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Episodes of low-dimensional self-organized dynamics from electroencephalographic alpha-signals.

Self-organized neuronal dynamics revealed by cortical alpha-rhythms occur as episodes, which are rarely observed without extraction of the alpha-band from the other spectral components. Three episodes of an unusually long duration of 10 s, two with no signal processing after data recording at the clinic, are described and show evidence of low-dimensional alpha-dynamics. The evidence is gained from an analysis of scaled structures appearing in families of slope curves of the correlation integrals and is checked against time reparametrization. The data for the two unprocessed 10-s episodes are used for a test of the methodology, as well as a re-examination of the adequacy of the model of an autonomous dynamic system in steady state and of the concept of an attractor in brain dynamics investigations. Striking evidence for the model's inadequacy is provided by the episode of subject S1. In this example five consecutive overlapping 6-s sections do show evidence for low-dimensional dynamics, whereas the 10-s section containing those sections does not. The episode of subject 1 provides an example of alpha-activity which may involve self-organized dynamics extending down to low frequencies. The system (the neuronal network) showing episodes of attractor-ruled dynamics, under conditions of blurred and smoothly fading out evidence that it stays on an attractor, is designated as being ruled by a 'shadow-attractor'. This concept is compared with that of a 'quasi-attractor' introduced by H. Haken in studies of physiological systems. One possible mechanism for the observed episodes is proposed, based on a time-dependent number of enslaved sub-systems.

Adult↗

Cortical memory dynamics.

Biological memories have a number of unique features, including (1) hierarchical, reciprocally interacting layers, (2) lateral inhibitory interactions within layers, and (3) Hebbian synaptic modifications. We incorporate these key features into a mathematical and computational model in which we derive and study Hebbian learning dynamics and recall dynamics. Introducing the construct of a feasible memory (a memory that formally responds correctly to a specified collection of noisy cues that are known in advance), we study stability and convergence of the two kinds of dynamics by both analytical and computational methods. A conservation law for memory feasibility under Hebbian dynamics is derived. An infomax net is one where the synaptic weights resolve the most uncertainty about a neural input based on knowledge of the output. The infomax notion is described and is used to grade memories and memory performance. We characterize the recall dynamics of the most favorable solutions from an infomax perspective. This characterization includes the dynamical behavior when the net is presented with external stimuli (noisy cues) and a description of the accuracy of recall. The observed richness of dynamical behavior, such as its initial state sensitivity, provides some hints for possible biological parallels to this model.

Animals↗

Modeling observed chaotic oscillations in bursting neurons: the role of calcium dynamics and IP3.

Chaotic bursting has been recorded in synaptically isolated neurons of the pyloric central pattern generating (CPG) circuit in the lobster stomatogastric ganglion. Conductance-based models of pyloric neurons typically fail to reproduce the observed irregular behavior in either voltage time series or state-space trajectories. Recent suggestions of Chay [Biol Cybern 75: 419-431] indicate that chaotic bursting patterns can be generated by model neurons that couple membrane currents to the nonlinear dynamics of intracellular calcium storage and release. Accordingly, we have built a two-compartment model of a pyloric CPG neuron incorporating previously described membrane conductances together with intracellular Ca2+ dynamics involving the endoplasmic reticulum and the inositol 1,4,5-trisphosphate receptor IP3R. As judged by qualitative inspection and quantitative, nonlinear analysis, the irregular voltage oscillations of the model neuron resemble those seen in the biological neurons. Chaotic bursting arises from the interaction of fast membrane voltage dynamics with slower intracellular Ca2+ dynamics and, hence, depends on the concentration of IP3. Despite the presence of 12 independent dynamical variables, the model neuron bursts chaotically in a subspace characterized by 3-4 active degrees of freedom. The critical aspect of this model is that chaotic oscillations arise when membrane voltage processes are coupled to another slow dynamic. Here we suggest this slow dynamic to be intracellular Ca2+ handling.

Action Potentials↗

Understanding wheel dynamics.

In five experiments, assessments were made of people's understandings about the dynamics of wheels. It was found that undergraduates make highly erroneous dynamical judgments about the motions of this commonplace event, both in explicit problem-solving contexts and when viewing ongoing events. These problems were also presented to bicycle racers and high-school physics teachers; both groups were found to exhibit misunderstandings similar to those of naive undergraduates. Findings were related to our account of dynamical event complexity. The essence of this account is that people encounter difficulties when evaluating the dynamics of any mechanical system that has more than one dynamically relevant object parameter. A rotating wheel is multidimensional in this respect: in addition to the motion of its center of mass, its mass distribution is also of dynamical relevance. People do not spontaneously form the essential multidimensional quantities required to adequately evaluate wheel dynamics.

Adult↗

Solvent effects on protein motion and protein effects on solvent motion. Dynamics of the active site region of lysozyme.

The stochastic boundary molecular dynamics methodology is applied to the active site of the enzyme lysozyme. A comparison is made of in vacuo dynamics results from the stochastic boundary method and a full conventional molecular dynamics simulation of lysozyme. Excellent agreement between the two approaches is obtained. The influence of solvent on the residues in the active site region is explored and it is shown that both the structure and dynamics are affected. Of particular importance for the structure of the protein is the solvation of polar residues and the stabilization of like-charged ion pairs. The magnitude of the fluctuations is only slightly altered by the solvent; the overall increase in the root-mean-square fluctuations, relative to the vacuum run, is 11%. The solvent effect on dynamical properties is found not to be simply related to the solvent viscosity. Both the solvent exposure and dynamic aspects of protein-solvent interactions, including the relative time scales of the motions, are shown to play a role. The effects of the protein on solvent dynamics and structure are also observed to be significant. The solvent molecules around atoms in charged, polar and apolar side-chains show markedly different diffusion coefficients as well as exhibiting different solvation structures. One key example is the water around apolar groups, which is much less mobile than bulk water, or water solvating polar groups.

Animals↗

Dynamics of brain extracellular fluid pH and phrenic nerve activity in cats after end-tidal CO2 forcing.

Ventilation is influenced by the interstitial [H+] of the brain. The pHecf, which in turn is determined largely by ventilation (via PaCO2) is sensed by the central chemoreceptors. In order to investigate the dynamics of both pHecf and neural tidal volume, we measured in cats with cut vagi and sinus nerves the dynamic medullary pHecf changes and the associated changes in integrated phrenic nerve activity after end-tidal CO2 forcing. The medullary surface ecf pH was measured with a glass electrode with a flat pH-sensitive surface. After CO2 up-steps, the pHecf changed with a time constant of about 43 sec, after down-steps 30 sec was found. The central time constant of the neural tidal volume response was 50 sec (mode) in both cases, whereas the overall response had a (modal) time constant of 80 sec. The results indicate that pHecf dynamics and the dynamic characteristics of the central neural respiratory organization are about equally important in determining the dynamic neural tidal volume response. It is argued that when PaCO2 changes, the dynamic pHecf change is perfusion limited and macroscopically homogeneous within the brainstem. Therefore, in our view it seems that the location of the central chemoreceptors within the brainstem is of minor importance in determining the dynamic neural tidal volume response to PaCO2 changes.

Animals↗

Gender- and age-related differences in heart rate dynamics: are women more complex than men?

OBJECTIVES: This study aimed to quantify the complex dynamics of beat-to-beat sinus rhythm heart rate fluctuations and to determine their differences as a function of gender and age. BACKGROUND: Recently, measures of heart rate variability and the nonlinear "complexity" of heart rate dynamics have been used as indicators of cardiovascular health. Because women have lower cardiovascular risk and greater longevity than men, we postulated that there are important gender-related differences in beat-to-beat heart rate dynamics. METHODS: We analyzed heart rate dynamics during 8-min segments of continuous electrocardiographic recording in healthy young (20 to 39 years old), middle-aged (40 to 64 years old) and elderly (65 to 90 years old) men (n = 40) and women (n = 27) while they performed spontaneous and metronomic (15 breaths/min) breathing. Relatively high (0.15 to 0.40 Hz) and low (0.01 to 0.15 Hz) frequency components of heart rate variability were computed using spectral analysis. The overall "complexity" of each heart rate time series was quantified by its approximate entropy, a measure of regularity derived from nonlinear dynamics ("chaos" theory). RESULTS: Mean heart rate did not differ between the age groups or genders. High frequency heart rate power and the high/low frequency power ratio decreased with age in both men and women (p < 0.05). The high/low frequency power ratio during spontaneous and metronomic breathing was greater in women than men (p < 0.05). Heart rate approximate entropy decreased with age and was higher in women than men (p < 0.05). CONCLUSIONS: High frequency heart rate spectral power (associated with parasympathetic activity) and the overall complexity of heart rate dynamics are higher in women than men. These complementary findings indicate the need to account for gender-as well as age-related differences in heart rate dynamics. Whether these gender differences are related to lower cardiovascular disease risk and greater longevity in women requires further study.

Adult↗

Studies of alphaB crystallin subunit dynamics by surface plasmon resonance.

The molecular chaperone activity of alphaB crystallin, an important stress protein in humans, is regulated by physiological factors, including temperature, pH, Ca2+, and ATP. In this study, the role of these factors in regulating the subunit dynamics of human alphaB crystallin was investigated using surface plasmon resonance (SPR). SPR experiments indicate that at temperatures above 37 degrees C, where alphaB crystallin has been reported to have higher chaperone activity, the subunit dynamics of alphaB crystallin were increased with faster association and dissociation rates. SPR experiments also indicate that interactions between alphaB crystallin subunits were enhanced with much faster association and slower dissociation rates at pH values below 7.0, where alphaB crystallin has been reported to have lower chaperone activity. The results suggest that the dynamic and rapid subunit exchange rate may regulate the chaperone activity of alphaB crystallin. The effect of Ca2+ and ATP on the subunit dynamics of alphaB crystallin was minimal, suggesting that Ca2+ and ATP modulate the chaperone activity of alphaB crystallin without altering the subunit dynamics. Based on the SPR results and previously reported biochemical data for the chaperone activity of alphaB crystallin under different conditions of temperature and pH, a model for the relationship between the subunit dynamics and chaperone activity of alphaB crystallin is established. The model is consistent with previous biochemical data for the chaperone activity and subunit dynamics of small heat shock proteins (sHSPs) and establishes a working hypothesis for the relationship between complex assembly and chaperone activity for sHSPs.

Adenosine Triphosphate↗

Usefulness of ventricular repolarization dynamicity in predicting arrhythmic deaths in patients with ischemic cardiomyopathy (from the European Myocardial Infarct Amiodarone Trial).

The European Myocardial Infarct Amiodarone Trial (EMIAT) investigated the effects of amiodarone versus placebo in patients after myocardial infarction who had left ventricular ejection fraction </=40% and were </=75 years of age. The present substudy examined whether ventricular repolarization (VR) dynamicity could differentiate patients who died from cardiac death from their matched survivors in this EMIAT population. In addition, we assessed whether VR dynamicity could differentiate patients who died from arrhythmic cardiac death (ACD) and from non-ACD. VR dynamicity (determined from Holter's recordings at baseline) was compared before antiarrhythmic therapy in 118 patients who had cardiac death and 118 matched survivors according to age, gender, left ventricular ejection fraction, and subsequent administration or nonadministration of amiodarone. VR dynamicity was compared within the cardiac death group between the 59 patients who died from ACD and the 59 who died from non-ACD. VR dynamicity was expressed as the slope of the linear regression between QTo (measured automatically) and stable RR intervals. Patients who died were found to have a significant steeper rate dependence of QTo intervals during the 3 periods than their matched survivors. In multivariate analysis, the QTo/RR nocturnal interval appeared to be the best independent predictor of cardiac death. In addition, patients who died from ACD were found to have a significant steeper rate dependence of QTo intervals during the morning period than those who died from non-ACD. In the multivariate analysis, the QTo/RR morning interval remained the best independent predictor of ACD. Thus, in the EMIAT trial, evaluation of QT dynamicity is a strong predictor of cardiac death. In addition, QT dynamicity could predict the occurrence of ACD in cases of cardiac death.

Aged↗

Picosecond dynamics of T and R forms of aspartate transcarbamylase: a neutron scattering study.

E. coli aspartate transcarbamylase (ATCase) is a 310 kDa allosteric enzyme which catalyses the first committed step in pyrimidine biosynthesis. The binding of its substrates, carbamylphosphate and aspartate, induces significant conformational changes. This enzyme shows homotropic cooperative interactions between the catalytic sites for the binding of aspartate. This property is explained by a quaternary structure transition from T state (aspartate low affinity) to R state (aspartate high affinity) accompanied by a 5% increase of radius of gyration of ATCase. The same quaternary structure change is observed upon binding of the bisubstrate analogue PALA (N-(phosphonacetyl)-L-aspartate. Owing to the large incoherent neutron scattering cross-section of the hydrogen atom and the abundance of this element in proteins, inelastic neutron scattering gives a global view of protein dynamics as sensed via the individual motions of its hydrogen atoms. We present neutron scattering results of the local dynamics (few angstroms), at short time (few tens of picoseconds), of ATCase in T and R forms. Compared to the T form, we observe an increased mobility of the protein in the R form that we associate to an increase of accessible surface area to the solvent. Beyond this specific result, this highlights the key role of the accessible surface area (ASA) in dynamic contribution to inelastic neutron data in the picosecond time scale. In particular, we want to stress out (i) that a difference at the picosecond time scale does not allow to conclude to a difference in the dynamics at a longer time scale and to address whether the T state is looser than the R state (ii) how challenging is, any comparison in terms of general dynamics (tense or relaxed) between dynamic values deduced from experimental neutron data on proteins with different sequences and therefore ASA. This caveat holds particularly when comparing dynamics of a mesophile with the corresponding extremophile.

Aspartate Carbamoyltransferase↗

Biomechanical evaluation of a dynamic pedicle screw fixation device.

BACKGROUND: Recent innovations in dynamic devices have promised a reduction in stress shielding, protection of adjacent segment degeneration, and decreased implant failure. However, there have been few studies comparing the biomechanical properties of a rigid device in comparison to a dynamic posterior fixation device. The purpose of this study was to compare the immediate stability of a new dynamic pedicle screw fixation device with an equivalent rigid device. METHODS: Six thoracolumbar cadaver spines (T10-L4) were fixed in a biomechanical testing frame. Pure moments of 10Nm were loaded in six directions: flexion, extension, right and left lateral bending, and right and left axial rotation. For each spine, four different stages were tested: intact, destabilization of the middle segment, fixation with the dynamic device, and fixation with the rigid device. Ranges of motion were measured using stereophotogrammetry. The specimens with each device were then subjected to flexion-compression loading for five cycles on a MTS 858 Universal Testing Machine. The average stiffness of the last three cycles was recorded. FINDINGS: Both dynamic and rigid devices were found to provide stability for the injured segment in flexion-extension and lateral bending. In axial rotation, the devices could restore the stability to levels similar to those in an intact spine. Results also indicated a slight increase in range of motion in flexion-extension and significant reduction in stiffness of flexion-compression with the dynamic device (P < 0.01), in comparison to the rigid device. INTERPRETATION: The dynamic device offers a system that may alter favorably the movement and load transmission of a spinal motion segment without sacrificing construct stability.

Adult↗

Effects of dynamic rotation on event-related brain potentials.

Event-related potentials were recorded during a mental rotation task. Subjects were shown pairs of letter-like shapes and were asked to make a parity judgment. The shape on the left was always in its canonical position and the shape on the right could either be in its canonical position or be a mirror image. Two variables were manipulated for the shape on the right. First, it could appear at different orientations (50 degrees , 100 degrees or 150 degrees ); second, it could be presented in a stationary position, in a dynamic congruent direction (the shape slowly rotating toward its normal upright position) or in a dynamic incongruent direction (the shape slowly rotating in the opposite direction to its normal upright position). Orientation- and direction-dependent modulations of a negative slow wave were found. For orientation, the typical amplitude effect over parietal sites was found, the amplitude becoming more negative as the rotational angle increased. For direction, the amplitude of the negative slow wave was larger for stationary and dynamic incongruent trials than for dynamic congruent trials at 100 degrees and 150 degrees . This result suggests that presentation of a stimulus in a dynamic congruent direction facilitates the mental rotation process. At 50 degrees , differences between dynamic incongruent trials and both stationary and dynamic congruent trials were found, suggesting that the incongruent movement elicits an obstructing effect over the mental rotation process. In summary, the present experiment provides new evidence in support of the idea that the amplitude modulation over the parietal cortex is a psychophysiological marker of the mental rotation process.

Adolescent↗

Spontaneous facial mimicry in response to dynamic facial expressions.

Based on previous neuroscientific evidence indicating activation of the mirror neuron system in response to dynamic facial actions, we hypothesized that facial mimicry would occur while subjects viewed dynamic facial expressions. To test this hypothesis, dynamic/static facial expressions of anger/happiness were presented using computer-morphing (Experiment 1) and videos (Experiment 2). The subjects' facial actions were unobtrusively videotaped and blindly coded using Facial Action Coding System [FACS; Ekman, P., & Friesen, W. V. (1978). Facial action coding system. Palo Alto, CA: Consulting Psychologist]. In the dynamic presentations common to both experiments, brow lowering, a prototypical action in angry expressions, occurred more frequently in response to angry expressions than to happy expressions. The pulling of lip corners, a prototypical action in happy expressions, occurred more frequently in response to happy expressions than to angry expressions in dynamic presentations. Additionally, the mean latency of these actions was less than 900 ms after the onset of dynamic changes in facial expression. Naive raters recognized the subjects' facial reactions as emotional expressions, with the valence corresponding to the dynamic facial expressions that the subjects were viewing. These results indicate that dynamic facial expressions elicit spontaneous and rapid facial mimicry, which functions both as a form of intra-individual processing and as inter-individual communication.

Adult↗

Dynamic MR imaging in Tolosa-Hunt syndrome.

OBJECTIVE: To evaluate the cavernous sinuses with dynamic magnetic resonance (MR) imaging in patients with Tolosa-Hunt syndrome (THS). METHODS: The sellar and parasellar regions of five patients with THS and 12 control subjects were examined with dynamic MR (1.5 T) imaging in the coronal plane. Dynamic images were obtained with spin-echo (SE) sequences in three patients, and with fast spin-echo (FSE) sequences in two patients and control subjects. Conventional MR images of the cranium including sellar and parasellar regions were also obtained on T1-weighted pre- and post-contrast SE, and T2-weighted FSE sequences in the coronal plane. RESULTS: MR images revealed affected cavernous sinus with bulged convex lateral wall in three patients and concave lateral wall in two patients. In all control subjects, cavernous sinuses were observed with concave lateral wall. The signal intensity on T1- and T2-weighted images and contrast enhancement on post-contrast images of the affected cavernous sinuses in patients were similar to those of the unaffected cavernous sinuses in patients and control subjects. The dynamic images in all patients disclosed small areas adjacent to the cranial nerve filling-defects within the enhanced venous spaces of the affected cavernous sinus, which showed slow and gradual enhancement from the early to the late dynamic images. No such gradually enhancing area was observed in control subjects except one. The follow-up dynamic MR images after corticosteroid therapy revealed complete resolution of the gradually enhancing areas in the previously affected cavernous sinus. CONCLUSION: Dynamic MR imaging may facilitate the diagnosis of THS.

Adult↗

Dynamic range relations for auditory primary afferents.

Dynamic range is one of four attributes typically assigned to the plot of firing rate vs. stimulus level of an auditory primary afferent. Dynamic range is generally understood to be the contiguous range of sound-pressure-level over which the neuron can indicate some small level change. Typically, however, dynamic range has been quantified as the width in decibels between the endpoints of the rate-level plot, which is not a measure of sensitivity to level change. A sensitivity measure is provided here by first deriving an equation for the intensity-difference limen (DL) in terms of attributes of the rate-level curve. The result is a generally U-shaped curve of DL vs. level. Any given criterion DL corresponds to a horizontal line cutting the DL curve at two points, with the separation in decibels between those points providing a dynamic range for that DL criterion. Plotting the dynamic ranges vs. the respective DLs yields a dynamic range curve. These were made for 62 afferents from the cat. The dynamic ranges of sloping-saturating rate-level plots do not exceed those for sigmoidal plots until the DL criterion reaches 50 dB, supporting the conclusion of Palmer and Evans [Cochlear fibre rate-intensity functions: no evidence for basilar membrane nonlinearities, Hearing Research 2 (1980) 319-326] that sloping saturation is not a reflection of cochlear nonlinearity.

Acoustic Stimulation↗

A new view on visuomotor channels: the case of the disappearing dynamics.

A considerable body of kinematic data supports the proposal that independent visuomotor channels are involved in the control of the transport and grip components of reach and grasp. These channels are seen as having separate perceptual inputs, outputs and internal processing and are thought by some to correspond to independent neuroanatomical pathways. The idea that different groups of muscles and biomechanical structures can be controlled independently is attractive, but this kinematically-inspired hypothesis fails to take into account the complexity of the dynamic relationships and their interactions within the neuromusculoskeletal system. Inertial, viscous, centrifugal, coriolis, gravitational and reflex cross couplings exist between efferent drives to muscles and resulting body movements. Rotation at even a single joint generates a complex set of dynamic reaction forces and requires coordinated activation of many muscles throughout the body to maintain posture and balance. In this theoretical paper we present a new view of independent visuomotor channels in the form of an adaptive neural controller that can compensate for the above interactions and decouple the relationships between efferent drives to muscles and resulting body movements. At the same time, the neural controller renders all the dynamics (linear and nonlinear), other than time delays, of the neuromusculoskeletal system, unobservable in the visuomotor relationships. Using the geometry of nonlinear dynamical systems we show that, providing certain constraints on the structure of time delays within the system are satisfied, there exists a neural controller that can render all the dynamics of the neuromusculoskeletal system (except for time delays) unobservable in the responses. The controller simultaneously decouples all the interactive dynamics so that each of the m independent inputs controls one and only one degree of freedom of the response. This means that each degree of freedom in a multi-joint response can be controlled by an independent component of the visual input, a behaviour that has long been observed in visual tracking experiments. The controller effectively establishes m independent visuomotor channels. However, rather than reflecting separate neuroanatomical pathways, the independent channels result from a neural controller with convergent and divergent connections to compensate for the interactive nonlinear dynamics within the neuromusculoskeletal system. This new view of visuomotor channels has implications for neural control processes involved in the acquisition and adaptability of skilled perceptual-motor behaviour in general, as well as for the design of robotic controllers.

Biomechanical Phenomena↗

Effects of dynamic compressive loading on chondrocyte biosynthesis in self-assembling peptide scaffolds.

Dynamic mechanical loading has been reported to affect chondrocyte biosynthesis in both cartilage explant and chondrocyte-seeded constructs. In this study, the effects of dynamic compression on chondrocyte-seeded peptide hydrogels were analyzed for extracellular matrix synthesis and retention over long-term culture. Initial studies were conducted with chondrocyte-seeded agarose hydrogels to explore the effects of various non-continuous loading protocols on chondrocyte biosynthesis. An optimized alternate day loading protocol was identified that increased proteoglycan (PG) synthesis over control cultures maintained in free-swelling conditions. When applied to chondrocyte-seeded peptide hydrogels, alternate day loading stimulated PG synthesis up to two-fold higher than that in free-swelling cultures. While dynamic compression also increased PG loss to the medium throughout the 39-day time course, total PG accumulation in the scaffold was significantly higher than in controls after 16 and 39 days of loading, resulting in an increase in the equilibrium and dynamic compressive stiffness of the constructs. Viable cell densities of dynamically compressed cultures differed from free-swelling controls by less than 20%, demonstrating that changes in PG synthesis were due to an increase in the average biosynthesis per viable cell. Protein synthesis was not greatly affected by loading, demonstrating that dynamic compression differentially regulated the synthesis of PGs. Taken together, these results demonstrate the potential of dynamic compression for stimulating PG synthesis and accumulation for applications to in vitro culture of tissue engineered constructs prior to implantation.

Animals↗

Dynamic shear properties of the porcine molar periodontal ligament.

The role of the periodontal ligament (PDL) is to support the tooth during function and resist external forces applied to it. The dominant vertical component of these forces is associated with shear in the PDL. Little information, however, is available on the dynamic behavior of the PDL in shear. Therefore, the present study was designed to determine the dynamic shear properties of the PDL in the porcine molar (n=10). From dissected mandibles transverse sections of the mesial root of the first molar were obtained at the apical and coronal levels and used for dynamic shear tests. Shear strain (0.5%, 1.0%, and 1.5%) was applied in superoinferior direction parallel to the root axis with a wide range of frequencies (0.01-100 Hz). The dynamic complex and storage moduli increased significantly with the loading frequency, the dynamic loss modulus showed only a small increase. The dynamic elasticity was significantly larger in the coronal region than in the apical region although the dynamic viscosity was similar in both regions. The present results suggest that non-linearities, compression/shear coupling, and intrinsic viscoelasticity affect the shear material behavior of the PDL, which might have important implications for load transmission from tooth to bone and vice versa.

Animals↗