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Mechanotransduction

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by Chowdhury TT and Agarwal S have shown that mechanical loading which represents physiological loading conditions will block the production of catabolic mediators (iNOS, COX-2, NO, PGE2) induced by inflammatory cytokines (IL-1) and restore anabolic activities. Thus an improved understanding of the interplay of biomechanics and cell signalling will help to develop therapeutic methods for blocking catabolic components of the mechanotransduction pathway. A better understanding of the optimal levels of in vivo mechanical forces are therefore necessary for maintaining the health and viability of cartilage, preventative techniques may be devised for the prevention of cartilage degradation and disease.
305:. Decreased mechanical loading over long periods, such as during extended bed-rest, causes a loss of matrix production. Static loads have been shown to be detrimental to biosynthesis while oscillatory loads at low frequencies (similar that of a normal walking gait) have been shown to be beneficial in maintaining health and increasing matrix synthesis. Due to the complexity of in-vivo loading conditions and the interplay of other mechanical and biochemical factors, the question of what an optimal loading regimen may be or whether one exists remain unanswered. 226: 97: 250: 340: 424:
act as a mechanoreceptor for the cell, transducing forces from the extracellular matrix into the cell. Each chondrocyte has one cilium and it is hypothesized to transmit mechanical signals by way of bending in response to ECM loading. Integrins have been identified on the upper shaft of the cilium,
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of the organ of Corti, causing the hair bundles that link the two to be deflected, initiating mechano-electrical transduction. When the basilar membrane is driven upward, shear between the hair cells and the tectorial membrane deflects hair bundles in the excitatory direction, toward their tall edge.
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It is important to examine the mechanotransduction pathways in chondrocytes since mechanical loading conditions which represent an excessive or injurious response upregulates synthetic activity and increases catabolic signalling cascades involving mediators such as NO and MMPs. In addition, studies
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When a deformation is imposed on a muscle, changes in cellular and molecular conformations link the mechanical forces with biochemical signals, and the close integration of mechanical signals with electrical, metabolic, and hormonal signaling may disguise the aspect of the response that is specific
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in FASEB Journal have demonstrated for the first time that primary cilia are required for chondrocyte mechanotransduction. Chondrocytes derived from IFT88 mutant mice did not express primary cilia and did not show the characteristic mechanosensitive up regulation of proteoglycan synthesis seen in
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to open and produce a transduction current that changes the membrane potential of the cell. Typically the mechanical stimulus gets filtered in the conveying medium before reaching the site of mechanotransduction. Cellular responses to mechanotransduction are variable and give rise to a variety of
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Integrin signaling is just one example of multiple pathways that are activated when cartilage is loaded. Some intracellular processes that have been observed to occur within these pathways include phosphorylation of ERK1/2, p38 MAPK, and SAPK/ERK kinase-1 (SEK-1) of the JNK pathway as well as
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is to act as a low-friction, load-bearing surface. Due to its unique location at joint surfaces, articular cartilage experiences a range of static and dynamic forces that include shear, compression and tension. These mechanical loads are absorbed by the cartilage
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Using the integrin-linked mechanotransduction pathway as an example (being one of the better studied pathways), it has been shown to mediate chondrocyte adhesion to cartilage surfaces, mediate survival signaling and regulate matrix production and degradation.
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complex (FAC). The activation of these FAC molecules in turn, triggers downstream events that up-regulate and /or down-regulate intracellular processes such as transcription factor activation and gene regulation resulting in apoptosis or differentiation.
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Although studies have shown that, like most biological tissues, cartilage is capable of mechanotransduction, the precise mechanisms by which this is done remain unknown. However, there exist a few hypotheses which begin with the identification of
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family. Chondrocytes have been shown to secrete TGF-b, and upregulate TGF-b receptors in response to mechanical stimulation; this secretion may be a mechanism for autocrine signal amplification within the tissue.
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Chondrocyte surface mechano-receptors include CD44, annexin V and integrins. Chondrocyte extracellular matrix components include collagens, proteoglycans (which consist of aggrecan and hyaluronan), fibronectin and
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processes is strongly influenced by the type of loading that cartilage experiences. High strain rates (such as which occurs during impact loading) cause tissue damage, degradation, decreased matrix production and
191:: near the base for high frequencies, near the apex for low frequencies. Hair cells in the cochlea are stimulated when the basilar membrane is driven up and down by differences in the fluid pressure between the 1416:
Loening, A; James, IE; Levenston, ME; Badger, AM; Frank, EH; Kurz, B; Nuttall, ME; Hung, HH; Blake, SM (2000). "Injurious Mechanical Compression of Bovine Articular Cartilage Induces Chondrocyte Apoptosis".
1773:"Mechanotransduction via integrins and interleukin–4 results in altered aggrecan and matrix metalloproteinase 3 gene expression in normal, but not osteoarthritic, human articular chondrocytes" 1546:
Sah, Robert L.-Y.; Kim, Young-Jo; Doong, Joe-Yuan H.; Grodzinsky, Alan J.; Plass, Anna H. K.; Sandy, John D. (1989). "Biosynthetic response of cartilage explants to dynamic compression".
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At the midpoint of an oscillation the hair bundles resume their resting position. When the basilar membrane moves downward, the hair bundles are driven in the inhibitory direction.
151:). They can also be suggested by sampling in extensive molecular dynamics trajectories and principal component analysis, or they can be directly observed using spectra measured by 107:
In this process, a mechanically gated ion channel makes it possible for sound, pressure, or movement to cause a change in the excitability of specialized sensory cells and
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Behrens, Fred; Kraft, Ellen L.; Oegema, Theodore R. (1989). "Biochemical changes in articular cartilage after joint immobilization by casting or external fixation".
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In order for mechanical signals to be sensed, there need to be mechanoreceptors on the surface of chondrocytes. Candidates for chondrocyte mechanoreceptors include
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Mouw, J. K.; Imler, S. M.; Levenston, M. E. (2006). "Ion-channel Regulation of Chondrocyte Matrix Synthesis in 3D Culture Under Static and Dynamic Compression".
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Biswas, Abhijit; Manivannan, M.; Srinivasan, Mandyam A. (2014). "Nonlinear two stage mechanotransduction model and neural response of Pacinian Corpuscle".
1295:"Mechanical Compression of Cartilage Explants Induces Multiple Time-dependent Gene Expression Patterns and Involves Intracellular Calcium and Cyclic AMP" 1855: 1689:
Kurtis, Melissa S.; Tu, Buu P.; Gaya, Omar A.; Mollenhauer, Jürgen; Knudson, Warren; Loeser, Richard F.; Knudson, Cheryl B.; Sah, Robert L. (2001).
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Torzilli, P. A.; Deng, X-H.; Ramcharan, M. (2006). "Effect of Compressive Strain on Cell Viability in Statically Loaded Articular Cartilage".
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Chondrocytes sense and convert the mechanical signals they receive into biochemical signals, which subsequently direct and mediate both
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Tavernarakis, Nektarios; Driscoll, Monica (1997). "Molecular Modeling of Mechanotransduction in the Nematode Caenorhabditis Elegans".
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Hackney, CM; Furness, DN (1995). "Mechanotransduction in vertebrate hair cells: Structure and function of the stereociliary bundle".
1336:"Shear and Compression Differentially Regulate Clusters of Functionally Related Temporal Transcription Patterns in Cartilage Tissue" 2267: 1789: 1772: 1125: 417:
changes in cAMP levels, actin re-organization and changes in the expression of genes which regulate cartilage ECM content.
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Millward-Sadler, S. J.; Salter, D. M. (2004). "Integrin-Dependent Signal Cascades in Chondrocyte Mechanotransduction".
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1. Kandel, E.R., Schwartz, J.H., Jessell, T.M., Principles of Neural Science. New York: McGraw-Hill ed, ed. 4th. 2000.
2297: 1210: 1988:"Primary cilia mediate mechanotransduction through control of ATP-induced Ca2+ signaling in compressed chondrocytes" 389: 183:
of the cochlea generates a pressure field within the cochlear fluids, imparting a pressure differential across the
453:"Vibrotactile Sensitivity Threshold: Nonlinear Stochastic Mechanotransduction Model of the Pacinian Corpuscle" 317: 148: 1143:"Stiffness gradient along the basilar membrane as a basis for spatial frequency analysis within the cochlea" 2546: 2290: 2165:"Putting ion channels to work: Mechanoelectrical transduction, adaptation, and amplification by hair cells" 200: 1375:
Kurz, Bodo; Jin, Moonsoo; Patwari, Parth; Cheng, Debbie M.; Lark, Michael W.; Grodzinsky, Alan J. (2001).
2272: 541:"Mechanical Strain Induces pp60src Activation and Translocation to Cytoskeleton in Fetal Rat Lung Cells" 155:
spectroscopy. Current findings indicate that the mechanotransduction channel in hair cells is a complex
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Biomedical Science and Engineering Center Conference (BSEC), 2014 Annual Oak Ridge National Laboratory
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Gillespie, Peter G.; Walker, Richard G. (2001). "Molecular basis of mechanosensory transduction".
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Howard, J; Roberts, W M; Hudspeth, A J (1988). "Mechanoelectrical Transduction by Hair Cells".
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Integrin receptors have an extracellular domain that binds to the ECM proteins (collagen,
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Single-molecule biomechanics studies of proteins and DNA, and mechanochemical coupling in
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Millward–Sadler, S. J.; Wright, M. O.; Davies, L. W.; Nuki, G.; Salter, D. M. (2000).
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have demonstrated the critical importance of molecular mechanics as a new frontier in
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acting as anchors to the collagen matrix around it. Recent studies published by Wann
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Fanning, P. J.; Emkey, G; Smith, RJ; Grodzinsky, AJ; Szasz, N; Trippel, SB (2003).
1841: 1821: 1784: 1743: 1702: 1653: 1645: 1618: 1598: 1575: 1555: 1512: 1469: 1426: 1388: 1347: 1306: 1265: 1257: 1216: 1198: 1187:"Hair cell transduction, tuning and synaptic transmission in the mammalian cochlea" 1146: 1086: 1076: 1027: 1019: 970: 960: 911: 903: 848: 809: 786: 766: 723: 688: 653: 626: 593: 580:
Bidhendi, Amir J; Altartouri, Bara; Gosselin, Frédérick P.; Geitmann, Anja (2019).
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Mammano, F.; Nobili, R (1993). "Biophysics of the cochlea: Linear approximation".
1947:"Localization of Extracellular Matrix Receptors on the Chondrocyte Primary Cilium" 727: 2503: 1115: 813: 598: 192: 136: 112: 58: 2131:
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Fitzgerald, J. B.; Jin, M; Dean, D; Wood, DJ; Zheng, MH; Grodzinsky, AJ (2004).
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Proceedings of the National Academy of Sciences of the United States of America
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Proceedings of the National Academy of Sciences of the United States of America
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and life sciences. Protein domains, connected by intrinsically disordered
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Conversion of mechanical stimulus of a cell into electrochemical activity
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In addition to binding to ECM ligands, integrins are also receptive to
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10.1002/1529-0131(200009)43:9<2091::AID-ANR21>3.0.CO;2-C
187:. A sinusoidal pressure wave results in localized vibrations of the 2433: 836: 452: 409: 381: 333: 293: 258: 249: 172: 1910:
Urban, J. P. G. (1994). "The Chondrocyte: A Cell Under Pressure".
837:"Multiscale Layered Biomechanical Model of the Pacinian Corpuscle" 579: 253:
Cartilage experience tension, compression and shear forces in vivo
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Air pressure changes in the ear canal cause the vibrations of the
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changes and sensations. Broader issues involved include molecular
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Pulai, Judit I.; Del Carlo, Marcello; Loeser, Richard F. (2002).
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Bu Z, Biehl R, Monkenbusch M, Richter D, Callaway DJ (Dec 2005).
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converting mechanical signals into electrical or chemical signals
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Biswas, Abhijit; Manivannan, M.; Srinivasan, Mandyam A. (2015).
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Biswas, Abhijit; Manivannan, M.; Srinivasan, Mandyam A. (2015).
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receptors (of which there exist several types on chondrocytes).
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More recent studies have hypothesized that chondrocyte primary
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Hudspeth, A. J.; Choe, Y.; Mehta, A. D.; Martin, P. (2000).
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Qin, Y.; Qin, Y; Liu, J; Tanswell, AK; Post, M (1996).
497: 89:. The basic mechanism of mechanotransduction involves 1113: 1811: 1545: 1502: 1292: 681:
Annual Review of Biophysics and Biophysical Chemistry
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Katsumi, A.; Orr, AW; Tzima, E; Schwartz, MA (2003).
392:) are recruited to this cluster, which is called the 175:. At the end of the ossicular chain, movement of the 2216:
Hudspeth, A. J. (1989). "How the ear's works work".
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Cambridge University Press. 742: 707: 672: 637: 614: 573: 532: 491: 444: 318:stretch-activated ion channels 115:causes mechanically sensitive 1: 1708:10.1016/S0736-0266(01)00051-1 1394:10.1016/S0736-0266(01)00033-X 802:Journal of Applied Physiology 728:10.1152/ajpcell.1995.268.1.C1 437: 149:Database of Molecular Motions 2273:Cellular+Mechanotransduction 2041:Khan, K M; Scott, A (2009). 1141:Ehret, Günter (2009-05-08). 841:IEEE Transactions on Haptics 814:10.1152/jappl.1986.60.4.1107 599:10.1016/j.celrep.2019.07.006 457:IEEE Transactions on Haptics 220: 7: 646:Annual Review of Physiology 625:. USA: IEEE. pp. 1–4. 139:domains, induce long-range 10: 2706: 1777:Arthritis & Rheumatism 1736:Arthritis & Rheumatism 229:Mechanically gated channel 217:to the mechanical forces. 18: 2604: 2574: 2537: 2512: 2489: 2480: 2452: 2406: 2399: 2324: 1603:10.1007/s10237-006-0034-1 1517:10.1007/s10237-006-0030-5 1024:10.1016/j.bpj.2010.09.058 908:10.1016/j.bpj.2009.03.051 631:10.1109/BSEC.2014.6867740 2632:Neuroendocrine signaling 2277:Medical Subject Headings 2190:10.1073/pnas.97.22.11765 2059:10.1136/bjsm.2008.054239 1191:Comprehensive Physiology 853:10.1109/TOH.2014.2369416 469:10.1109/TOH.2014.2369422 1964:10.1369/jhc.5A6866.2006 1828:(inactive 2024-03-26). 1250:Frontiers in Bioscience 1244:Burkholder, TJ (2007). 1117:Molecular Communication 1082:10.1073/pnas.0503388102 966:10.1073/pnas.1207892109 558:10.1074/jbc.271.12.7066 277:, protease inhibitors, 145:protein domain dynamics 111:. The stimulation of a 61:activity. This form of 2494:cAMP-dependent pathway 1888:10.1074/jbc.M305107200 1560:10.1002/jor.1100070502 1474:10.1002/jor.1100070305 1431:10.1006/abbi.2000.1988 1353:10.1074/jbc.M510858200 1312:10.1074/jbc.M400437200 1166:Cite journal requires 517:10.1074/jbc.R300038200 345: 261:(matrix building) and 254: 230: 162: 104: 2620:Synaptic transmission 2616:Neurocrine signaling 2539:Transcription factors 2386:PI3K/AKT/mTOR pathway 1650:10.1007/s000180050069 374:focal adhesion kinase 342: 279:transcription factors 252: 228: 179:footplate within the 99: 2005:10.1096/fj.11-193649 1203:10.1002/cphy.c160049 240:extracellular matrix 63:sensory transduction 45:) is any of various 2649:Mechanotransduction 2637:Exocrine signalling 2444:Signaling molecules 2318:Signal transduction 2230:1989Natur.341..397H 2181:2000PNAS...9711765H 2145:1952ASAJ...24...72V 2098:1993ASAJ...93.3320M 1151:10.18725/OPARU-1218 1073:2005PNAS..10217646B 1067:(49): 17646–17651. 1016:2010BpJ....99.3473F 1004:Biophysical Journal 957:2012PNAS..10912509B 900:2009BpJ....96.4993P 888:Biophysical Journal 763:2001Natur.413..194G 246:(cartilage cells). 102:biological machines 55:mechanical stimulus 31:mechanotransduction 2659:Ion channel gating 2391:Integrin receptors 2325:Signaling pathways 346: 255: 231: 201:tectorial membrane 157:biological machine 105: 2672: 2671: 2654:Phototransduction 2612:Intracrine action 2570: 2569: 2533: 2532: 2421:Neurotransmitters 2224:(6241): 397–404. 2153:10.1121/1.1906851 1749:10.1002/art.10334 1346:(34): 24095–103. 1127:978-1-107-02308-6 1010:(10): 3473–3482. 894:(12): 4993–5002. 757:(6852): 194–202. 722:(1 Pt 1): C1–13. 169:tympanic membrane 153:neutron spin echo 2697: 2625:Chemical synapse 2525:Scaffold protein 2487: 2486: 2482:Second messenger 2408:Receptor ligands 2404: 2403: 2307: 2300: 2293: 2284: 2283: 2257: 2238:10.1038/341397a0 2212: 2202: 2192: 2175:(22): 11765–72. 2156: 2127: 2117: 2106:10.1121/1.405716 2080: 2070: 2028: 2027: 2017: 2007: 1983: 1977: 1976: 1966: 1942: 1936: 1935: 1907: 1901: 1900: 1890: 1866: 1860: 1859: 1853: 1845: 1809: 1803: 1802: 1792: 1783:(9): 2091–2099. 1768: 1762: 1761: 1751: 1727: 1721: 1720: 1710: 1686: 1680: 1679: 1661: 1629: 1623: 1622: 1586: 1580: 1579: 1543: 1537: 1536: 1500: 1494: 1493: 1457: 1451: 1450: 1413: 1407: 1406: 1396: 1372: 1366: 1365: 1355: 1331: 1325: 1324: 1314: 1305:(19): 19502–11. 1290: 1284: 1283: 1273: 1241: 1235: 1234: 1224: 1197:(4): 1197–1227. 1182: 1176: 1175: 1169: 1164: 1162: 1154: 1138: 1132: 1131: 1111: 1105: 1104: 1094: 1084: 1052: 1046: 1045: 1035: 995: 989: 988: 978: 968: 951:(31): 12509–14. 936: 930: 929: 919: 879: 873: 872: 832: 826: 825: 797: 791: 790: 771:10.1038/35093011 746: 740: 739: 711: 705: 704: 676: 670: 669: 641: 635: 634: 618: 612: 611: 601: 592:(5): 1237–1250. 577: 571: 570: 560: 536: 530: 529: 519: 495: 489: 488: 448: 430:wild type cells 378:adapter proteins 311:mechanoreceptors 267:type II collagen 205:reticular lamina 185:basilar membrane 129:molecular motors 27:cellular biology 21:Mechanosensation 2705: 2704: 2700: 2699: 2698: 2696: 2695: 2694: 2675: 2674: 2673: 2668: 2600: 2566: 2529: 2508: 2504:Lipid signaling 2476: 2448: 2395: 2320: 2311: 2264: 2037: 2035:Further reading 2032: 2031: 1984: 1980: 1943: 1939: 1918:(10): 901–908. 1908: 1904: 1881:(51): 50940–8. 1867: 1863: 1847: 1846: 1810: 1806: 1769: 1765: 1728: 1724: 1687: 1683: 1630: 1626: 1587: 1583: 1544: 1540: 1511:(2–3): 123–32. 1501: 1497: 1458: 1454: 1414: 1410: 1373: 1369: 1332: 1328: 1291: 1287: 1242: 1238: 1213: 1183: 1179: 1167: 1165: 1156: 1155: 1139: 1135: 1128: 1112: 1108: 1053: 1049: 996: 992: 937: 933: 880: 876: 833: 829: 798: 794: 747: 743: 712: 708: 677: 673: 642: 638: 619: 615: 578: 574: 551:(12): 7066–71. 537: 533: 510:(13): 12001–4. 496: 492: 449: 445: 440: 388:, aka Tal, and 223: 214: 212:Skeletal muscle 193:scala vestibuli 171:and middle ear 165: 137:flexible linker 113:mechanoreceptor 109:sensory neurons 59:electrochemical 23: 17: 12: 11: 5: 2703: 2693: 2692: 2690:Cell signaling 2687: 2670: 2669: 2667: 2666: 2661: 2656: 2651: 2646: 2645: 2644: 2634: 2629: 2628: 2627: 2622: 2614: 2608: 2606: 2605:Other concepts 2602: 2601: 2599: 2598: 2593: 2584: 2578: 2576: 2572: 2571: 2568: 2567: 2565: 2564: 2559: 2554: 2549: 2543: 2541: 2535: 2534: 2531: 2530: 2528: 2527: 2522: 2516: 2514: 2510: 2509: 2507: 2506: 2501: 2496: 2490: 2484: 2478: 2477: 2475: 2474: 2469: 2464: 2458: 2456: 2450: 2449: 2447: 2446: 2441: 2439:Growth factors 2436: 2431: 2418: 2412: 2410: 2401: 2397: 2396: 2394: 2393: 2388: 2383: 2378: 2373: 2368: 2363: 2358: 2357: 2356: 2351: 2341: 2336: 2334: 2328: 2326: 2322: 2321: 2314:Cell signaling 2310: 2309: 2302: 2295: 2287: 2281: 2280: 2270: 2263: 2262:External links 2260: 2259: 2258: 2213: 2160: 2157: 2128: 2092:(6): 3320–32. 2081: 2036: 2033: 2030: 2029: 1998:(4): 1663–71. 1978: 1957:(9): 1005–14. 1937: 1902: 1861: 1804: 1763: 1742:(6): 1528–35. 1722: 1701:(6): 1122–30. 1681: 1624: 1597:(1–2): 33–41. 1581: 1538: 1495: 1452: 1408: 1367: 1326: 1285: 1236: 1211: 1177: 1168:|journal= 1133: 1126: 1106: 1047: 990: 931: 874: 827: 808:(4): 1107–15. 792: 741: 706: 671: 636: 613: 572: 531: 490: 463:(1): 102–113. 442: 441: 439: 436: 394:focal adhesion 287:growth factors 222: 219: 213: 210: 189:organ of Corti 164: 161: 133:bioengineering 75:proprioception 15: 9: 6: 4: 3: 2: 2702: 2691: 2688: 2686: 2683: 2682: 2680: 2665: 2662: 2660: 2657: 2655: 2652: 2650: 2647: 2643: 2640: 2639: 2638: 2635: 2633: 2630: 2626: 2623: 2621: 2618: 2617: 2615: 2613: 2610: 2609: 2607: 2603: 2597: 2594: 2592: 2588: 2585: 2583: 2580: 2579: 2577: 2573: 2563: 2560: 2558: 2555: 2553: 2550: 2548: 2545: 2544: 2542: 2540: 2536: 2526: 2523: 2521: 2518: 2517: 2515: 2511: 2505: 2502: 2500: 2497: 2495: 2492: 2491: 2488: 2485: 2483: 2479: 2473: 2470: 2468: 2467:Intracellular 2465: 2463: 2460: 2459: 2457: 2455: 2451: 2445: 2442: 2440: 2437: 2435: 2432: 2430: 2429:Neurohormones 2426: 2425:Neuropeptides 2422: 2419: 2417: 2414: 2413: 2411: 2409: 2405: 2402: 2398: 2392: 2389: 2387: 2384: 2382: 2379: 2377: 2376:Fas apoptosis 2374: 2372: 2369: 2367: 2364: 2362: 2359: 2355: 2352: 2350: 2347: 2346: 2345: 2342: 2340: 2337: 2335: 2333: 2330: 2329: 2327: 2323: 2319: 2315: 2308: 2303: 2301: 2296: 2294: 2289: 2288: 2285: 2278: 2274: 2271: 2269: 2266: 2265: 2255: 2251: 2247: 2243: 2239: 2235: 2231: 2227: 2223: 2219: 2214: 2210: 2206: 2201: 2196: 2191: 2186: 2182: 2178: 2174: 2170: 2166: 2161: 2158: 2154: 2150: 2146: 2142: 2138: 2134: 2129: 2125: 2121: 2116: 2115:11577/2467023 2111: 2107: 2103: 2099: 2095: 2091: 2087: 2082: 2078: 2074: 2069: 2064: 2060: 2056: 2053:(4): 247–52. 2052: 2048: 2044: 2039: 2038: 2025: 2021: 2016: 2011: 2006: 2001: 1997: 1993: 1989: 1982: 1974: 1970: 1965: 1960: 1956: 1952: 1948: 1941: 1933: 1929: 1925: 1921: 1917: 1913: 1906: 1898: 1894: 1889: 1884: 1880: 1876: 1872: 1865: 1857: 1851: 1843: 1839: 1835: 1831: 1827: 1823: 1820:(3): 435–46. 1819: 1815: 1808: 1800: 1796: 1791: 1786: 1782: 1778: 1774: 1767: 1759: 1755: 1750: 1745: 1741: 1737: 1733: 1726: 1718: 1714: 1709: 1704: 1700: 1696: 1692: 1685: 1677: 1673: 1669: 1665: 1660: 1655: 1651: 1647: 1644:(6): 539–45. 1643: 1639: 1635: 1628: 1620: 1616: 1612: 1608: 1604: 1600: 1596: 1592: 1585: 1577: 1573: 1569: 1565: 1561: 1557: 1554:(5): 619–36. 1553: 1549: 1542: 1534: 1530: 1526: 1522: 1518: 1514: 1510: 1506: 1499: 1491: 1487: 1483: 1479: 1475: 1471: 1468:(3): 335–43. 1467: 1463: 1456: 1448: 1444: 1440: 1436: 1432: 1428: 1425:(2): 205–12. 1424: 1420: 1412: 1404: 1400: 1395: 1390: 1387:(6): 1140–6. 1386: 1382: 1378: 1371: 1363: 1359: 1354: 1349: 1345: 1341: 1337: 1330: 1322: 1318: 1313: 1308: 1304: 1300: 1296: 1289: 1281: 1277: 1272: 1267: 1263: 1259: 1255: 1251: 1247: 1240: 1232: 1228: 1223: 1218: 1214: 1212:9780470650714 1208: 1204: 1200: 1196: 1192: 1188: 1181: 1173: 1160: 1152: 1148: 1144: 1137: 1129: 1123: 1119: 1118: 1110: 1102: 1098: 1093: 1088: 1083: 1078: 1074: 1070: 1066: 1062: 1058: 1051: 1043: 1039: 1034: 1029: 1025: 1021: 1017: 1013: 1009: 1005: 1001: 994: 986: 982: 977: 972: 967: 962: 958: 954: 950: 946: 942: 935: 927: 923: 918: 913: 909: 905: 901: 897: 893: 889: 885: 878: 870: 866: 862: 858: 854: 850: 846: 842: 838: 831: 823: 819: 815: 811: 807: 803: 796: 788: 784: 780: 776: 772: 768: 764: 760: 756: 752: 745: 737: 733: 729: 725: 721: 717: 710: 702: 698: 694: 690: 686: 682: 675: 667: 663: 659: 655: 651: 647: 640: 632: 628: 624: 617: 609: 605: 600: 595: 591: 587: 583: 576: 568: 564: 559: 554: 550: 546: 542: 535: 527: 523: 518: 513: 509: 505: 501: 494: 486: 482: 478: 474: 470: 466: 462: 458: 454: 447: 443: 435: 431: 428: 423: 418: 414: 411: 407: 403: 398: 395: 391: 387: 383: 379: 375: 371: 367: 363: 359: 355: 350: 341: 337: 335: 331: 327: 323: 319: 314: 312: 306: 304: 299: 295: 290: 288: 284: 280: 276: 272: 271:proteoglycans 268: 264: 260: 251: 247: 245: 241: 236: 227: 218: 209: 206: 202: 198: 197:scala tympani 194: 190: 186: 182: 178: 174: 170: 160: 158: 154: 150: 146: 142: 138: 134: 130: 125: 123: 118: 114: 110: 103: 98: 94: 92: 88: 84: 80: 76: 72: 71:physiological 68: 64: 60: 56: 52: 48: 44: 43: 38: 37: 32: 28: 22: 2664:Gap junction 2648: 2499:Ca signaling 2462:Cell surface 2221: 2217: 2172: 2168: 2139:(1): 72–76. 2136: 2132: 2089: 2085: 2050: 2046: 1995: 1991: 1981: 1954: 1950: 1940: 1915: 1912:Rheumatology 1911: 1905: 1878: 1874: 1864: 1850:cite journal 1817: 1813: 1807: 1780: 1776: 1766: 1739: 1735: 1725: 1698: 1694: 1684: 1641: 1637: 1627: 1594: 1590: 1584: 1551: 1547: 1541: 1508: 1504: 1498: 1465: 1461: 1455: 1422: 1418: 1411: 1384: 1380: 1370: 1343: 1339: 1329: 1302: 1298: 1288: 1262:10.2741/2057 1253: 1249: 1239: 1194: 1190: 1180: 1159:cite journal 1136: 1116: 1109: 1064: 1060: 1050: 1007: 1003: 993: 948: 944: 934: 891: 887: 877: 847:(1): 31–42. 844: 840: 830: 805: 801: 795: 754: 750: 744: 719: 715: 709: 684: 680: 674: 649: 645: 639: 622: 616: 589: 586:Cell Reports 585: 575: 548: 544: 534: 507: 503: 493: 460: 456: 446: 432: 426: 419: 415: 399: 351: 347: 315: 307: 291: 256: 244:chondrocytes 232: 215: 166: 126: 122:biomechanics 117:ion channels 106: 42:transduction 40: 34: 30: 24: 2575:By distance 2513:Assistants: 2472:Co-receptor 384:, aka Pax, 376:, FAK) and 366:osteopontin 362:vitronectin 354:fibronectin 320:(SAC), the 181:oval window 2685:Biophysics 2679:Categories 2642:Pheromones 2582:Juxtacrine 1256:: 174–91. 687:: 99–124. 652:: 659–89. 438:References 322:hyaluronan 47:mechanisms 19:See also: 2596:Endocrine 2591:Paracrine 2587:Autocrine 2454:Receptors 2434:Cytokines 406:paracrine 402:autocrine 330:annexin V 324:receptor 303:apoptosis 298:catabolic 283:cytokines 275:proteases 263:catabolic 235:cartilage 221:Cartilage 141:allostery 49:by which 2416:Hormones 2366:JAK-STAT 2354:MAPK/ERK 2349:TGF beta 2254:33117543 2209:11050207 2077:19244270 2024:22223751 1973:16651393 1897:12952976 1834:15095818 1799:11014361 1758:12115183 1717:11781014 1676:21313045 1659:11147192 1611:16767453 1533:39216430 1525:16506016 1490:34651862 1447:21964244 1439:11032407 1403:11781016 1362:16782710 1321:14960571 1280:17127292 1231:28915323 1101:16306270 1042:21081097 985:22802671 926:19527659 869:24658742 861:25398182 779:11557988 608:31365867 526:14960578 485:15326972 477:25398183 410:TGF-beta 382:paxillin 334:integrin 294:anabolic 259:anabolic 203:and the 173:ossicles 53:convert 2547:General 2371:Akt/PKB 2246:2677742 2226:Bibcode 2177:Bibcode 2141:Bibcode 2124:8326060 2094:Bibcode 2068:2662433 2015:3316893 1992:FASEB J 1932:7921748 1842:1717838 1668:9230933 1619:7270995 1576:1933220 1568:2760736 1482:2703926 1271:2043154 1222:5658794 1092:1345721 1069:Bibcode 1033:2980739 1012:Bibcode 976:3411975 953:Bibcode 917:2712024 896:Bibcode 822:2422151 787:4388399 759:Bibcode 736:7840137 701:3293600 666:9074782 567:8636139 380:(e.g., 372:(e.g., 370:kinases 358:laminin 87:hearing 83:balance 36:mechano 2400:Agents 2279:(MeSH) 2252:  2244:  2218:Nature 2207:  2197:  2122:  2075:  2065:  2022:  2012:  1971:  1930:  1895:  1840:  1832:  1797:  1756:  1715:  1674:  1666:  1656:  1617:  1609:  1574:  1566:  1531:  1523:  1488:  1480:  1445:  1437:  1401:  1360:  1319:  1278:  1268:  1229:  1219:  1209:  1124:  1099:  1089:  1040:  1030:  983:  973:  924:  914:  867:  859:  820:  785:  777:  751:Nature 734:  699:  664:  606:  565:  524:  483:  475:  427:et al. 422:cilium 177:stapes 85:, and 67:senses 2562:TFIIH 2557:TFIID 2381:Hippo 2361:Notch 2250:S2CID 2200:34347 1838:S2CID 1672:S2CID 1615:S2CID 1572:S2CID 1529:S2CID 1486:S2CID 1443:S2CID 865:S2CID 783:S2CID 481:S2CID 386:talin 344:COMP. 100:Some 79:touch 57:into 51:cells 2332:GPCR 2242:PMID 2205:PMID 2120:PMID 2073:PMID 2020:PMID 1969:PMID 1928:PMID 1893:PMID 1856:link 1830:PMID 1795:PMID 1754:PMID 1713:PMID 1664:PMID 1607:PMID 1564:PMID 1521:PMID 1478:PMID 1435:PMID 1399:PMID 1358:PMID 1317:PMID 1276:PMID 1227:PMID 1207:ISBN 1172:help 1122:ISBN 1097:PMID 1038:PMID 981:PMID 922:PMID 857:PMID 818:PMID 775:PMID 732:PMID 697:PMID 662:PMID 604:PMID 563:PMID 522:PMID 473:PMID 404:and 364:and 326:CD44 296:and 285:and 269:and 195:and 143:via 69:and 2344:RTK 2339:Wnt 2234:doi 2222:341 2195:PMC 2185:doi 2149:doi 2110:hdl 2102:doi 2063:PMC 2055:doi 2010:PMC 2000:doi 1959:doi 1920:doi 1883:doi 1879:278 1822:doi 1785:doi 1744:doi 1703:doi 1654:PMC 1646:doi 1599:doi 1556:doi 1513:doi 1470:doi 1427:doi 1423:381 1389:doi 1348:doi 1344:281 1307:doi 1303:279 1266:PMC 1258:doi 1217:PMC 1199:doi 1147:doi 1087:PMC 1077:doi 1065:102 1028:PMC 1020:doi 971:PMC 961:doi 949:109 912:PMC 904:doi 849:doi 810:doi 767:doi 755:413 724:doi 720:268 689:doi 654:doi 627:doi 594:doi 553:doi 549:271 512:doi 508:279 465:doi 390:Shc 273:), 163:Ear 25:In 2681:: 2589:/ 2316:/ 2248:. 2240:. 2232:. 2220:. 2203:. 2193:. 2183:. 2173:97 2171:. 2167:. 2147:. 2137:24 2135:. 2118:. 2108:. 2100:. 2090:93 2088:. 2071:. 2061:. 2051:43 2049:. 2045:. 2018:. 2008:. 1996:26 1994:. 1990:. 1967:. 1955:54 1953:. 1949:. 1926:. 1916:33 1914:. 1891:. 1877:. 1873:. 1852:}} 1848:{{ 1836:. 1818:32 1816:. 1793:. 1781:43 1779:. 1775:. 1752:. 1740:46 1738:. 1734:. 1711:. 1699:19 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Index

Mechanosensation
cellular biology
mechano
transduction
mechanisms
cells
mechanical stimulus
electrochemical
sensory transduction
senses
physiological
proprioception
touch
balance
hearing
converting mechanical signals into electrical or chemical signals

biological machines
sensory neurons
mechanoreceptor
ion channels
biomechanics
molecular motors
bioengineering
flexible linker
allostery
protein domain dynamics
Database of Molecular Motions
neutron spin echo
biological machine

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