title: Torque measurement at the single-molecule level.
abstract: Methods for exerting and measuring forces on single molecules have revolutionized the study of the physics of biology. However, it is often the case that biological processes involve rotation or torque generation, and these parameters have been more difficult to access experimentally. Recent advances in the single-molecule field have led to the development of techniques that add the capability of torque measurement. By combining force, displacement, torque, and rotational data, a more comprehensive description of the mechanics of a biomolecule can be achieved. In this review, we highlight a number of biological processes for which torque plays a key mechanical role. We describe the various techniques that have been developed to directly probe the torque experienced by a single molecule, and detail a variety of measurements made to date using these new technologies. We conclude by discussing a number of open questions and propose systems of study that would be well suited for analysis with torsional measurement techniques.
authors.affiliation: Laboratory of Chemistry and Cell Biology, The Rockefeller University, New York, New York 10065, USA. sforth@rockefeller.edu
authors.forename: Scott
authors.initials: S
authors.lastname: Forth
authors.forename: Maxim Y
authors.initials: MY
authors.lastname: Sheinin
authors.forename: James
authors.initials: J
authors.lastname: Inman
authors.forename: Michelle D
authors.initials: MD
authors.lastname: Wang
mesh_terms: D000818:Animals; D002468:Cell Physiological Phenomena; D004247:DNA / Q000737:chemistry*; D005057:Eukaryotic Cells / Q000378:metabolism; D006801:Humans; D049329:Nanostructures / Q000737:chemistry; D011387:Prokaryotic Cells / Q000378:metabolism; D011506:Proteins / Q000737:chemistry*; D012399:Rotation; D019415:Torque
chemical_list: D011506:Proteins; D004247:DNA
references.citation: Curr Opin Microbiol. 2003 Apr;6(2):93-100
references.pmid: 12732296
references.citation: Nature. 2008 Apr 3;452(7187):598-603
references.pmid: 18327250
references.citation: Proc Natl Acad Sci U S A. 2009 Dec 29;106(52):22564-8
references.pmid: 20018699
references.citation: Mol Cell. 2012 Mar 30;45(6):710-8
references.pmid: 22464441
references.citation: Proc Natl Acad Sci U S A. 1997 Dec 23;94(26):14433-7
references.pmid: 9405630
references.citation: Nano Lett. 2011 Dec 14;11(12):5558-63
references.pmid: 22047401
references.citation: Mol Cell Biol. 2011 Feb;31(3):482-94
references.pmid: 21098118
references.citation: Proc Natl Acad Sci U S A. 2002 Feb 19;99(4):1960-5
references.pmid: 11854495
references.citation: Nat Rev Mol Cell Biol. 2006 Aug;7(8):580-8
references.pmid: 16936698
references.citation: Nature. 2000 Mar 2;404(6773):103-6
references.pmid: 10716452
references.citation: Biophys J. 2010 Apr 7;98(7):1267-76
references.pmid: 20371326
references.citation: Biochim Biophys Acta. 2012 Jul;1819(7):632-8
references.pmid: 22233557
references.citation: Phys Rev Lett. 2006 Aug 4;97(5):058301
references.pmid: 17026144
references.citation: Nature. 2000 Apr 20;404(6780):901-4
references.pmid: 10786800
references.citation: Phys Chem Chem Phys. 2009 Jun 28;11(24):4800-3
references.pmid: 19506753
references.citation: Nat Struct Mol Biol. 2008 Feb;15(2):146-54
references.pmid: 18193062
references.citation: Opt Express. 2003 Mar 10;11(5):446-51
references.pmid: 19461751
references.citation: Biochemistry. 2003 Sep 16;42(36):10718-25
references.pmid: 12962496
references.citation: Proc Natl Acad Sci U S A. 2012 Apr 17;109(16):6106-11
references.pmid: 22474350
references.citation: Phys Rev Lett. 2006 May 5;96(17):178102
references.pmid: 16712339
references.citation: Crit Rev Biochem Mol Biol. 1991;26(2):151-226
references.pmid: 1914495
references.citation: Q Rev Biophys. 2008 May;41(2):103-32
references.pmid: 18812014
references.citation: Biophys J. 1997 Mar;72(3):1335-46
references.pmid: 9138579
references.citation: Proc Natl Acad Sci U S A. 2011 Oct 11;108(41):16992-7
references.pmid: 21969536
references.citation: Science. 1996 Feb 9;271(5250):795-9
references.pmid: 8628994
references.citation: Mol Microbiol. 1992 Feb;6(4):425-33
references.pmid: 1313943
references.citation: Phys Rev E Stat Nonlin Soft Matter Phys. 2001 May;63(5 Pt 1):051903
references.pmid: 11414929
references.citation: Front Biosci. 2007 May 01;12:4409-23
references.pmid: 17485385
references.citation: Trends Cell Biol. 2003 Mar;13(3):114-21
references.pmid: 12628343
references.citation: Nature. 2006 Aug 17;442(7104):836-9
references.pmid: 16862122
references.citation: Nat Methods. 2007 Mar;4(3):223-5
references.pmid: 17322891
references.citation: Genetica. 1999;106(1-2):57-62
references.pmid: 10710710
references.citation: Biophys J. 1993 Nov;65(5):2201-16
references.pmid: 8298044
references.citation: Nucleic Acids Res. 2010 Oct;38(19):e182
references.pmid: 20685815
references.citation: Cell. 1989 Jan 13;56(1):111-8
references.pmid: 2535966
references.citation: Rev Sci Instrum. 2011 Mar;82(3):034302
references.pmid: 21456769
references.citation: Nature. 2006 Jan 5;439(7072):100-104
references.pmid: 16397501
references.citation: Phys Rev Lett. 2009 Jan 16;102(2):028302
references.pmid: 19257322
references.citation: Proc Natl Acad Sci U S A. 1975 Nov;72(11):4275-9
references.pmid: 172901
references.citation: J Mol Biol. 2003 Apr 11;327(5):1043-51
references.pmid: 12662929
references.citation: Proc Natl Acad Sci U S A. 2007 Jan 30;104(5):1500-5
references.pmid: 17242356
references.citation: Nat Methods. 2010 Dec;7(12):977-80
references.pmid: 20953173
references.citation: Trends Biochem Sci. 2002 Mar;27(3):154-60
references.pmid: 11893513
references.citation: Science. 1998 Oct 30;282(5390):902-7
references.pmid: 9794753
references.citation: Proc Natl Acad Sci U S A. 2009 Oct 6;106(40):17007-12
references.pmid: 19805111
references.citation: Nat Struct Mol Biol. 2007 Oct;14(10):968-73
references.pmid: 17891151
references.citation: Cell. 1985 Oct;42(3):799-808
references.pmid: 2996776
references.citation: Nat Rev Mol Cell Biol. 2010 Mar;11(3):208-19
references.pmid: 20177396
references.citation: Phys Rev E Stat Nonlin Soft Matter Phys. 2009 Oct;80(4 Pt 1):040901
references.pmid: 19905264
references.citation: Nature. 1973 Oct 19;245(5425):380-2
references.pmid: 4593496
references.citation: Proc Natl Acad Sci U S A. 2006 May 23;103(21):8066-71
references.pmid: 16698936
references.citation: Opt Lett. 2010 Sep 1;35(17):2949-51
references.pmid: 20808379
references.citation: Science. 2001 May 4;292(5518):912-4
references.pmid: 11340200
references.citation: Phys Rev Lett. 2011 Sep 2;107(10):108102
references.pmid: 21981534
references.citation: Phys Rev E Stat Nonlin Soft Matter Phys. 2007 Aug;76(2 Pt 1):021926
references.pmid: 17930084
references.citation: Proc Natl Acad Sci U S A. 1992 Sep 15;89(18):8784-8
references.pmid: 1326763
references.citation: Opt Express. 2006 Jul 24;14(15):6963-70
references.pmid: 19516880
references.citation: Proc Natl Acad Sci U S A. 1998 Nov 24;95(24):14152-7
references.pmid: 9826669
references.citation: Mol Cell. 2012 Apr 27;46(2):115-24
references.pmid: 22541554
references.citation: Science. 1996 Mar 29;271(5257):1835-7
references.pmid: 8596951
references.citation: Proc Natl Acad Sci U S A. 1976 Nov;73(11):3872-6
references.pmid: 186775
references.citation: EMBO J. 1989 Dec 20;8(13):4335-44
references.pmid: 2556269
references.citation: Science. 2001 Apr 27;292(5517):733-7
references.pmid: 11326101
references.citation: Biophys J. 2000 Feb;78(2):1036-41
references.pmid: 10653817
references.citation: J Phys Chem B. 2010 Dec 23;114(50):16929-35
references.pmid: 21090816
references.citation: Methods Enzymol. 2010;475:377-404
references.pmid: 20627165
references.citation: EMBO J. 2006 Jun 7;25(11):2575-83
references.pmid: 16710299
references.citation: Biophys J. 2011 Jul 20;101(2):L5-7
references.pmid: 21767475
references.citation: Nature. 1994 Oct 27;371(6500):752
references.pmid: 7935835
references.citation: Proc Natl Acad Sci U S A. 2004 Apr 6;101(14):4776-80
references.pmid: 15037753
references.citation: Nat Rev Cancer. 2009 Dec;9(12):849-61
references.pmid: 19907434
references.citation: Cell. 2010 Aug 20;142(4):519-30
references.pmid: 20723754
references.citation: Proc Natl Acad Sci U S A. 2011 Nov 1;108(44):17951-6
references.pmid: 21997211
references.citation: Annu Rev Biochem. 2003;72:19-54
references.pmid: 12500982
references.citation: Cell. 2007 Jun 29;129(7):1299-309
references.pmid: 17604719
references.citation: J Biol Chem. 1990 Oct 5;265(28):16821-9
references.pmid: 1698768
references.citation: EMBO J. 1991 Feb;10(2):387-95
references.pmid: 1991452
references.citation: Nature. 2001 Jan 4;409(6816):113-5
references.pmid: 11343125
references.citation: Nucleic Acids Res. 1997 Apr 1;25(7):1412-8
references.pmid: 9060437
references.citation: Cell. 1998 Jun 26;93(7):1117-24
references.pmid: 9657145
references.citation: Nature. 2003 Jul 17;424(6946):338-41
references.pmid: 12867987
references.citation: Biochemistry. 2005 Apr 12;44(14):5357-72
references.pmid: 15807529
references.citation: Nature. 2001 Oct 18;413(6857):748-52
references.pmid: 11607035
references.citation: Biochem Biophys Res Commun. 2008 Feb 22;366(4):951-7
references.pmid: 18083117
references.citation: Chromosoma. 2011 Aug;120(4):323-34
references.pmid: 21567156
references.citation: J Mol Biol. 1999 Oct 8;292(5):1149-60
references.pmid: 10512709
references.citation: Nature. 1997 Mar 20;386(6622):299-302
references.pmid: 9069291
references.citation: Nature. 2009 May 21;459(7245):364-70
references.pmid: 19458712
references.citation: Phys Rev Lett. 2004 May 14;92(19):190801
references.pmid: 15169392
references.citation: Annu Rev Genet. 2002;36:175-203
references.pmid: 12429691
references.citation: Nature. 1997 Sep 18;389(6648):251-60
references.pmid: 9305837
references.citation: Rev Sci Instrum. 2011 Oct;82(10):103707
references.pmid: 22047303
references.citation: Nature. 1953 Apr 25;171(4356):737-8
references.pmid: 13054692
references.citation: Proc Natl Acad Sci U S A. 1987 Oct;84(20):7024-7
references.pmid: 2823250
references.citation: Phys Rev Lett. 2008 Apr 11;100(14):148301
references.pmid: 18518075
references.citation: J Biol Chem. 2001 Jan 26;276(4):2790-6
references.pmid: 11056156
references.citation: Nature. 1987 Dec 24-31;330(6150):769-71
references.pmid: 3320757
references.citation: Curr Opin Struct Biol. 2012 Jun;22(3):304-12
references.pmid: 22658779
references.citation: Proc Natl Acad Sci U S A. 1992 Jul 1;89(13):6055-9
references.pmid: 1631091
references.citation: Annu Rev Biophys Biomol Struct. 1994;23:609-43
references.pmid: 7919794
references.citation: IEEE Eng Med Biol Mag. 2003 Nov-Dec;22(6):33-42
references.pmid: 15007989
references.citation: J Biol Chem. 1990 Oct 5;265(28):16830-40
references.pmid: 2170357
references.citation: Opt Lett. 2009 Jan 15;34(2):139-41
references.pmid: 19148234
references.citation: Nat Commun. 2011 Aug 23;2:439
references.pmid: 21863006
references.citation: Nano Lett. 2012 Jul 11;12(7):3634-9
references.pmid: 22642488
references.citation: Phys Rev Lett. 2009 Feb 20;102(7):078301
references.pmid: 19257716
references.citation: Nat Struct Mol Biol. 2006 May;13(5):444-50
references.pmid: 16622406
references.citation: Proc Natl Acad Sci U S A. 1997 Dec 23;94(26):14418-22
references.pmid: 9405627
references.citation: Opt Lett. 2002 May 1;27(9):743-5
references.pmid: 18007918
references.citation: ACS Nano. 2011 Feb 22;5(2):1418-27
references.pmid: 21280614
references.citation: Proc Natl Acad Sci U S A. 2001 Jul 17;98(15):8219-26
references.pmid: 11459956
references.citation: Nano Lett. 2009 Apr;9(4):1720-5
references.pmid: 19301859