干细胞的生物力学研究

吕东媛; 周吕文; 龙勉

doi:10.6052/1000-0992-16-041

干细胞的生物力学研究

doi: 10.6052/1000-0992-16-041

中国科学院力学研究所微重力重点实验室/工程化构建与力学生物学北京市重点实验室, 北京100190

详细信息

作者简介:
龙勉, 工学博士, 中科院力学所研究员, 博士生导师. 1995 年晋升为教授,2000 年入选中科院"百人计划", 2002 年获国家杰出青年基金, 2004 年入选"新世纪百千万人才工程国家级人选". 先后任世界生物力学理事会(WCB) 执委、国际生物流变学学会(ISB) 理事、世界华人生物医学工程协会(WACBE) 理事、开云棋牌官方副理事长等, 担任C ell. Mol. Bioengi.,Biophys. Rep. 副主编及相关杂志编委. 研究兴趣为蛋白质间相互作用与组装动力学、细胞-亚细胞力学-生物学耦合、空间细胞-分子生物力学工程. 在Nature, Biomaterials, J Immunol. 等杂志发表学术论文124篇. 获教育部首届青年教师奖、开云棋牌官方青年科技奖等. mlong@imech.ac.cn

中图分类号: R 318.01
计量
- 文章访问数: 2651
- HTML全文浏览量: 512
- PDF下载量: 1929
- 被引次数: 0
出版历程
- 收稿日期: 2016-11-15
- 网络出版日期: 2017-01-24
- 刊出日期: 2017-02-24

Biomechanics of stem cells

Key Laboratory of Microgravity (National Microgravity Laboratory) and Beijing Key Laboratory of Engineered Construction and Mechanobiology, Institute of Mechanics, Chinese Academy of Sciences, Beijing 100190, China

摘要

摘要: 干细胞生物力学作为生物力学的重要分支和前沿学科, 近年来在力学-生物学、力学-化学耦合等方面取得了重大进展, 已成为生物力学乃至生物医学工程最活跃的领域之一, 并对发育生物学、干细胞生物学、组织修复、再生医学等相关领域产生重要影响. 干细胞具有独特的力学性质, 可感知、传递、转导和响应生理力学微环境的改变, 从而调控干细胞的生长、分化等功能, 体现出典型的力学-生物学耦合特征. 本文将对干细胞的力学性质与细胞力学模型、在体力学环境对干细胞生长和分化的影响、干细胞对外界力学刺激的响应等方面加以综述.
- 干细胞 /
- 生物力学 /
- 力学-生物学耦合 /
- 力学微环境 /
- 信号传递与转导
Abstract: As an important branch of biomechanics, stem cell biomechanics has made great progress in mechano-biological, mechano-chemical coupling, and other related felds in recent years. It has become one of the most active felds in biomechanics and biomedical engineering, and made signifcant impacts on the areas of stem cell biology, tissue repair, regenerative medicine, and so on. Stem cells have unique mechanical properties, which can sense, transmit, transduce and respond to changes in physiological and mechanical microen-vironment, thus regulating the function of stem cell growth and differentiation, showing a typical mechano-biological coupling characteristics. This paper reviews the mechanical properties of stem cells and cell mechanics models, the effects of in vivo mechanical mi-croenvironment on stem cell growth and differentiation, and the responses of stem cells to external mechanical stimuli.
- stem cell /
- biomechanics /
- mechano-biological coupling /
- mechanical microenvi-ronment /
- signal transmission and transduction

HTML全文

图 1 典型的4种连续介质细胞模型.(a)线弹性固体模型(b)黏弹性固体模型(c)牛顿液滴模型,(d)复合液滴模型

下载: 全尺寸图片幻灯片

图 2 典型的3种细胞结构模型.(a)开孔泡沫模型,(b)索网模型(c)六杆张力整合模型

下载: 全尺寸图片幻灯片

图 3 组织微观力学影响干细胞分化并与细胞核内富含的骨架蛋白lamins相关( Swift et al. 2013)

下载: 全尺寸图片幻灯片

图 4 线粒体动力学通过调控核转录程序影响干细胞命运( Khacho et al. 2016)

下载: 全尺寸图片幻灯片

图 5 组织形态建成中的力学因素( Patwari & Lee 2008)

下载: 全尺寸图片幻灯片

图 6 干细胞"壁龛"中的力学作用模式( Conway & Schaffer 2012)

下载: 全尺寸图片幻灯片

图 7 硬度范围与相应的组织类型( Buxboim et al. 2010)

下载: 全尺寸图片幻灯片

图 8 基底硬度耦合拓扑结构对MSC的影响.(a)MSC在不同基底上的形态、细胞骨架及黏附斑蛋白的表达,(b)结构模式示意图

下载: 全尺寸图片幻灯片

图 9 不同基底硬度耦合拓扑结构基底对胚胎干细胞干性的影响(Lü et al. 2014)

下载: 全尺寸图片幻灯片

图 10 细胞力学信号转导的媒介( Ingber 2006)

下载: 全尺寸图片幻灯片

图 11 黏着斑复合物结构示意图( Mitra et al. 2005)

下载: 全尺寸图片幻灯片

图 12 力学组学概念示意图

下载: 全尺寸图片幻灯片

map

参考文献(232)

[1]	龙勉. 2014. 如何在地球表面模拟空间微重力环境或效应: 从空间细胞生长对微重力响应谈起. 科学通报, 59: 2004-2015 (Long M. 2014. How to stimulate a space microgravity environment or effect on Earth: From the viewpoint of responses of space cell growth to microgravity. Chinese Science Bulletin, 59: 2004-2015) . http://www.cnki.com.cn/Article/CJFDTOTAL-KXTB201420013.htm
[2]	Agresar G, Linderman J J, Tryggvason G, Powell K G. 1998. An adaptive, cartesian, front-tracking method for the motion, deformation and adhesion of circulating cells. Journal of Computational Physics, 143: 346-380. doi: 10.1006/jcph.1998.5967
[3]	Aikawa R, Nagai T, Kudoh S, Zou Y, Tanaka M, Tamura M, Akazawa H, Takano H, Nagai R, Komuro I. 2002. Integrins play a critical role in mechanical stress-induced p38 MAPK activation. Hypertension, 39: 233-238. doi: 10.1161/hy0202.102699
[4]	Alcaraz J, Buscemi L, Grabulosa M, Trepat X, Fabry B, Farré R, Navajas D. 2003. Microrheology of human lung epithelial cells measured by atomic force microscopy. Biophysical Journal, 84: 2071-2079. doi: 10.1016/S0006-3495(03)75014-0
[5]	Altman A M, Gupta V, Rios C N, Alt E U, Mathur A B. 2010. Adhesion, migration and mechanics of human adipose-tissue-derived stem cells on silk fibroin-chitosan matrix. Acta Biomaterialia, 6: 1388-1397. doi: 10.1016/j.actbio.2009.10.034
[6]	Amy C R, Jan L, Harald H, Aebi U. 2008. Towards an integrated understanding of the structure and mechanics of the cell nucleus. Bioessays, 30: 226-236. doi: 10.1002/(ISSN)1521-1878
[7]	Anneren C, Cowan C A, Melton D A. 2004. The Src family of tyrosine kinases is important for embryonic stem cell self-renewal. The Journal of Biological Chemistry, 279: 31590-31598. doi: 10.1074/jbc.M403547200
[8]	Arnsdorf E J, Tummala P, Kwon R Y, Jacobs C R. 2009. Mechanically induced osteogenic differentiation--the role of RhoA, ROCKII and cytoskeletal dynamics. Journal of Cell Science, 122: 546-553. doi: 10.1242/jcs.036293
[9]	Bachrach N M, Valhmu W B, Stazzone E, Ratcliffe A, Lai W M, Mow V C. 1995. Changes in proteoglycan synthesis of chondrocytes in articular cartilage are associated with the time-dependent changes in their mechanical environment. Journal of Biomechanics, 28: 1561-1569. doi: 10.1016/0021-9290(95)00103-4
[10]	Balaban N Q, Schwarz U S, Riveline D, Goichberg P, Tzur G, Sabanay I, Mahalu D, Safran S, Bershadsky A, Addadi L and Geiger B. 2001. Force and focal adhesion assembly: a close relationship studied using elastic micropatterned substrates. Nature Cell Biology, 3: 466-472. doi: 10.1038/35074532
[11]	Bandyopadhyay S K, Dutta A. 2005. Mitochondrial hepatopathies. The Journal of the Association of Physicians of India, 53: 973-978.
[12]	Bao G, Suresh S. Cell and molecular mechanics of biological materials. 2003. Nature materials, 2: 715-725.
[13]	Betschinger J, Knoblich J A. 2004. Dare to be different: asymmetric cell division in Drosophila, C. elegans and vertebrates. Current Biology, 14: R674-R685.
[14]	Bhat M B, Zhao J, Zang W, Balke W, Takeshima H, Wier W G, Ma J. 1997. Caffeine-induced release of intracellular Ca ²⁺ from chinese hamster ovary cells expressing skeletal muscle ryanodine receptor effects on full-length and carboxyl-terminal portion of Ca 1 release channels. The Journal of General Physiology, 110: 749-762. doi: 10.1085/jgp.110.6.749
[15]	Blaber E A, Dvorochkin N, Torres M L, Yousuf R, Burns B P, Globus R K, Almeida E A. 2014. Mechanical unloading of bone in microgravity reduces mesenchymal and hematopoietic stem cell-mediated tissue regeneration. Stem Cell Research, 13: 181-201. doi: 10.1016/j.scr.2014.05.005
[16]	Brafman D A, Phung C, Kumar N, Willert K. 2013. Regulation of endodermal differentiation of human embryonic stem cells through integrin-ECM interactions. Cell Death and Differentiation, 20: 369-381. doi: 10.1038/cdd.2012.138
[17]	Buscemi L, Ramonet D, Klingberg F, Formey A, Smith-Clerc J, Meister J J, Hinz B. 2011. The single-molecule mechanics of the latent TGF-beta1 complex. Current Biology, 21: 2046-2054. doi: 10.1016/j.cub.2011.11.037
[18]	Buschmann M D, Gluzband Y A, Grodzinsky A J, Hunziker E B. 1995. Mechanical compression modulates matrix biosynthesis in chondrocyte/agarose culture. Journal of Cell Science, 108: 1497-1508.
[19]	Buxboim A, Ivanovska I L, Discher D E. 2010. Matrix elasticity, cytoskeletal forces and physics of the nucleus: how deeply do cells 'feel' outside and in? Journal of Cell Science, 123: 297-308.
[20]	Carey S P, D'Alfonso T M, Shin S J, Reinhart-King C A. 2012. Mechanobiology of tumor invasion: Engineering meets oncology. Critical Reviews in Oncology Hematology, 83: 170-183. doi: 10.1016/j.critrevonc.2011.11.005
[21]	Chachisvilis M, Zhang Y L, Frangos J A. 2006. G protein-coupled receptors sense fluid shear stress in endothelial cells. Proceedings of the National Academy of Sciences of the United States of America, 103: 15463-15468. doi: 10.1073/pnas.0607224103
[22]	Chada S R, Hollenbeck P J. 2003. Mitochondrial movement and positioning in axons: The role of growth factor signaling. The Journal of Cell Biology, 206: 1985-1992.
[23]	Chalut K J, Hopfler M, Lautenschlager F, Boyde L, Chan C J, Ekpenyong A, Martinez-Arias A, Guck J. 2012. Chromatin decondensation and nuclear softening accompany Nanog downregulation in embryonic stem cells. Biophysical Journal, 103: 2060-2070. doi: 10.1016/j.bpj.2012.10.015
[24]	Chasis J A, Mohandas N. 2008. Erythroblastic islands: niches for erythropoiesis. Blood, 112: 470-478. doi: 10.1182/blood-2008-03-077883
[25]	Chaudhari A A, Seol J W, Kim S, Lee Y, Kang H, Kim I, Kim N, Park S Y. 2007. Reactive oxygen species regulate Bax translocation and mitochondrial transmembrane potential, a possible mechanism for enhanced TRAIL-induced apoptosis by CCCP. Oncology Report, 18: 71-76.
[26]	Chen C Y, Byrne H M, King J R. 2001. The influence of growth-induced stress from the surrounding medium on the development of multicell spheroids. Journal of Mathematical Biology, 43: 191-220. doi: 10.1007/s002850100091
[27]	Cheng H, Lederer W J, Cannell M B. 1993. Calcium sparks: Elementary events underlying excitation contraction coupling in heart muscle. Science, 262: 740-744. doi: 10.1126/science.8235594
[28]	Cherbuin T, Movahednia M M, Toh W S, Cao T. 2015. Investigation of human embryonic stem cell-derived keratinocytes as an in vitroresearch model for mechanical stress dynamic response. Stem Cell Reviews and Reports, 11: 460-473.
[29]	Chowdhury F, Li Y, Poh Y C, Yokohama-Tamaki T, Wang N, Tanaka T S. 2010a. Soft Substrates Promote Homogeneous Self-Renewal of Embryonic Stem Cells via Downregulating Cell-Matrix Tractions. PLoS One, 5: e15655. doi: 10.1371/journal.pone.0015655
[30]	Chowdhury F, Na S, Li D, Poh Y C, Tanaka T S, Wang F, Wang N. 2010b. Material properties of the cell dictate stress-induced spreading and differentiation in embryonic stem cells. Nature Materials, 9: 82-88. doi: 10.1038/nmat2563
[31]	Chrzanowska-Wodnicka M, Burridge K. 1996. Rho-stimulated contractility drives the formation of stress fibers and focal adhesions. Journal of Cell Biology, 133: 1403-1415. doi: 10.1083/jcb.133.6.1403
[32]	Conway A, Schaffer D V. 2012. Biophysical regulation of stem cell behavior within the niche. Stem Cell Research & Therapy, 3: 1-7.
[33]	Coughlin M F, Stamenović D. 1997. A tensegrity structure with buckling compression elements: application to cell mechanics. Journal of Applied Mechanic, 64: 480-486. doi: 10.1115/1.2788918
[34]	Coughlin M F, Stamenović D. 1998. A tensegrity model of the cytoskeleton in spread and round cells. Journal of Biomechanical Engineering, 120: 770-777. doi: 10.1115/1.2834892
[35]	Coughlin M F, StamenovićD. 2003. A prestressed cable network model of the adherent cell cytoskeleton. Biophysical journal, 84: 1328-1336. doi: 10.1016/S0006-3495(03)74948-0
[36]	Dalby M J, Gadegaard N, Tare R, Andar A, Riehle M O, Herzyk P, Wilkinson C D, Oreffo R O. 2007. The control of human mesenchymal cell differentiation using nanoscale symmetry and disorder. Nature Materials, 6: 997-1003. doi: 10.1038/nmat2013
[37]	D'Souza-Schorey C, Boettner B, Aelst L V. 1998. Rac regulates integrin-mediated spreading and increased adhesion of T lymphocytes. Molecular and Cellular Biology, 18: 3936-3946. doi: 10.1128/MCB.18.7.3936
[38]	Darin N, Kollberg G, Moslemi A, Tulinius M, Holme E, Grönlund M A, Andersson S, Oldfors A. 2006. Mitochondrial myopathy with exercise intolerance and retinal dystrophy in a sporadic patient with a G583A mutation in the mt tRNA ^phe gene. Neuromuscular Disorders, 16: 504-506. doi: 10.1016/j.nmd.2006.05.010
[39]	de Giorgi F, Lartigue L, Ichas F. 2000. Electrical coupling and plasticity of the mitochondrial network. Cell Calcium, 28: 365-370. doi: 10.1054/ceca.2000.0177
[40]	Desprat N, Supatto W, Pouille P A, Beaurepaire E, Farge E. 2008. Tissue deformation modulates twist expression to determine anterior midgut differentiation in Drosophila embryos. Developmental Cell, 15: 470-477. doi: 10.1016/j.devcel.2008.07.009
[41]	Di Bucchianico S, Poma A M, Giardi M F, Di Leandro L, Valle F, Biscarini F, Botti D. 2011. Atomic force microscope nanolithography on chromosomes to generate single-cell genetic probes. Journal of Nanobiotechnology, 9: 511.
[42]	Dingal P C, Bradshaw A M, Cho S, Raab M, Buxboim A, Swift J, Discher D E. 2015. Fractal heterogeneity in minimal matrix models of scars modulates stiff-niche stem-cell responses via nuclear exit of a mechanorepressor. Nature Materials, 14: 951-960. doi: 10.1038/nmat4350
[43]	Discher D E, Boal D H, Boey S K. 1998. Simulations of the erythrocyte cytoskeleton at large deformation. ii. micropipette aspiration. Biophysical Journal, 75: 1584-1597.
[44]	Discher D E, Janmey P, Wang Y L. 2005. Tissue cells feel and respond to the stiffness of their substrate. Science, 310: 1139-1143. doi: 10.1126/science.1116995
[45]	Du J, Chen X, Liang X, Zhang G, Xu J, He L, Zhan Q, Feng X Q, Chien S, Yang C. 2011. Integrin activation and internalization on soft ECM as a mechanism of induction of stem cell differentiation by ECM elasticity. Proceedings of the National Academy of Sciences of the United States of America, 108: 9466-9471. doi: 10.1073/pnas.1106467108
[46]	Dupont S, Morsut L, Aragona M, Enzo E, Giulitti S, Cordenonsi M, Zanconato F, Le Digabel J, Forcato M, Bicciato S, Elvassore N, Piccolo S. 2011. Role of YAP/TAZ in mechanotransduction. Nature, 474: 179-183. doi: 10.1038/nature10137
[47]	Elbediwy A, Vincent-Mistiaen Z I, Thompson B J. 2016. YAP and TAZ in epithelial stem cells: A sensor for cell polarity, mechanical forces and tissue damage. Bioessays, 38: 644-653. doi: 10.1002/bies.v38.7
[48]	Engler A J, Sen S, Sweeney H L, Discher D E. 2006. Matrix elasticity directs stem cell lineage specification. Cell, 126: 677-689. doi: 10.1016/j.cell.2006.06.044
[49]	Evans E, Yeung A. 1989. Apparent viscosity and cortical tension of blood granulocytes determined by micropipet aspiration. Biophysical Journal, 56: 151. doi: 10.1016/S0006-3495(89)82660-8
[50]	Eyal-Giladi H, Fabian B. 1980. Axis determination in uterine chick blastocysts under changing spatial positions during the sensitive period of polarity. Developmental Biology, 77: 228-232. doi: 10.1016/0012-1606(80)90470-4
[51]	Fabry B, Maksym G N, Butler J P, Glogauer M, Navajas D, Fredberg J J. 2001. Scaling the microrheology of living cells. Physical Review Letters, 87: 148102. doi: 10.1103/PhysRevLett.87.148102
[52]	Fabry B, Maksym G N, Butler J P, Glogauer M, Navajas D, Taback N A, Millet E J, Fredberg J J. 2003. Time scale and other invariants of integrative mechanical behavior in living cells. Physical Review E, 68: 041914. doi: 10.1103/PhysRevE.68.041914
[53]	Farge E. 2003. Mechanical induction of twist in the Drosophila foregut/stomodeal primordium. Current Biology, 13: 1365-1377. doi: 10.1016/S0960-9822(03)00576-1
[54]	Friedland J C, Lee M H, Boettiger D. 2009. Mechanically activated integrin switch controls alpha5beta1 function. Science, 323: 642-644. doi: 10.1126/science.1168441
[55]	Foty R A, Steinberg M S. 2005. The differential adhesion hypothesis: a direct evaluation. Developmental Biology, 278: 255-263. doi: 10.1016/j.ydbio.2004.11.012
[56]	Fu J, Wang Y K, Yang M T, Desai R A, Yu X, Liu Z, Chen C S. 2010. Mechanical regulation of cell function with geometrically modulated elastomeric substrates. Nature Methods, 7: 733-736. doi: 10.1038/nmeth.1487
[57]	Galluzzi L, Larochette N, Zamzami N, Kroemer G. 2006. Mitochondria as therapeutic targets for cancer chemotherapy. Oncogene, 25: 4812-4830. doi: 10.1038/sj.onc.1209598
[58]	Geiger B, Bershadsky A. 2002. Exploring the neighborhood: adhesion-coupled cell mechanosensors. Cell, 110: 139-142. doi: 10.1016/S0092-8674(02)00831-0
[59]	Gibson L J, Ashby M F. 1982. The mechanics of three-dimensional cellular materials//Proceedings of the Royal Society of London A: Mathematical, Physical and Engineering Sciences, 382: 43-59.
[60]	Grashoff C, Hoffman B D, Brenner M D, Zhou R, Parsons M, Yang M T, McLean M A, Sligar S G, Chen C S, Ha T, Schwartz M A. 2010. Measuring mechanical tension across vinculin reveals regulation of focal adhesion dynamics. Nature, 466: 263-266. doi: 10.1038/nature09198
[61]	Gorsich S W, Shaw J M. 2004. Importance of mitochondrial dynamics during meiosis and sporulation. Molecular Biology of the Cell, 15: 4369-4381. doi: 10.1091/mbc.E03-12-0875
[62]	Guilak F, Jones W R, Ting-Beall H P, Lee G M. 1999. The deformation behavior and mechanical properties of chondrocytes in articular cartilage. Osteoarthritis and Cartilage, 7: 59-70. doi: 10.1053/joca.1998.0162
[63]	Guilluy C, Swaminathan V, Garcia-Mata R, O'Brien E T, Superfine R, Burridge K. 2011. The Rho GEFs LARG and GEF-H1 regulate the mechanical response to force on integrins. Nature Cell Biology, 13: 722-727.
[64]	Han J, He H. 2015. Expression and function of osteogenic genes runt-related transcription factor 2 and osterix in orthodontic tooth movement in rats. International Journal of Clinical and Experimental Pathology, 8: 11895-11900.
[65]	Han Y, Wang L, Yao Q P, Zhang P, Liu B, Wang G L, Shen B R, Cheng B, Wang Y, Jiang Z L, Qi Y X. 2015. Nuclear envelope proteins Nesprin2 and LaminA regulate proliferation and apoptosis of vascular endothelial cells in response to shear stress. Biochimica et Biophysica Acta, 1853: 1165-1173. doi: 10.1016/j.bbamcr.2015.02.013
[66]	Harden N. 2002. Signaling pathways directing the movement and fusion of epithelial sheets: lessons from dorsal closure in Drosophila. Differentiation, 70: 181-203. doi: 10.1046/j.1432-0436.2002.700408.x
[67]	Harris A K, Wild P, Stopak D. 1980. Silicone rubber substrata: a new wrinkel in the study of cell locomotion. Science, 208: 177-179. doi: 10.1126/science.6987736
[68]	Heisenberg C P, Bellaiche Y. 2013. Forces in tissue morphogenesis and patterning. Cell, 153: 948-962. doi: 10.1016/j.cell.2013.05.008
[69]	Henderson J H, Cater D R. 2002. Mechanical induction in limb morphogenesis: the role of growth-generated strain and pressures. Bone, 31: 645-653. doi: 10.1016/S8756-3282(02)00911-0
[70]	Hochmuth R M, Ting-Beall H P, Beaty B B, Needham D, Tran-Son-Tay R. 1993. Viscosity of passive human neutrophils undergoing small deformations. Biophysical Journal, 64: 1596. doi: 10.1016/S0006-3495(93)81530-3
[71]	Hoffman B D, Grashoff C, Schwartz M A. 2011. Dynamic molecular processes mediate cellular mechanotransduction. Nature, 475: 316-323. doi: 10.1038/nature10316
[72]	Horner V L, Wolfner M F. 2008. Mechanical stimulation by osmotic and hydrostatic pressure activates Drosophila oocytes in vitro in a calcium-dependent manner. Developmental Biology, 316: 100-109. doi: 10.1016/j.ydbio.2008.01.014
[73]	Huang H, Kamm R D, Lee R T. 2004. Cellmechanicsandmechanotransduction: pathways, probes, and physiology. American Journal of Physiology-Cell Physiology, 287: C1-C11. doi: 10.1152/ajpcell.00559.2003
[74]	Huang C, Holfeld J, Schaden W, Orgill D, Ogawa R. 2013. Mechanotherapy: revisiting physical therapy and recruiting mechanobiology for a new era in medicine. Trends in Molecular Medicine, 19: 555-564. doi: 10.1016/j.molmed.2013.05.005
[75]	Hutson M S, Tokutake Y, Chang M S, Bloor J W, Venakides S, Kiehart D P, Edwards G S. 2003. Forces for morphogenesis investigated with laser microsurgery and quantitative modeling. Science, 300: 145-149. doi: 10.1126/science.1079552
[76]	Ingber D E. 1993. Cellular tensegrity: defining new rules of biological design that govern the cytoskeleton. Journal of Cell Science, 104: 613-627.
[77]	Ingber D E. 2003. Tensegrity i. cell structure and hierarchical systems biology. Journal of cell science, 116: 1157-1173.
[78]	Ingber D E. 2000. Opposing views on tensegrity as a structural framework for understanding cell mechanics. Journal of Applied Physiology, 89: 1663-1678.
[79]	Ingber D E. 2006. Cellular mechanotransduction: putting all the pieces together again. Faseb Journal, 20: 811-827. doi: 10.1096/fj.05-5424rev
[80]	Jakkaraju S, Zhe X, Pan D, Choudhury R, Schuger L. 2005. TIPs are tension-responsive proteins involved in myogenic versus adipogenic differentiation. Developmental Biology, 9: 39-49.
[81]	Janmey P A, McCulloch C A. 2007. Cell mechanics: integrating cell responses to mechanical stimuli. Annual Review of Biomedical Engineering, 9: 1-34. doi: 10.1146/annurev.bioeng.9.060906.151927
[82]	Jen C J, Jhiang S J, Chen H I. 2000. Invited review: effects of flow on vascular endothelial intracellular calcium signaling of rat aortas ex vivo. Journal of Applied Physiology, 89: 1657-1662.
[83]	Johnson C P, Tang H Y, Carag C, Speicher D W, Discher D E. 2007. Forced unfolding of protein swith in cells. Science, 317: 663-666. doi: 10.1126/science.1139857
[84]	Jones W R, Ting-Beall H P, Lee G M, Kelley S S, Hochmuth R M, Guilak F. 1999. Alterations in the young's modulus and volumetric properties of chondrocytes isolated from normal and osteoarthritic human cartilage. Journal of biomechanics, 32: 119-127. doi: 10.1016/S0021-9290(98)00166-3
[85]	Jufri N F, Mohamedali A, Avolio A, Baker M S. 2015. Mechanical stretch: physiological and pathological implications for human vascular endothelial cells. Vascular Cell, 7: 8. doi: 10.1186/s13221-015-0033-z
[86]	Kahn J, Shwartz Y, Blitz E, Krief S, Sharir A, Breitel D A, Rattenbach R, Relaix F, Maire P, Rountree R B, Kingsley D M, Zelzer E. 2009. Muscle contraction is necessary to maintain joint progenitor cell fate. Developmental Cell, 16: 734-743. doi: 10.1016/j.devcel.2009.04.013
[87]	Kan H C, Shyy W, Udaykumar H S, Vigneron P, Tran-Son-Tay R. 1999. Effects of nucleus on leukocyte recovery. Annals of Biomedical Engineering, 27: 648-655. doi: 10.1114/1.214
[88]	Karbowski M, Youle RJ. 2003. Dynamics of mitochondrial morphology in healthy cells and during apoptosis. Cell Death and Differentiation, 10: 870-880. doi: 10.1038/sj.cdd.4401260
[89]	Kasza K E, Rowat A C, Liu J, Angelini T E, Brangwynne C P, Koenderink G H, Weitz D A. 2007. The cell as a material. Current Opinion in Cell Biology, 19: 101-107. doi: 10.1016/j.ceb.2006.12.002
[90]	Khacho M, Clark A, Svoboda D S, Azzi J, MacLaurin J G, Meghaizel C, Sesaki H, Lagace D C, Germain M, Harper M E, Park D S, Slack R S. 2016. Mitochondrial dynamics impacts stem cell identity and fate decisions by regulating a nuclear transcriptional program. Cell Stem Cell, 19: 232-247. doi: 10.1016/j.stem.2016.04.015
[91]	Kim M C, Kim C, Wood L, Neal D, Kamm R D, Asada H H. 2012. Integrating focal adhesion dynamics, cytoskeleton remodeling, and actin motor activity for predicting cell migration on 3d curved surfaces of the extracellular matrix. Integrative Biology, 4: 1386-1397. doi: 10.1039/c2ib20159c
[92]	Kim M C, Neal D M, Kamm R D, Asada H H. 2013. Dynamic modeling of cell migration and spreading behaviors on fibronectin coated planar substrates and micropatterned geometries. PLoS Computational Biology, 9: e1002926. doi: 10.1371/journal.pcbi.1002926
[93]	Kim T J, Joo C, Seong J, Vafabakhsh R, Botvinick E L, Berns M W, Palmer A E, Wang N, Ha T, Jakobsson E, Sun J, Wang Y. 2015. Distinct mechanisms regulating mechanical force-induced Ca ²⁺ signals at the plasma membrane and the ER in human MSCs. Elife, 4: e04876.
[94]	Kim T J, Seong J, Ouyang M, Sun J, Lu S, Hong J P, Wang N, Wang Y. 2009. Substrate rigidity regulates Ca ²⁺ oscillation via rhoa pathway in stem cells. Journal of Cellular Physiology, 218: 285-293. doi: 10.1002/jcp.v218:2
[95]	Knowles M K, Guenza M G, Capaldi R A, Marcus A H. 2002. Cytoskeletal-assisted dynamics of the mitochondrial reticulum in living cells. Proceedings of the National Academy of Sciences of the United States of America, 99: 14772-14777. doi: 10.1073/pnas.232346999
[96]	Krieg M, Arboleda-Estudillo Y, Puech P H, Kafer J, Graner F, Muller D J, Heisenberg C P. 2008. Tensile forces govern germ-layer organization in zebrafish. Nature Cell Biology, 10: 429-436. doi: 10.1038/ncb1705
[97]	Kris N D, Alexandre J S, Jan L. 2008. Nuclear Shape, Mechanics, and Mechanotransduction. Circulation Research, 102: 1307-1318. doi: 10.1161/CIRCRESAHA.108.173989
[98]	Kochav S M, Eyal-Giladi H. 1971. Bilateral symmetry in the chick embryo determination by gravity. Science, 171: 1027-1029. doi: 10.1126/science.171.3975.1027
[99]	Kondo M, Wagers A J, Manz M G, Prohaska S S, Scherer D C, Beilhack G F, Shizuru J A, Weissman I L. 2003. Annual Review of Immunology, 21: 759-806.
[100]	Lammerding J, Hsiao J, Schulze P C, Kozlov S, Stewart C L, Lee R T. 2005. Abnormal nuclear shape and impaired mechanotransduction in emerin-deficient cells. Journal of Cell Biology, 170: 781-791. doi: 10.1083/jcb.200502148
[101]	Lang M. 2007. Lighting up the mechanome. Bridge, 37: 11-16.
[102]	Lawrie C H, Gal S, Dunlop H M, Pushkaran B, Liggins A P, Pulford K, Banham A H, Pezzella F, Boultwood J, Wainscoat J S, Hatton C S, Harris A L. 2008. Detection of elevated levels of tumour-associated microRNAs in serum of patients with diffuse large B-cell lymphoma. British Journal of Haematology, 2008, 141: 672-675.
[103]	Lee D Y, Lee C I, Lin T E, Lim S H, Zhou J, Tseng Y C, Chien S, Chiu J J. 2012. Role of histone deacetylases in transcription factor regulation and cell cycle modulation in endothelial cells in response to disturbed flow. Proceedings of the National Academy of Sciences of the United States of America, 109: 1967-1972. doi: 10.1073/pnas.1121214109
[104]	Li D, Zhou J, Wang L, Shin M E, Su P, Lei X, Kuang H, Guo W, Yang H, Cheng L, Tanaka T S, Leckband D E, Reynolds A B, Duan E, Wang F. 2010. Integrated biochemical and mechanical signals regulate multifaceted human embryonic stem cell functions. The Journal of Cell Biology, 191: 631-644. doi: 10.1083/jcb.201006094
[105]	Li H, Chen J, Zhang Y, Sun S J, Tao Z L, Long M. 2010. Effects of oriented substrates on cell morphology, the cell cycle, and the cytoskeleton in ros 17/2.8 cells. Science China Life Sciences, 53: 1085-1091. doi: 10.1007/s11427-010-4057-6
[106]	Li J, Dao M, Lim C T, Suresh S. 2005. Spectrin-level modeling of the cytoskeleton and optical tweezers stretching of the erythrocyte. Biophysical Journal, 88: 3707-3719. doi: 10.1529/biophysj.104.047332
[107]	Li Y, Chu J S, Kurpinski K, Li X, Bautista D M, Yang L, Sung K L, Li S. 2011. Biophysical regulation of histone acetylation in mesenchymal stem cells. Biophysical Journal, 100: 1902-1909. doi: 10.1016/j.bpj.2011.03.008
[108]	Li Z, Gong Y W, Sun S J, Du Y, Lü D Y, Liu X F, Long M. 2013. Differential regulation of stiffness, topography, and dimension of substrates in rat mesenchymal stem cells. Biomaterials, 34: 7616-7625. doi: 10.1016/j.biomaterials.2013.06.059
[109]	Liang Z, Liu C, Li L, Xu P, Luo G, Ding M, Liang Q. 2016. Double-network hydrogel with tunable mechanical performance and biocompatibility for the fabrication of stem cells-encapsulated fibers and 3D assemble. Scientific Reports, 6: 33462. doi: 10.1038/srep33462
[110]	Lim C T, Zhou E H, Quek S T. 2006. Mechanical models for living cells--a review. Journal of Biomechanics, 39: 195-216. doi: 10.1016/j.jbiomech.2004.12.008
[111]	Liu H, Wen J, Xiao Y, Liu J, Hopyan S, Radisic M, Simmons C A, Sun Y. 2014a. In situ mechanical characterization of the cell nucleus by atomic force microscopy. ACS Nano, 8: 3821-3828. doi: 10.1021/nn500553z
[112]	Liu S Q. 1998. Influence of tensile strain on smooth muscle cell orientation in rat blood vessels. Journal of Biomechanical Engineering, 120: 313-320. doi: 10.1115/1.2797996
[113]	Liu X F, Vargas D A, Lü D Y, Zhang Y, Zaman M H, and Long M. 2014. Computational modeling of stem cell migration: A mini review. Cellular and Molecular Bioengineering, 7: 196-204. doi: 10.1007/s12195-014-0330-2
[114]	Long M, Wang Y R, Zheng H Q, Shang P, Duan E K, Lü D Y. 2015. Mechano-biological coupling of cellular responses to microgravity. Microgravity Science and Technology, 27: 505-514. doi: 10.1007/s12217-015-9464-7
[115]	Long M, Sato M, Lim C T, Wu J H, Adachi A, and Inoue Y. 2011. Advances in experiments and modeling in micro-and nano-biomechanics---A mini review. Cellular and Molecular Bioengineering, 4: 327-339. doi: 10.1007/s12195-011-0183-x
[116]	Lozoya O A, Gilchrist C L, Guilak F. 2016. Universally conserved relationships between nuclear shape and cytoplasmic mechanical properties in human stem cells. Scientific Reports, 6: 23047. doi: 10.1038/srep23047
[117]	Lucia U. 2015. Bioengineering thermodynamics: An engineering science for thermodynamics of biosystems. International Journal of Thermophysics, 18: 254-265.
[118]	Lü D Y, Li Z, Gao Y X, Huo H, Kang Y Y, Chen J, Sun S J, Luo X D, Long M. 2013. Asymmetric migration of human keratinocytes under mechanical stretch and cocultured fibroblasts in a wound repairing model. PLoS One, 8: e74563. doi: 10.1371/journal.pone.0074563
[119]	Lü D Y, Luo C H, Zhang C, Li Z, Long M. 2014. Differential regulation of morphology and stemness of mouse embryonic stem cells by substrate stiffness and topography. Biomaterials, 35: 3945-3955. doi: 10.1016/j.biomaterials.2014.01.066
[120]	Lü D Y, Li Z, Gao Y X, Luo C H, Zhang F, Zheng L, Wang J W, Sun S J, Long M. 2016. β1 integrin signaling in asymmetric migration of keratinocytes under mechanical stretch in a co-cultured wound repair model. Biomedical Engneering Online. 15: 605-620.
[121]	MacKenna D A, Dolfi F, Vuori K, Ruoslahti E. 1998. Extracellular signal-regulated kinase and c-Jun NH2-terminal kinase activation by mechanical stretch is integrin-dependent and matrix-specific in rat cardiac fibroblasts. Journal Clinical Investigation, 101: 301-310. doi: 10.1172/JCI1026
[122]	MacQueen L, Chebotarev O, Simmons C A, Sun Y. 2012. Miniaturized platform with on-chip strain sensors for compression testing of arrayed materials. Lab on a Chip, 12: 4178-4184. doi: 10.1039/c2lc40670e
[123]	Mahaffy R E, Shih C K, MacKintosh F C, Käs J. 2000. Scanning probe-based frequency-dependent microrheology of polymer gels and biological cells. Physical Review Letters, 85: 880. doi: 10.1103/PhysRevLett.85.880
[124]	Mammoto T, Mammoto A, Torisawa Y S, Tat T, Gibbs A, Derda R, Mannix R, de Bruijn M, Yung C W, Huh D, Ingber D E. 2011. Mechanochemical control of mesenchymal condensation and embryonic tooth organ formation. DevelopmentalCell, 21: 758-769.
[125]	Marella S, Udaykumar H S. 2004. Computational analysis of the deformability of leukocytes modeled with viscous and elastic structural components. Physics of Fluids, 16: 244-264. doi: 10.1063/1.1629691
[126]	Martin A C, Kaschube M, Wieschaus E F. 2009. Pulsed contractions of an actin-myosin network drive apical constriction. Nature, 457: 495-499. doi: 10.1038/nature07522
[127]	Matthews B D, Overby D R, Mannix R, Ingber D E. 2006. Cellular adaptation to mechanical stress: role of integrins, Rho, cytoskeletal tension and mechanosensitive ion channels. Journal of Cell Science, 119: 508-518. doi: 10.1242/jcs.02760
[128]	Meeusen S, McCaffery M, Nunnari J. 2004. Mitochondrial fusion intermediates revealed in vitro. Science, 305: 1747-1752. doi: 10.1126/science.1100612
[129]	Miralles F, Posern G, Zaromytidou A I, Treisman R. 2003. Actin dynamics control SRF activity by regulation of its coactivator MAL. Cell, 113: 329-342. doi: 10.1016/S0092-8674(03)00278-2
[130]	Mitra S K, Hanson D A, Schlaepfer D D. 2005. Focal adhesion kinase: in command and control of cell motility. Nature Reviews Molecular Cell Biology, 6: 56-68. doi: 10.1038/nrm1549
[131]	Miyamoto S, Teramoto H, Coso O A, Gutkind J S, Burbelo P D, Akiyama S K, Yamada K M. 1995. Integrin function: molecular hierarchies of cytoskeletal and signaling molecules. Journal of Cell Biology, 131: 791-805. doi: 10.1083/jcb.131.3.791
[132]	Moore S W, Roca-Cusachs P, Sheetz M P. 2010. Stretchy proteins on stretchy substrates: the important elements of integrin-mediated rigidity sensing. Developmental Cell, 19: 194-206. doi: 10.1016/j.devcel.2010.07.018
[133]	Moraes C, Likhitpanichkul M, Lam C J, Beca B M, Sun Y, Simmons C A. 2013. Microdevice array-based identification of distinct mechanobiological response profiles in layer-specific valve interstitial cells. Integrative Biology, 5: 673-680. doi: 10.1039/c3ib20254b
[134]	N'Dri N A, Shyy W, Tran-Son-Tay R. 2003. Computational modeling of cell adhesion and movement using a continuum-kinetics approach. Biophysical Journal, 85: 2273-2286. doi: 10.1016/S0006-3495(03)74652-9
[135]	Na S, Collin O, Chowdhury F, Tay B, Ouyang M, Wang Y, Wang N. 2008. Rapid signal transduction in living cells is a unique feature of mechanotransduction. Proceedings of the National Academy of Sciences of the United States of America, 105: 6626-6631. doi: 10.1073/pnas.0711704105
[136]	Nix D A, Beckerle M C. 1997. Nuclear-cytoplasmic shuttling of the focal contact protein, zyxin: A potential mechanism for communication between sites of cell adhesion and the nucleus. Journal of Cell Biology, 138: 1139-1147. doi: 10.1083/jcb.138.5.1139
[137]	Novochadov V V, Bovol'skaya K A, Lipnitzkaya S A, Perevalova E V, Shuvalova E Y, Zagrebina Z N, Zaytzev V G. 2014. Different phenotype of chondrocytes in articular c'artilage: Mapping, possible mechanisms, and impact to implant healing. European Journal of Molecular Biotechology, 6: 210-222.
[138]	Obataya I, Nakamura C, Han S, Nakamura N, Miyake J. 2005. Nanoscale operation of a living cell using an atomic force microscope with a nanoneedle. Nano letters, 5: 27-30. doi: 10.1021/nl0485399
[139]	Okada Y, Nonaka S, Tanaka Y, Saijoh Y, Hamada H, Hirokawa N. 1999. Abnormal nodal flow precedes situs inversus in ivand invmice. Molecular Cell, 4:459-468. doi: 10.1016/S1097-2765(00)80197-5
[140]	Ozolek J A, Jane E P, Krowsoski L A, Sammak P J. 2007. Human embryonic stem cells (HSF-6) show greater proliferation and apoptoses when grown on glioblastoma cells than mouse embryonic fibroblasts at day 19 in culture: comparison of proliferation, survival, and neural differentiation on two different feeder cell types. Stem Cells Development, 16: 403-412. doi: 10.1089/scd.2006.0109
[141]	Pagliara S, Franze K, McClain C R, Wylde G W, Fisher C L, Franklin R J, Kabla A J, Keyser U F, Chalut K J. 2014. Auxetic nuclei in embryonic stem cells exiting pluripotency. Nature Materials, 13: 638-644. doi: 10.1038/nmat3943
[142]	Pardanaud L, Eichmann A. 2009. Stem cells: The stress of forming blood cells. Nature, 459: 1068-1069. doi: 10.1038/4591068a
[143]	Pajerowski J D, Dahl K N, Zhong F L, Sammak P J, Discher D E. 2007. Physical plasticity of the nucleus in stem cell differentiation. Proceedings of the National Academy of Sciences of the United States of America, 104: 15619-15624. doi: 10.1073/pnas.0702576104
[144]	Park W, Han Y, Kim S, Kim S, Cho K, Kim S. 2007. Antimycin A induces apoptosis in As4.1 juxtaglomerular cells. Cancer Letters, 251: 68-77.
[145]	Patwari P, Lee R T. 2008. Mechanical control of tissue morphogenesis. Circulation Research, 103: 234-243. doi: 10.1161/CIRCRESAHA.108.175331
[146]	Peric-Hupkes D, Meuleman W, Pagie L, Bruggeman S W, Solovei I, Brugman W, Gräf S, Flicek P, Kerkhoven R M, van Lohuizen M, Reinders M, Wessels L, van Steensel B. 2010. Molecular maps of the reorganization of genome-nuclear lamina interactions during differentiation. Molecular Cell, 38: 603-613. doi: 10.1016/j.molcel.2010.03.016
[147]	Petersen N O, McConnaughey W B, Elson E L. 1982. Dependence of locally measured cellular deformability on position on the cell, temperature, and cytochalasin b. Proceedings of the National Academy of Sciences, 79: 5327-5331. doi: 10.1073/pnas.79.17.5327
[148]	Poh Y C, Chen J, Hong Y, Yi H, Zhang S, Chen J, Wu D C, Wang L, Jia Q, Singh R, Yao W, Tan Y, Tajik A, Tanaka T S, Wang N. 2014. Generation of organized germ layers from a single mouse embryonic stem cell. Nature Communications, 5: 4000.
[149]	Pouille P A, Ahmadi P, Brunet A C, Farge E. 2009. Mechanical signals trigger Myosin-II redistribution and mesoderm invagination in Drosophila embryos. Science Signalling, 2: ra16.
[150]	Pourati J, Maniotis A, Spiegel D, Schaffer J L, Butler J P, Fredberg J J, Ingber D E, 1998. Stamenović D, Wang N. Is cytoskeletal tension a major determinant of cell deformability in adherent endothelial cells? American Journal of Physiology-Cell Physiology. 274: C1283-1289.
[151]	Price L S, Leng J, Schwartz M A, Bokoch G M. 1998. Activation of rac and cdc42 by integrins mediates cell spreading. Molecular Biology of the Cell, 9: 1863-1871. doi: 10.1091/mbc.9.7.1863
[152]	Pritz T. 1996. Analysis of four-parameter fractional derivative model of real solid materials. Journal of Sound and Vibration, 195: 103-115. doi: 10.1006/jsvi.1996.0406
[153]	Puig-De-Morales M, Grabulosa M, Alcaraz J, Mullol J, Maksym G N, Fredberg J J, Navajas V. 2001. Measurement of cell microrheology by magnetic twisting cytometry with frequency domain demodulation. Journal of Applied Physiology, 91: 1152-1159.
[154]	Qi Y X, Jiang J, Jiang X H, Wang X D, Ji S Y, Han Y, Long D K, Shen B R, Yan Z Q, Chien S, Jiang Z L. 2011. PDGF-BB and TGF- β1 on cross-talk between endothelial and smooth muscle cells in vascular remodeling induced by low shear stress. Proceedings of the National Academy of Sciences of the United States of America, 108: 1908-1913. doi: 10.1073/pnas.1019219108
[155]	Qi Y X, Yao Q P, Huang K, Shi Q, Zhang P, Wang G L, Han Y, Bao H, Wang L, Li H P, Shen B R, Wang Y, Chien S, Jiang Z L. 2016. Nuclear envelope proteins modulate proliferation of vascular smooth muscle cells during cyclic stretch application. Proceedings of the National Academy of Sciences of the United States of America, 113: 5293-5298. doi: 10.1073/pnas.1604569113
[156]	Radmacher M. 1997. Measuring the elastic properties of biological samples with the afm. IEEE Engineering in Medicine and Biology Magazine, 16: 47-57.
[157]	Rauskolb C, Pan G, Reddy B V, Oh H, Irvine K D. 2011. Zyxin links fat signaling to the hippo pathway. PLoS Biology, 9: e1000624. doi: 10.1371/journal.pbio.1000624
[158]	Resnick A, Hopfer U. 2007. Force-response considerations in ciliary mechanosensation. Biophysical Journal, 93: 1380-1390. doi: 10.1529/biophysj.107.105007
[159]	Rizzo V, Sung A, Schnitzer J E. 1998. Rapid mechanotransduction in situ at the luminal cell surface of vascular endothelium and its caveolae. The Journal of Biological Chemistry, 273: 26323-26329. doi: 10.1074/jbc.273.41.26323
[160]	Rodrigues I, Sanches J. 2008. Temporal 2D reconstruction of cell nucleus from fluorescence confocal Microscopy images with anisotropic filtering. Conference Proceedings IEEE Engineering in Medicine and Biology Society, 2008: 2197-2200.
[161]	Rolin G L, Binda D, Tissot M, Viennet C, Saas P, Muret P, Humbert P. 2014. In vitro study of the impact of mechanical tension on the dermal fibroblast phenotype in the context of skin wound headling. Journal of Biomechanics, 47: 3555-3561. doi: 10.1016/j.jbiomech.2014.07.015
[162]	Rosowski K A, Mertz A F, Norcross S, Dufresne E R, Horsley V. 2015. Edges of human embryonic stem cell colonies display distinct mechanical properties and differentiation potential. Scientific Reports, 5: 14218. doi: 10.1038/srep14218
[163]	Salmeen I, Zacmanidis P, Jesion G, Feldkamp LA. 1985. Motion of mitochondria in cultured cells quantified by analysis of digitized images. Biophysical Journal, 48: 681-686. doi: 10.1016/S0006-3495(85)83825-X
[164]	Sanders M C, Way M, Sakai J, Matsudaira P. 1996. Characterization of the actin cross-linking properties of the scruin-calmodulin complex from the acrosomal process of Limulu sperm. The Journal of Biological Chemistry, 271: 2651-2657. doi: 10.1074/jbc.271.5.2651
[165]	Sansores-Garcia L, Bossuyt W, Wada K, Yonemura S, Tao C, Sasaki H, Halder G. 2011. Modulating F-actin organization induces organ growth by affecting the Hippo pathway. The EMBO Journal, 30: 2325-2335. doi: 10.1038/emboj.2011.157
[166]	Satcher R L Jr, Dewey C F Jr. 1996. Theoretical estimates of mechanical properties of the endothelial cell cytoskeleton. Biophysical Journal, 71: 109. doi: 10.1016/S0006-3495(96)79206-8
[167]	Satcher R, Dewey C F, Hartwig J H. 1997. Mechanical remodeling of the endothelial surface and actin cytoskeleton induced by fluid flow. Microcirculation, 4: 439-453. doi: 10.3109/10739689709146808
[168]	Sato M, Levesque M J, Nerem R M. 1987. Micropipette aspiration of cultured bovine aortic endothelial cells exposed to shear stress. Arteriosclerosis, Thrombosis, and Vascular Biology, 7: 276-286. doi: 10.1161/01.ATV.7.3.276
[169]	Schmid-Schönbein G W, Sung K L, Tözeren H, Skalak R, Chien S. 1981. Passive mechanical properties of human leukocytes. Biophysical Journal, 36: 243. doi: 10.1016/S0006-3495(81)84726-1
[170]	Sem D S, Yu L, Coutts S M, Jack R. 2001. Object-oriented approach to drug design enabled by NMR SOLVE: First real-time structural tool for characterizing protein-ligand interactions. Journal of Cellular Biochemistry, 84: 99-105. doi: 10.1002/(ISSN)1097-4644
[171]	Sesaki H, Jensen RE. 1999. Division versus fusion: Dnm1p and Fzo1p antagonistically regulate mitochondrial shape. The Journal of Cell Biology, 147: 699-706. doi: 10.1083/jcb.147.4.699
[172]	Shieh A C, Athanasiou K A. 2002. Biomechanics of single chondrocytes and osteoarthritis. Critical Reviews in Biomedical Engineering, 30: 307-343. doi: 10.1615/CritRevBiomedEng.v30.i456
[173]	Shieh A C, Athanasiou K A. 2003. Principles of cell mechanics for cartilage tissue engineering. Annals of Biomedical Engineering, 31: 1-11. doi: 10.1114/1.1535415
[174]	Shin J H, Tam B K, Brau R R, Lang M J, Mahadevan L, Matsudaira P. 2007. Force of an actin spring. Biophysical Journal, 92: 3729-3733. doi: 10.1529/biophysj.106.099994
[175]	Shin J W, Discher D E. 2015. Blood and immune cell engineering: cytoskeletal contractility and nuclear rheology impact cell lineage and localization: biophysical regulation of hematopoietic differentiation and trafficking. Bioessays, 37: 633-642. doi: 10.1002/bies.v37.6
[176]	Shin J W, Spinler K R, Swift J, Chasis J A, Mohandas N, Discher D E. 2013. Lamins regulate cell trafficking and lineage maturation of adult human hematopoietic cells. Proceedings of the National Academy of Sciences of the United States of America, 110: 18892-18897. doi: 10.1073/pnas.1304996110
[177]	Shiroky A A, Volkov A V, Novochadov V V. 2014. Crucial processes' interaction during the renewal of articular cartilage: the mathematical modeling. European of Journal Molecular Biotechology, 4: 86-94. doi: 10.13187/issn.2310-6255
[178]	Shroff S G, Saner D R, Lal R. 1995. Dynamic micromechanical properties of cultured rat atrial myocytes measured by atomic force microscopy. American Journal of Physiology-Cell Physiology, 269: C286-C292.
[179]	Simpson C F, Kling J M. 1967. The mechanism of denucleation in circulating erythroblasts. Journal of Cell Biology, 35: 237-45. doi: 10.1083/jcb.35.1.237
[180]	Sims N R. 1990. Rapid isolation of metabolically active mitochondria from rat brain and subregions using percoll density gradient centrifugation. Journal of Neurochemistry, 55: 698-707. doi: 10.1111/jnc.1990.55.issue-2
[181]	Snelson K. 1996. Snelson on the tensegrity invention. International Journal of Space Structures, 11: 43-48. doi: 10.1177/026635119601-207
[182]	Song M J, Brady-Kalnay S M, McBride S H, Phillips-Mason P, Dean D, Tate ML. 2012. Mapping the mechanome of live stem cells using a novel method to measure local strain fields in situ at the fluid-cell interface. PLoS One, 7: e43601. doi: 10.1371/journal.pone.0043601
[183]	Staal J A, Dickson T C, Chung R S, Vickers J C. 2007. Cyclosporin-A treatment attenuates delayed cytoskeletal alterations and secondary axotomy following mild axonal stretch injury. Developmental Neurobiology, 67: 1831-1842. doi: 10.1002/(ISSN)1932-846X
[184]	Stamenović D, Ingber D E. 2002. Models of cytoskeletal mechanics of adherent cells. Biomechanics and Modeling in Mechanobiology, 1: 95-108. doi: 10.1007/s10237-002-0009-9
[185]	Stamenović D, Mijailovich S M, Tolic-N Ørrelykke I M, Chen J, Wang N. 2002. Cell prestress. II. Contribution of microtubules. American Journal of Physiology-Cell Physiology, 282: C617-624. doi: 10.1152/ajpcell.00271.2001
[186]	Sukharev S, Corey D P. 2004 .Mechanosensitive channels: multiplicity of families and gating paradigms. Science's STKE: Signal Transduction Knowledge Environment, 2004: re4.
[187]	Swift J, Ivanovska I L, Buxboim A, Harada T, Dingal P C, Pinter J, Pajerowski J D, Spinler K R, Shin J W, Tewari M, Rehfeldt F, Speicher D W, Discher D E. 2013. Nuclear lamin-A scales with tissue stiffness and enhances matrix-directed differentiation. Science, 341: 1240104. doi: 10.1126/science.1240104
[188]	Taguchi N, Ishihara M, Jofuku A, Oka T, Mihara K. 2007. Mitotic phosphorylation of dynamin-related GTPase Drp1 participates in mitochondrial fission. The Journal of Biological Chemistry, 282: 11521-11529. doi: 10.1074/jbc.M607279200
[189]	Tajik A, Zhang Y, Wei F, Sun J, Jia Q, Zhou W, Singh R, Khanna N, Belmont A S, Wang N. 2016. Transcription upregulation via force-induced direct stretching of chromatin. Nature Materials, 15: 1287-1296. doi: 10.1038/nmat4729
[190]	Tan J L, Tien J, Pirone D M, Gray D S, Bhadriraju K, Chen C S. 2003. Cells lying on a bed of microneedles: an approach to isolate mechanical force. Proceedings of the National Academy of Sciences of the United States of America, 100: 1484-1489. doi: 10.1073/pnas.0235407100
[191]	Tavakolinejad A, Rabbani M, Janmaleki M. 2015. Effects of hypergravity on adipose-derived stem cell morphology, mechanical property and proliferation. Biochemical and Biophysical Research Communications, 464: 473-479. doi: 10.1016/j.bbrc.2015.06.160
[192]	Theret D P, Levesque M J, Sato M, Nerem R M, Wheeler L T. 1988. The application of a homogeneous half-space model in the analysis of endothelial cell micropipette measurements. Journal of Biomechanical Engineering, 110: 190-199. doi: 10.1115/1.3108430
[193]	Titushkin I, Cho M. 2007. Modulation of cellular mechanics during osteogenic differentiation of human mesenchymal stem cells. Biophysical Journal, 93: 3693-3702 doi: 10.1529/biophysj.107.107797
[194]	Tran-Son-Tay R, Kan H C, Udaykumar H S, Damay E, Shyy W. 1998. Rheological modelling of leukocytes. Medical and Biological Engineering and Computing, 36: 246-250. doi: 10.1007/BF02510753
[195]	Tsai J W, Chen Y, Kriegstein A R, Vallee R B. 2005. LIS1 RNA interference blocks neural stem cell division, morphogenesis, and motility at multiple stages. Journal of Cell Biology, 170: 935-945. doi: 10.1083/jcb.200505166
[196]	Tschumperlin D J, Liu F, Tager A M. 2013. Biomechanical regulation of mesenchymal cell function. Current Opinion in Rheumatology, 25: 92-100. doi: 10.1097/BOR.0b013e32835b13cd
[197]	Tuleuova N, Lee J Y, Lee J, Ramanculov E, Zern M A, Revzin A. 2010. Using growth factor arrays and micropatterned co-cultures to induce hepatic differentiation of embryonic stem cells. Biomaterials, 31: 9221-9231. doi: 10.1016/j.biomaterials.2010.08.050
[198]	Uzer G, Fuchs R K, Rubin J, Thompson W R. 2016. Concise Review: Plasma and Nuclear Membranes Convey Mechanical Information to Regulate Mesenchymal Stem Cell Lineage. Stem Cells, 34:1455-1463. doi: 10.1002/stem.v34.6
[199]	van Loon JJ. 2009. Mechanomics and physicomics in gravisensing. Microgravity Science Technology, 21: 159-167.
[200]	Vogel V, Sheetz M. 2006. Local force and geometry sensing regulate cell functions. Nature Reviews Molecular Cell Biology, 7:265-275. doi: 10.1038/nrm1890
[201]	Volokh K Y, Vilnay O, Belsky M. 2000. Tensegrity architecture explains linear stiffening and predicts softening of living cells. Journal of Biomechanics, 3: 1543-1549.
[202]	Vorselen D, Roos W H, MacKintosh F C, Wuite G J, van Loon J J. 2014. The role of the cytoskeleton in sensing changes in gravity by nonspecialized cells. The FASEB Journal, 28: 536-547. doi: 10.1096/fj.13-236356
[203]	Wada K, Itoga K, Okano T, Yonemura S, Sasaki H. 2011. Hippo pathway regulation by cell morphology and stress fibers. Development, 138: 3907-3914. doi: 10.1242/dev.070987
[204]	Wagner O I, Lifshitz J, Janmey P A, Linden M, McIntosh T K, Leterrier J. 2003. Mechanisms of mitochondria-neurofilament interactions. The Journal of Neuroscience, 23: 9046-9058.
[205]	Wang H, Ip W, Boissy R, Grood E S. 1995. Cell orientation response to cyclically deformed substrates: experimental validation of a cell model. Journal of Biomechanics, 28: 1543-1552. doi: 10.1016/0021-9290(95)00101-8
[206]	Wang J W, Lü D Y, Mao D B, Long M. 2014. Mechanomics: an emerging field between biology and biomechanics. Protein & Cells, 5: 518-531.
[207]	Wang J G, Miyazu M, Matsushita E, Sokabe M, Naruse K. 2001. Uniaxial cyclic stretch induces focal adhesion kinase (FAK) tyrosine phosphorylation followed by mitogen-activated protein kinase (MAPK) activation. Biochemical and Biophysical Research Communications, 288: 356-361. doi: 10.1006/bbrc.2001.5775
[208]	Wang N, Naruse K, Stamenović D, Fredberg J J, Mijailovich S M, Tolic-N Ørrelykke I M, Polte T, Mannix R, Ingber D E. 2001. Mechanical behavior in living cells consistent with the tensegrity model. Proceedings of the National Academy of Sciences, 98: 7765-7770. doi: 10.1073/pnas.141199598
[209]	Wang N, Butler J P, Ingber D E. 1993. Mechanotransduction across the cell surface and through the cytoskeleton. Science, 260: 1124-1127. doi: 10.1126/science.7684161
[210]	Wang N, Stamenović D. 2000. Contribution of intermediate filaments to cell stiffness, stiffening, and growth. American Journal of Physiology-cell Physiology, 279: 188-194.
[211]	Wang N, Tolic-N Ørrelykke I M, Chen J, Mijailovich S M, Butler J P, Fredberg J J, StamenovićD. 2002. Cell prestress. I. Stiffness and prestress are closely associated in adherent contractile cells. American Journal of Physiology-Cell Physiology, 282: C606-616.
[212]	Wang N, Tytell J D, Ingber D E. 2009. Mechanotransduction at a distance: mechanically coupling the extracellular matrix with the nucleus. Nature Reviews Molecular Cell Biology, 10: 75-82. doi: 10.1038/nrm2594
[213]	Wang S Q, Xiao W M, Shan S C, Jiang C S, Chen M, Zhang Y, Lü S Q, Chen J, Zhang C M, Chen Q, Long M. 2012. Multi-patterned dynamics of mitochondrial fission and fusion in a living cell. PLoS One, 7: e19879. doi: 10.1371/journal.pone.0019879
[214]	Wang Y, Botvinick E L, Zhao Y, Berns M W, Usami S, Tsien R Y, Chien S. 2005. Visualizing the mechanical activation of Src. Nature, 434: 1040-1045. doi: 10.1038/nature03469
[215]	Wang Y K, Yu X, Cohen D M, Wozniak M A, Yang M T, Gao L, Eyckmans J, Chen C S. 2012. Bone morphogenetic protein-2-induced signaling and osteogenesis is regulated by cell shape, RhoA/ROCK, and cytoskeletal tension. Stem Cells and Development, 21: 1176-1186. doi: 10.1089/scd.2011.0293
[216]	Wang Y X, Shyy J Y, Chien S. 2008. Fluorescence proteins, live-cell imaging, and mechanobiology: Seeing is believing. Annual Review of Biomedical Engineering, 10: 1-38. doi: 10.1146/annurev.bioeng.010308.161731
[217]	Warren W E, Kraynik A M. 1997. Linear elastic behavior of a low-density kelvin foam with open cells. Journal of Applied Mechanics, 64: 787-794. doi: 10.1115/1.2788983
[218]	Weis W I. 2004. How to build a cell junction. Nature, 430: 513-514. doi: 10.1038/430513a
[219]	Watanabe K, Ueno M, Kamiya D, Nishiyama A, Matsumura M, Wataya T, Takahashi J B, Nishikawa S, Nishikawa S, Muguruma K, Sasai Y. 2007. A ROCK inhibitor permits survival of dissociated human embryonic stem cells. Nature Biotechnology, 25: 681-686. doi: 10.1038/nbt1310
[220]	Wernig F, Mayr M, Xu Q. 2003. Mechanical stretch-induced apoptosis in smooth muscle cells is mediated by beta1-integrin signaling pathways. Hypertension, 41: 903-911. doi: 10.1161/01.HYP.0000062882.42265.88
[221]	Wipff P J, Rifkin D B, Meister J J, Hinz B. 2007. Myofibroblast contraction activates latent TGF-beta1 from the extracellular matrix. Journal of Cell Biology, 179: 1311-1323. doi: 10.1083/jcb.200704042
[222]	Wong J J, Ritchie W, Ebner O A, Selbach M, Wong J W, Huang Y, Gao D, Pinello N, Gonzalez M, Baidya K, Thoeng A, Khoo T L, Bailey C G, Holst J, Rasko J E. 2013. Orchestrated intron retention regulates normal granulocyte differentiation. Cell, 154: 583-595. doi: 10.1016/j.cell.2013.06.052
[223]	Wuest S L, Richard S, Kopp S, Grimm D, Egli M. 2015. Simulated microgravity: critical review on the use of random positioning machines for mammalian cell culture. Biomed Research International, 2015: 971474.
[224]	Yaffe, M.P., Harata, D., Verde, F., Eddison, M., Toda, T., Nurse, P. 1996. Microtubules mediate mitochondrial distribution in fission yeast. Proceedings of the National Academy of Sciences of the United States of America, 93: 11664-11668.
[225]	Yang Y, Relan N K, Przywara D A, Schuger L. 1999. Embryonic mesenchymal cells share the potential for smooth muscle differentiation: myogenesis is controlled by the cell's shape. Development, 126: 3027-3033.
[226]	Yasuhiro S, Masako T, Dubin-Thaler B J, Cherniavskaya O, Sakai R, Tanaka S, Sheetz M P. 2006. Force sensing by mechanical extension of the Src family kinase substrate p130Cas. Cell, 127: 1015-1026. doi: 10.1016/j.cell.2006.09.044
[227]	Yeung A, Evans E. 1989. Cortical shell-liquid core model for passive flow of liquid-like spherical cells into micropipets. Biophysical Journal, 56: 139-149. doi: 10.1016/S0006-3495(89)82659-1
[228]	Zermati Y, Garrido C, Amsellem S, Fishelson S, Bouscary D, Valensi F, Varet B, Solary E, Hermine O. 2001. Caspase activation is required for terminal erythroid differentiation. The Journal of Experimental Medicine, 193: 247-254. doi: 10.1084/jem.193.2.247
[229]	Zhang C, Zhou L W, Zhang F, Lü D Y, Li N, Zheng L, Xu Y H, Li Z, Sun S J, Long M. 2016. Mechanical remodeling of normally-sized mammalian cells under a gravity vector. FASEB Journal, 2016 Nov 8. pii: fj.201600897RR.
[230]	Zhang L, Liu W, Zhao J, Ma X, Shen L, Zhang Y, Jin F, Jin Y. 2016. Mechanical stress regulates osteogenic differentiation and RANKL/OPG ratio in periodontal ligament stem cells by the Wnt/ β-catenin pathway. Biochimica et Biophysica Acta, 1860: 2211-2219. doi: 10.1016/j.bbagen.2016.05.003
[231]	Zhang W, Kong C W, Tong M H, Chooi W H, Huang N, Li R A, Chan B P. 2016. Maturation of human embryonic stem cell-derived cardiomyocytes (hESC-CMs) in 3D collagen matrix: Effects of niche cell supplementation and mechanical stimulation. Acta Biomaterialia, 24, pii: S1742-7061(16) 30654-7.
[232]	Zhu Z, Gan X, Fan H, Yu H. 2015. Mechanical stretch endows mesenchymal stem cells stronger angiogenic and anti-apoptotic capacities via NF κB activation. Biochemical and Biophysical Research Communications, 468: 601-605. doi: 10.1016/j.bbrc.2015.10.157