technology based on microfluidics plays an increasingly promising role in intracellular delivery. To deepen our understanding of microfluidic chip design and optimization, it is essential to explore the underlying physics required in the generation of the appropriate tension for gating capsule membrane. In this investigation, an immersed finite element method (IFEM) has been adopted to simulate the interaction between capsule and fluid fields in microchannel with variable section. Having obtained the numerical results, the gating region on the membrane can be determined by the non-uniform tension distribution based on the critical gating membrane tension during the capsule translocates the microchannel. In addition, the gating integral, which is defined to measure the degree of gating, shows that the setting of driven pressure might be crucial for the design and optimization of the system. The numerical results demonstrate that the occurrence of secondary peak in deformed energy after the capsule passes through the confined microchannel under certain flow condition might be ascribed to the elastic recovery of the membrane. To investigate the effects of initial orientations, the translocation of an ellipsoid capsule through the channel has also been simulated numerically, which indicates that both flow shear force and compressive force due to the constrained solid wall have significant effects on membrane gating. Therefore, to improve the gating efficiency of capsule membrane, it is necessary to optimize various factors to achieve the balance among the compressive force, shear force and the translocation time.

Effect of hydrogen peroxide concentration on the maintenance and differentiation of cultured skeletal muscle cells

We have studied the effects of hydrogen peroxide (H2O2) on the differentiation and maintenance of C2C12 myoblasts. The effects of H2O2 were evaluated by cell viability, total protein concentration, the relative amount of muscle-related proteins, sarcomere structure, and active tension generation. Oxidative stress is one of the major causes of myopathy after exercise and thus establishing the method to evaluate the effects on muscle function is essential. The primary function of striated muscle is to generate force, thus, the measurement of active tension is important in assessing the effect of chemicals on muscle. Among the indices we tested, the sarcomere structure was the most sensitive to the H2O2 exposure while the cell viability was less sensitive. The effects of H2O2 on active tension correlated with a decrease in the amount of muscle proteins. In this study, our results showed that the effect of chemicals on muscle should be measured in multiple ways, including active tension generation, for a better understanding of its physiological impact.

A Perspective on Personalized Therapies in Hypertrophic Cardiomyopathy.

A dominant mechanism of sudden cardiac death in the young is the progression of maladaptive responses to genes encoding proteins linked to hypertrophic cardiomyopathy. Most are mutant sarcomere proteins that trigger the progression by imposing a biophysical defect on the dynamics and levels of myofilament tension generation. We discuss approaches for personalized treatments that are indicated by recent advanced understanding of the progression.

Evaluation of Strain Transition Properties between Cast-In Fibre Bragg Gratings and Cast Aluminium during Uniaxial Straining.

Current testing methods are capable of measuring strain near the surface on structural parts, for example by using strain gauges. However, stress peaks often occur within the material and can only be approximated. An alternative strain measurement incorporates fibre-optical strain sensors (Fiber Bragg Gratings, FBG) which are able to determine strains within the material. The principle has already been verified by using embedded FBGs in tensile specimens. The transition area between fibre and aluminium, however, is not yet properly investigated. Therefore, strains in tensile specimens containing FBGs were measured by neutron diffraction in gauge volumes of two different sizes around the Bragg grating. As a result, it is possible to identify and decouple elastic and plastic strains affecting the FBGs and to transfer the findings into a fully descriptive FE-model of the strain transition area.We thus accomplished closing the gap between the external load and internal straining obtained from cast-in FBG and generating valuable information about the mechanisms within the strain transition area.It was found that the porosity within the casting has a significant impact on the stiffness of the tensile specimen, the generation of excess microscopic tensions and thus the formation of permanent plastic strains, which are well recognized by the FBG. The knowledge that FBG as internal strain sensors function just as well as common external strain sensors will now allow for the application of FBG in actual structural parts and measurements under real load conditions. In the future, applications for long-term monitoring of cast parts will also be enabled and are currently under development.

Animal Cell Cytokinesis: The Rho-Dependent Actomyosin-Anilloseptin Contractile Ring as a Membrane Microdomain Gathering, Compressing, and Sorting Machine.

Cytokinesis is the last step of cell division that partitions the cellular organelles and cytoplasm of one cell into two. In animal cells, cytokinesis requires Rho-GTPase-dependent assembly of F-actin and myosin II (actomyosin) to form an equatorial contractile ring (CR) that bisects the cell. Despite 50 years of research, the precise mechanisms of CR assembly, tension generation and closure remain elusive. This hypothesis article considers a holistic view of the CR that, in addition to actomyosin, includes another Rho-dependent cytoskeletal sub-network containing the scaffold protein, Anillin, and septin filaments (collectively termed anillo-septin). We synthesize evidence from our prior work in Drosophila S2 cells that actomyosin and anillo-septin form separable networks that are independently anchored to the plasma membrane. This latter realization leads to a simple conceptual model in which CR assembly and closure depend upon the micro-management of the membrane microdomains to which actomyosin and anillo-septin sub-networks are attached. During CR assembly, actomyosin contractility gathers and compresses its underlying membrane microdomain attachment sites. These microdomains resist this compression, which builds tension. During CR closure, membrane microdomains are transferred from the actomyosin sub-network to the anillo-septin sub-network, with which they flow out of the CR as it advances. This relative outflow of membrane microdomains regulates tension, reduces the circumference of the CR and promotes actomyosin disassembly all at the same time. According to this hypothesis, the metazoan CR can be viewed as a membrane microdomain gathering, compressing and sorting machine that intrinsically buffers its own tension through coordination of actomyosin contractility and anillo-septin-membrane relative outflow, all controlled by Rho. Central to this model is the abandonment of the dogmatic view that the plasma membrane is always readily deformable by the underlying cytoskeleton. Rather, the membrane resists compression to build tension. The notion that the CR might generate tension through resistance to compression of its own membrane microdomain attachment sites, can account for numerous otherwise puzzling observations and warrants further investigation using multiple systems and methods.

Musical Tension Associated With Violations of Hierarchical Structure.

Tension is one of the core principles of emotion evoked by music, linking objective musical events and subjective experience. The present study used continuous behavioral rating and electroencephalography (EEG) to investigate the dynamic process of tension generation and its underlying neurocognitive mechanisms; specifically, tension induced by structural violations at different music hierarchical levels. In the experiment, twenty-four musicians were required to rate felt tension continuously in real-time, while listening to music sequences with either well-formed structure, phrase violations, or period violations. The behavioral data showed that structural violations gave rise to increasing and accumulating tension experience as the music unfolded; tension was increased dramatically by structural violations. Correspondingly, structural violations elicited N5 at GFP peaks, and induced decreasing neural oscillations power in the alpha frequency band (8–13 Hz). Furthermore, compared to phrase violations, period violations elicited larger N5 and induced a longer-lasting decrease of power in the alpha band, suggesting a hierarchical manner of musical processing. These results demonstrate the important role of musical structure in the generation of the experience of tension, providing support to the dynamic view of musical emotion and the hierarchical manner of tension processing.