CoChat
Sign Up
Cited by 409Cellular Mechanics and InteractionsSkin and Cellular Biology ResearchCell Adhesion Molecules Research

Focal adhesions, stress fibers and mechanical tension

Open access
Burridge, Keith, Guilluy, Christophe·2016·Experimental Cell Research
DOI 10.1016/j.yexcr.2015.10.029PMC PMC4891215
Open this paper in CoChat — free accountOpen PDF

Abstract

Stress fibers and focal adhesions are complex protein arrays that produce, transmit and sense mechanical tension. Evidence accumulated over many years led to the conclusion that mechanical tension generated within stress fibers contributes to the assembly of both stress fibers themselves and their associated focal adhesions. However, several lines of evidence have recently been presented against this model. Here we discuss the evidence for and against the role of mechanical tension in driving the assembly of these structures. We also consider how their assembly is influenced by the rigidity of the substratum to which cells are adhering. Finally, we discuss the recently identified connections between stress fibers and the nucleus, and the roles that these may play, both in cell migration and regulating nuclear function.

The claims and statistics of this paper haven’t been checked yet.

Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.

A few free checks a day · papers already verified open instantly · results are public

Integrity signals

Registry

Crossref · 10.1016/j.yexcr.2015.10.029

Cites

4 of 4 references verified0 citation errors · 0 not checkable

Statistics

Claims the paper makes

The claims and statistics of this paper haven’t been checked yet.

Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.

A few free checks a day · papers already verified open instantly · results are public

Claims other papers make about this one

Each is checked against this paper’s own text: does it say what the citing paper says it does?

  • Contractile actin filament assembly, stress fiber assembly, and regulation of actin filament-based processes are molecular processes centrally involved in focal adhesion biology.

    “all of which are molecular processes centrally involved in focal adhesion biology (Burridge and Guilluy, 2016; Ciobanasu et al., 2012; Geiger et al., 2009; Wozniak et al., 2004).”

    From Alzheimer’s genetic risk factor CASS4 controls microglial focal adhesion biology and amyloidosis (2026) · cites this paper as Burridge and Guilluy, 2016

Self-check

Percentages, sums and reported differences re-computed from the paper’s own numbers.

The self-check needs the paper’s full text.

Cites

Flagged first, then by citations

Research works printed but not linked: 87

  1. [1]Burridge K Are stress fibres contractile? Nature 294 1981 691 692 7198718
  2. [2]Geiger B Bershadsky A Exploring the neighborhood: adhesion-coupled cell mechanosensors Cell 110 2002 139 142 12150922
  3. [3]Hoffman BD Grashoff C Schwartz MA Dynamic molecular processes mediate cellular mechanotransduction Nature 475 2011 316 323 21776077 PMC6449687
  4. [4]Case LB Waterman CM Integration of actin dynamics and cell adhesion by a three-dimensional, mechanosensitive molecular clutch Nat Cell Biol 17 2015 955 963 26121555 PMC6300998
  5. [5]Burridge K Wittchen ES The tension mounts: stress fibers as force-generating mechanotransducers J Cell Biol 200 2013 9 19 23295347 PMC3542796
  6. [6]Riveline D Zamir E Balaban NQ Schwarz US Ishizaki T Narumiya S Kam Z Geiger B Bershadsky AD Focal contacts as mechanosensors: externally applied local mechanical force induces growth of focal contacts by an mDia1-dependent and ROCK-independent mechanism J Cell Biol 153 2001 1175 1186 11402062 PMC2192034
  7. [7]Chrzanowska-Wodnicka M Burridge K Rho-stimulated contractility drives the formation of stress fibers and focal adhesions J Cell Biol 133 1996 1403 1415 8682874 PMC2120895
  8. [8]Choi CK Vicente-Manzanares M Zareno J Whitmore LA Mogilner A Horwitz AR Actin and alpha-actinin orchestrate the assembly and maturation of nascent adhesions in a myosin II motor-independent manner Nat Cell Biol 2008 PMC2827253 19160484
  9. [9]Stricker J Aratyn-Schaus Y Oakes PW Gardel ML Spatiotemporal constraints on the force-dependent growth of focal adhesions Biophys J 100 2011 2883 2893 21689521 PMC3123981
  10. [10]Oakes PW Beckham Y Stricker J Gardel ML Tension is required but not sufficient for focal adhesion maturation without a stress fiber template J Cell Biol 196 2012 363 374 22291038 PMC3275371
  11. [11]Thievessen I Thompson PM Berlemont S Plevock KM Plotnikov SV Zemljic-Harpf A Ross RS Davidson MW Danuser G Campbell SL Vinculin–actin interaction couples actin retrograde flow to focal adhesions, but is dispensable for focal adhesion growth J Cell Biol 202 2013 163 177 23836933 PMC3704983
  12. [12]Herman IM Crisona NJ Pollard TD Relation between cell activity and the distribution of cytoplasmic actin and myosin J Cell Biol 90 1981 84 91 7019223 PMC2111826
  13. [13]Small JV Rottner K Kaverina I Anderson KI Assembling an actin cytoskeleton for cell attachment and movement Biochim Et Biophys Acta 1404 1998 271 281 9739149
  14. [14]Hotulainen P Lappalainen P Stress fibers are generated by two distinct actin assembly mechanisms in motile cells J Cell Biol 173 2006 383 394 16651381 PMC2063839
  15. [15]Webb DJ Donais K Whitmore LA Thomas SM Turner CE Parsons JT Horwitz AF FAK-Src signalling through paxillin, ERK and MLCK regulates adhesion disassembly Nat Cell Biol 6 2004 154 161 14743221
  16. [16]Burnette DT Shao L Ott C Pasapera AM Fischer RS Baird MA Der Loughian C Delanoe-Ayari H Paszek MJ Davidson MW A contractile and counterbalancing adhesion system controls the 3D shape of crawling cells J Cell Biol 205 2014 83 96 24711500 PMC3987145
  17. [17]Khatau SB Hale CM Stewart-Hutchinson PJ Patel MS Stewart CL Searson PC Hodzic D Wirtz D A perinuclear actin cap regulates nuclear shape Proc Natl Acad Sci USA 106 2009 19017 19022 19850871 PMC2776434
  18. [18]Ridley AJ Hall A The small GTP-binding protein rho regulates the assembly of focal adhesions and actin stress fibers in response to growth-factors Cell 70 1992 389 399 1643657
  19. [19]Burridge K Chrzanowska-Wodnicka M Focal adhesions, contractility, and signaling Annu Rev Cell Dev Biol 12 1996 463 518 8970735
  20. [20]Pasapera AM Schneider IC Rericha E Schlaepfer DD Waterman CM Myosin II activity regulates vinculin recruitment to focal adhesions through FAK-mediated paxillin phosphorylation J Cell Biol 188 2010 877 890 20308429 PMC2845065
  21. [21]Even-Ram S Doyle AD Conti MA Matsumoto K Adelstein RS Yamada KM Myosin IIA regulates cell motility and actomyosin-microtubule crosstalk Nat Cell Biol 9 2007 299 309 17310241
  22. [22]Vicente-Manzanares M Zareno J Whitmore L Choi CK Horwitz AF Regulation of protrusion, adhesion dynamics, and polarity by myosins IIA and IIB in migrating cells J Cell Biol 176 2007 573 580 17312025 PMC2064016
  23. [23]Vicente-Manzanares M Horwitz AR Adhesion dynamics at a glance J Cell Sci 124 2011 3923 3927 22194302 PMC3244977
  24. [24]Ponti A Machacek M Gupton SL Waterman-Storer CM Danuser G Two distinct actin networks drive the protrusion of migrating cells Science 305 2004 1782 1786 15375270
  25. [25]Alexandrova AY Arnold K Schaub S Vasiliev JM Meister JJ Bershadsky AD Verkhovsky AB Comparative dynamics of retrograde actin flow and focal adhesions: formation of nascent adhesions triggers transition from fast to slow flow Plos One 3 2008 e3234 18800171 PMC2535565
  26. [26]Smilenov LB Mikhailov A Pelham RJ Marcantonio EE Gundersen GG Focal adhesion motility revealed in stationary fibroblasts Science 286 1999 1172 1174 10550057
  27. [27]Nobes CD Hall A Rho, Rac, and Cdc42 GTPases regulate the assembly of multimolecular focal complexes associated with actin stress fibers, lamelli-podia, and filopodia Cell 81 1995 53 62 7536630
  28. [28]Franke RP Grafe M Schnittler H Seiffge D Mittermayer C Drenckhahn D Induction of human vascular endothelial stress fibres by fluid shear stress Nature 307 1984 648 649 6537993
  29. [29]Balaban NQ Schwarz US Riveline D Goichberg P Tzur G Sabanay I Mahalu D Safran S Bershadsky A Addadi L Force and focal adhesion assembly: a close relationship studied using elastic micropatterned substrates. [see comments.] Nat Cell Biol 3 2001 466 472 11331874
  30. [30]Schwarz US Balaban NQ Riveline D Bershadsky A Geiger B Safran SA Calculation of forces at focal adhesions from elastic substrate data: the effect of localized force and the need for regularization Biophys J 83 2002 1380 1394 12202364 PMC1302237
  31. [31]Galbraith CG Yamada KM Sheetz MP The relationship between force and focal complex development J Cell Biol 159 2002 695 705 12446745 PMC2173098
  32. [32]Kuo JC Han X Hsiao CT Yates JR 3rd Waterman CM Analysis of the myosin-II-responsive focal adhesion proteome reveals a role for beta-Pix in negative regulation of focal adhesion maturation Nat Cell Biol 13 2011 383 393 21423176 PMC3279191
  33. [33]Schiller HB Friedel CC Boulegue C Fassler R Quantitative proteomics of the integrin adhesome show a myosin II-dependent recruitment of LIM domain proteins EMBO Rep 12 2011 259 266 21311561 PMC3059911
  34. [34]Choquet D Felsenfeld DP Sheetz MP Extracellular matrix rigidity causes strengthening of integrin-cytoskeleton linkages Cell 88 1997 39 48 9019403
  35. [35]Wang N Butler JP Ingber DE Mechanotransduction across the cell surface and through the cytoskeleton Science 260 1993 1124 1127 7684161
  36. [36]Matthews BD Overby DR Mannix R Ingber DE Cellular adaptation to mechanical stress: role of integrins, Rho, cytoskeletal tension and mechanosensitive ion channels J Cell Sci 119 2006 508 518 16443749
  37. [37]Guilluy C Swaminathan V Garcia-Mata R O’Brien ET Superfine R Burridge K The Rho GEFs LARG and GEF-H1 regulate the mechanical response to force on integrins Nat Cell Biol 13 2011 722 727 21572419 PMC3107386
  38. [38]Zhao XH Laschinger C Arora P Szaszi K Kapus A McCulloch CA Force activates smooth muscle alpha-actin promoter activity through the Rho signaling pathway J Cell Sci 120 2007 1801 1809 17456553
  39. [39]Watanabe N Madaule P Reid T Ishizaki T Watanabe G Kakizuka A Saito Y Nakao K Jockusch BM Narumiya S p140mDia, a mammalian homolog of Drosophila diaphanous, is a target protein for Rho small GTPase and is a ligand for profilin EMBO J 16 1997 3044 3056 9214622 PMC1169923
  40. [40]Amano M Ito M Kimura K Fukata Y Chihara K Nakano T Matsuura Y Kaibuchi K Phosphorylation and activation of myosin by Rho-associated kinase (Rho-kinase) J Biol Chem 271 1996 20246 20249 8702756
  41. [41]Kimura K Ito M Amano M Chihara K Fukata Y Nakafuku M Yamamori B Feng JH Nakano T Okawa K Regulation of myosin phosphatase by Rho and Rho-Associated kinase (Rho-kinase) Science 273 1996 245 248 8662509
  42. [42]Uyeda TQ Iwadate Y Umeki N Nagasaki A Yumura S Stretching actin filaments within cells enhances their affinity for the myosin II motor domain Plos One 6 2011 e26200 22022566 PMC3192770
  43. [43]Hayakawa K Tatsumi H Sokabe M Actin filaments function as a tension sensor by tension-dependent binding of cofilin to the filament J Cell Biol 195 2011 721 727 22123860 PMC3257564
  44. [44]Zhong C Chrzanowska-Wodnicka M Brown J Shaub A Belkin AM Burridge K Rho-mediated contractility exposes a cryptic site in fibronectin and induces fibronectin matrix assembly J Cell Biol 141 1998 539 551 9548730 PMC2148448
  45. [45]del Rio A Perez-Jimenez R Liu R Roca-Cusachs P Fernandez JM Sheetz MP Stretching single talin rod molecules activates vinculin binding Science 323 2009 638 641 19179532 PMC9339221
  46. [46]Tamada M Sheetz MP Sawada Y Activation of a signaling cascade by cy-toskeleton stretch Dev Cell 7 2004 709 718 15525532
  47. [47]Sawada Y Tamada M Dubin-Thaler BJ Cherniavskaya O Sakai R Tanaka S Sheetz MP Force sensing by mechanical extension of the Src family kinase substrate p130Cas Cell 127 2006 1015 1026 17129785 PMC2746973
  48. [48]Yoshigi M Hoffman LM Jensen CC Yost HJ Beckerle MC Mechanical force mobilizes zyxin from focal adhesions to actin filaments and regulates cytoskeletal reinforcement J Cell Biol 171 2005 209 215 16247023 PMC2171187
  49. [49]Colombelli J Besser A Kress H Reynaud EG Girard P Caussinus E Haselmann U Small JV Schwarz US Stelzer EH Mechanosensing in actin stress fibers revealed by a close correlation between force and protein localization J Cell Sci 122 2009 1665 1679 19401336
  50. [50]Smith MA Blankman E Gardel ML Luettjohann L Waterman CM Beckerle MC A zyxin-mediated mechanism for actin stress fiber maintenance and repair Dev Cell 19 2010 365 376 20833360 PMC2954498
  51. [51]Evans EA Calderwood DA Forces and bond dynamics in cell adhesion Science 316 2007 1148 1153 17525329
  52. [52]Friedland JC Lee MH Boettiger D Mechanically activated integrin switch controls alpha5beta1 function Science 323 2009 642 644 19179533
  53. [53]Kong F Garcia AJ Mould AP Humphries MJ Zhu C Demonstration of catch bonds between an integrin and its ligand J Cell Biol 185 2009 1275 1284 19564406 PMC2712956
  54. [54]Plotnikov SV Pasapera AM Sabass B Waterman CM Force fluctuations within focal adhesions mediate ECM-rigidity sensing to guide directed cell migration Cell 151 2012 1513 1527 23260139 PMC3821979
  55. [55]Beningo KA Dembo M Kaverina I Small JV Wang YL Nascent focal adhesions are responsible for the generation of strong propulsive forces in migrating fibroblasts J Cell Biol 153 2001 881 888 11352946 PMC2192381
  56. [56]Tan JL Tien J Pirone DM Gray DS Bhadriraju K Chen CS Cells lying on a bed of microneedles: an approach to isolate mechanical force Proc Natl Acad Sci USA 100 2003 1484 1489 12552122 PMC149857
  57. [57]Gardel ML Sabass B Ji L Danuser G Schwarz US Waterman CM Traction stress in focal adhesions correlates biphasically with actin retrograde flow speed J Cell Biol 183 2008 999 1005 19075110 PMC2600750
  58. [58]Grashoff C Hoffman BD Brenner MD Zhou R Parsons M Yang MT McLean MA Sligar SG Chen CS Ha T Measuring mechanical tension across vinculin reveals regulation of focal adhesion dynamics Nature 466 2010 263 266 20613844 PMC2901888
  59. [59]Aratyn-Schaus Y Oakes PW Gardel ML Dynamic and structural signatures of lamellar actomyosin force generation Mol Biol Cell 22 2011 1330 1339 21307339 PMC3078065
  60. [60]Stricker J Beckham Y Davidson MW Gardel ML Myosin II-mediated focal adhesion maturation is tension insensitive Plos One 8 2013 e70652 23923013 PMC3726642
  61. [61]Machesky LM Hall A Role of actin polymerization and adhesion to extra-cellular matrix in Rac- and rho-induced cytoskeletal reorganization J Cell Biol 138 1997 913 926 9265656 PMC2138040
  62. [62]Schiller HB Hermann MR Polleux J Vignaud T Zanivan S Friedel CC Sun Z Raducanu A Gottschalk KE Thery M Beta1- and alphav-class integrins cooperate to regulate myosin II during rigidity sensing of fibronectin-based microenvironments Nat Cell Biol 15 2013 625 636 23708002
  63. [63]Mochitate K Pawelek P Grinnell F Stress relaxation of contracted collagen gels: disruption of actin filament bundles, release of cell surface fibronectin, and down-regulation of DNA and protein synthesis Exp Cell Res 193 1991 198 207 1995294
  64. [64]Grinnell F Fibroblasts, myofibroblasts, and wound contraction J Cell Biol 124 1994 401 404 8106541 PMC2119916
  65. [65]Tomasek JJ Haaksma CJ Eddy RJ Vaughan MB Fibroblast contraction occurs on release of tension in attached collagen lattices: dependency on an organized actin cytoskeleton and serum Anat Rec 232 1992 359 368 1543260
  66. [66]Pelham RJ Wang YL Cell locomotion and focal adhesions are regulated by substrate flexibility Proc Natl Acad Sci USA 94 1997 13661 13665 9391082 PMC28362
  67. [67]Wozniak MA Desai R Solski PA Der CJ Keely PJ ROCK-generated contractility regulates breast epithelial cell differentiation in response to the physical properties of a three-dimensional collagen matrix J Cell Biol 163 2003 583 595 14610060 PMC2173660
  68. [68]Paszek MJ Zahir N Johnson KR Lakins JN Rozenberg GI Gefen A Reinhart-King CA Margulies SS Dembo M Boettiger D Tensional homeostasis and the malignant phenotype Cancer Cell 8 2005 241 254 16169468
  69. [70]Maniotis AJ Chen CS Ingber DE Demonstration of mechanical connections between integrins, cytoskeletal filaments, and nucleoplasm that stabilize nuclear structure Proc Natl Acad Sci USA 94 1997 849 854 9023345 PMC19602
  70. [73]McBeath R Pirone DM Nelson CM Bhadriraju K Chen CS Cell shape, cytoskeletal tension, and RhoA regulate stem cell lineage commitment Dev Cell 6 2004 483 495 15068789
  71. [74]Wang N Tytell JD Ingber DE Mechanotransduction at a distance: mechanically coupling the extracellular matrix with the nucleus Nat Rev Mol Cell Biol 10 2009 75 82 19197334
  72. [75]Swift J Ivanovska IL Buxboim A Harada T Dingal PC Pinter J Pajerowski JD Spinler KR Shin JW Tewari M Nuclear lamin-A scales with tissue stiffness and enhances matrix-directed differentiation Science 341 2013 1240104 23990565 PMC3976548
  73. [76]Guilluy C Osborne LD Van Landeghem L Sharek L Superfine R Garcia-Mata R Burridge K Isolated nuclei adapt to force and reveal a mechanotransduction pathway in the nucleus Nat Cell Biol 2014 PMC4085695 24609268
  74. [77]Lombardi ML Jaalouk DE Shanahan CM Burke B Roux KJ Lammerding J The interaction between nesprins and sun proteins at the nuclear envelope is critical for force transmission between the nucleus and cytoskeleton J Biol Chem 286 2011 26743 26753 21652697 PMC3143636
  75. [78]Starr DA Fridolfsson HN Interactions between nuclei and the cytoskeleton are mediated by SUN-KASH nuclear-envelope bridges Annu Rev Cell Dev Biol 26 2010 421 444 20507227 PMC4053175
  76. [79]Crisp M Liu Q Roux K Rattner JB Shanahan C Burke B Stahl PD Hodzic D Coupling of the nucleus and cytoplasm: role of the LINC complex J Cell Biol 172 2006 41 53 16380439 PMC2063530
  77. [80]Gomes ER Jani S Gundersen GG Nuclear movement regulated by Cdc42, MRCK, myosin, and actin flow establishes MTOC polarization in migrating cells Cell 121 2005 451 463 15882626
  78. [81]Luxton GW Gomes ER Folker ES Vintinner E Gundersen GG Linear arrays of nuclear envelope proteins harness retrograde actin flow for nuclear movement Science 329 2010 956 959 20724637 PMC3938394
  79. [82]Chang W Folker ES Worman HJ Gundersen GG Emerin organizes actin flow for nuclear movement and centrosome orientation in migrating fibroblasts Mol Biol Cell 24 2013 3869 3880 24152738 PMC3861083
  80. [83]Kim DH Khatau SB Feng Y Walcott S Sun SX Longmore GD Wirtz D Actin cap associated focal adhesions and their distinct role in cellular mechanosensing Sci Rep 2 2012 555 22870384 PMC3412326
  81. [84]Kim DH Cho S Wirtz D Tight coupling between nucleus and cell migration through the perinuclear actin cap J Cell Sci 2014 PMC4038945 24639463
  82. [86]Goldman RD Pollack R Hopkins NH Preservation of normal behavior by enucleated cells in culture Proc Natl Acad Sci USA 70 1973 750 754 4197626 PMC433350
  83. [87]Verkhovsky AB Svitkina TM Borisy GG Self-polarization and directional motility of cytoplasm Curr Biol 9 1999 11 20 9889119
  84. [88]Webster KD Ng WP Fletcher DA Tensional homeostasis in single fibroblasts Biophys J 107 2014 146 155 24988349 PMC4119263
  85. [89]Trepat X Deng L An SS Navajas D Tschumperlin DJ Gerthoffer WT Butler JP Fredberg JJ Universal physical responses to stretch in the living cell Nature 447 2007 592 595 17538621 PMC2440511
  86. [90]Krishnan R Park CY Lin YC Mead J Jaspers RT Trepat X Lenormand G Tambe D Smolensky AV Knoll AH Reinforcement versus fluidization in cytoskeletal mechanoresponsiveness plos One 4 2009 e5486 19424501 PMC2675060
  87. [91]Dubash AD Wennerberg K Garcia-Mata R Menold MM Arthur WT Burridge K A novel role for Lsc/p115 RhoGEF and LARG in regulating RhoA activity downstream of adhesion to fibronectin J Cell Sci 120 2007 3989 3998 17971419

Cited by

Sorted by citations

Authors
Burridge, K., & Guilluy, C.
Affiliations
  • University of North Carolina at Chapel Hill (US)
  • Centre National de la Recherche Scientifique (FR)
Corresponding author
Burridge, Keith
ORCID
Venue
Experimental Cell Research
Publisher
Elsevier BV
Published
2016-04
Volume
343
Issue
1
Pages
14-20
ISSN
0014-4827 (ISSN-L)
DOI
10.1016/j.yexcr.2015.10.029
PMC
PMC4891215
OpenAlex
W2191883975
Cited by
409
References
4
Citations by year
2026: 31 · 2025: 30 · 2024: 45 · 2023: 39 · 2022: 42 · 2021: 60 · 2020: 53 · 2019: 41 · 2018: 31 · 2017: 26 · 2016: 11
FWCI
2.26
Type
journal article
Language
en
Open access
Licence
Locations
  • Experimental Cell Research (publishedVersion, cc-by-nc-nd)
  • PubMed (publishedVersion)
  • PubMed Central (acceptedVersion, cc-by-nc-nd)
  • Carolina Digital Repository (University of North Carolina at Chapel Hill) (submittedVersion)
  • HAL (submittedVersion, other-oa)
  • UNC Libraries
Funders
  • Kenan Foundation
  • NIH
  • Agence National de la Recherche (ANR-13-JSV1-0008)
  • European Union Seventh Framework Programme (304162)
Topics
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Cell Biology › Cellular Mechanics and Interactions
  • Life Sciences › Biochemistry, Genetics and Molecular Biology › Cell Biology › Skin and Cellular Biology Research
  • Health Sciences › Medicine › Immunology and Allergy › Cell Adhesion Molecules Research
Keywords
focal adhesions, stress fibers, mechanical tension, cell adhesion, substrate stiffness, cell migration
MeSH terms
Animals, Humans, Models, Biological, Stress, Mechanical, Focal Adhesions, Stress Fibers

About these checks

Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.

Statistics and arithmetic are recomputed, not judged.

Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.

What counts as a claim ›How we measure confidence ›How we check this ›

Are you an author, or do you see a problem with this page? Request a review