Mechanobiology of epidermal keratinocytes: Desmosomes, hemidesmosomes, keratin intermediate filaments, and blistering skin diseases

Mechanobiology of Cell-Cell and Cell-Matrix Interactions

Volumn , Issue , 2011, Pages 169-210

  Selby, John C ^a  

^a UNIVERSITY OF ILLINOIS AT URBANA CHAMPAIGN   (United States)

Author keywords

Adherens junctions; Desmosome; Epidermolysis bullosa; Focal adhesion; Hemidesmosome; Keratin intermediate filament; Keratinocyte; Pemphigus

Indexed keywords

EID: 84895042328     PISSN: None     EISSN: None     Source Type: Book    
DOI: 10.1007/978-1-4419-8083-0_9     Document Type: Chapter

References (221)

1. Kligman AM (1964) The biology of the stratum corneum. In: Montagna W, Lobitz Jr WC (eds) The Epidermis. Academic Press, New York
2. Gartner LP, Hiatt JL (2000) Color Atlas of Histology, 3rd edn. Lippincott Williams & Wilkins, Philadelphia
3. Kierszenbaum AL (2002) Histology and Cell Biology: An Introduction to Pathology. Mosby, St. Louis
4. Montagna W, Parakkal PF (1974) The Structure and Function of Skin, 3rd edn. Academic Press, New York
5. Burns T, Breathnach S, Cox N, Griffiths C (2010) Rook's Textbook of Dermatology. Wiley-Blackwell, Oxford
6. Fitzpatrick TB, Freedberg IM, Austen KF, et al (2003) Fitzpatrick's Dermatology in General Medicine, 6th edn. McGraw-Hill, New York
7. Junqueira LC, Carneiro J, Kelley RO (1998) Basic Histology, 9th edn. Appleton & Lange, Stamford
8. Williams BD, Kokko-Cunningham A, Cvetko S, et al (2003) Internet Atlas of Histology. https://histo.life.illinois.edu/histo/atlas/index.php. Accessed June 2003
9. Welch MP, Odland GF, Clark RA (1990) Temporal relationships of F-Actin bundle formation, collagen and fibronectin matrix assembly, and fibronectin receptor expression to wound contraction. J Cell Biol 110:133-145
10. Gabbiani G, Chaponnier C, Huttner I (1978) Cytoplasmic filaments and gap junctions in epithelial cells and myofibroblasts during wound healing. J Cell Biol 76:561-568
11. Hinz B, Pittet P, Smith-Clerc J, et al (2004) Myofibroblast development is characterized by specific cell-cell adherens junctions. Mol Biol Cell 15:4310-4320
12. Gabbiani G, Rungger-Brandle E (1981) The fibroblast. In: Glynn LE (ed) Handbook of Inflammation, Vol 3: Tissue Repair and Regeneration. Elsevier, Amsterdam
13. Wilkes GL, Brown IA, Wildnauer RH (1973) The biomechanical properties of skin. CRC Crit Rev Bioeng 1:453-495
14. Mackenzie IC (1983) The cellular architecture of the stratum corneum. In: Marks R, PlewigG (eds) Stratum Corneum. Springer, Berlin
15. Kenedi RM, Gibson T, Evans JH, Barbenel JC (1975) Tissue mechanics. Phys Med Biol 20:699-717
16. Alberts B, Johnson A, Lewis J, et al (2002) Molecular Biology of the Cell, 4th edn. Garland Science, New York
17. Goldman RD (2001) Worms reveal essential functions for intermediate filaments. Proc Natl Acad Sci U S A 98:7659-7661
18. Mitchison TJ, Cramer LP (1996) Actin-based cell motility and cell locomotion. Cell 84:371-379
19. Bretscher A (1991) Microfilament structure and function in the cortical cytoskeleton. Annu Rev Cell Biol 7:337-374
20. Nogales E (2000) Structural insights into microtubule function. Annu Rev Biochem 69:277-302
21. Gelfand VI, Bershadsky AD (1991) Microtubule dynamics-mechanism, regulation, and function. Annu Rev Cell Biol 7:93-116
22. Lodish H, Baltimore D, Berk A, et al (1995) Molecular Cell Biology, 3rd edn. Scientific American Books, New York
23. Fuchs E, Cleveland DW (1998) A structural scaffolding of intermediate filaments in health and disease. Science 279:514-519
24. Lazarides E (1980) Intermediate filaments as mechanical integrators of cellular space. Nature 283:249-256
25. Coulombe PA, Bousquet O, Ma LL, et al (2000) The "ins" and "outs" of intermediate filament organization. Trends Cell Biol 10:420-428
26. Svitkina TM, Verkhovsky AB, Borisy GG (1996) Plectin sidearms mediate interaction of intermediate filaments with microtubules and other components of the cytoskeleton. J Cell Biol 135:991-1007
27. Goldman RD, Khuon S, Chou YH, et al (1996) The function of intermediate filaments in cell shape and cytoskeletal integrity. J Cell Biol 134:971-983
28. Goldman RD, Clement S, Khuon S, et al (1998) Intermediate filament cytoskeletal system: Dynamic and mechanical properties. Biol Bull 194:361-362
29. Helfand BT, Chang L, Goldman RD (2004) Intermediate filaments are dynamic and motile elements of cellular architecture. J Cell Sci 117:133-141
30. Yoon KH, Yoon M, Moir RD, et al (2001) Insights into the dynamic properties of keratin intermediate filaments in living epithelial cells. J Cell Biol 153:503-516
31. Parry DAD, Steinert PM (1999) Intermediate filaments: Molecular architecture, assembly, dynamics and polymorphism. Q Rev Biophys 32:99-187
32. Fuchs E, Weber K (1994) Intermediate filaments-structure, dynamics, function, and disease. Annu Rev Biochem 63:345-382
33. Hesse M, Magin TM, Weber K (2001) Genes for intermediate filament proteins and the draft sequence of the human genome: Novel keratin genes and a surprisingly high number of pseudogenes related to keratin genes 8 and 18. J Cell Sci 114:2569-2575
34. Szeverenyi I, Cassidy AJ, Chung CW, et al (2008) The human intermediate filament database: Comprehensive information on a gene family involved in many human diseases. Hum Mutat 29:351-360
35. Langbein L, Rogers MA, Winter W, et al (1999) The catalog of human hair keratins I. Expression of the nine type I members in the hair follicle. J Biol Chem 274:19874-19884
36. Coulombe PA, Tong XM, Mazzalupo S, et al (2004) Great promises yet to be fulfilled: Defining keratin intermediate filament function in vivo. Eur J Cell Biol 83:735-746
37. Kirfel J, Peters B, Grund C, et al (2002) Ectopic expression of desmin in the epidermis of transgenic mice permits development of a normal epidermis. Differentiation 70:56-68
38. Kartenbeck J, Schwechheimer K, Moll R, et al (1984) Attachment of vimentin filaments to desmosomal plaques in human meningiomal cells and arachnoidal tissue. J Cell Biol 98:1072-1081
39. Kowalczyk AP, Stappenbeck TS, Parry DAD, et al (1994) Structure and function of desmosomal transmembrane core and plaque molecules. Biophys Chem 50:97-112
40. Yonemura S, Itoh M, Nagafuchi A, et al (1995) Cell-To-cell adherens junction formation and actin filament organization-similarities and differences between non-polarized fibroblasts and polarized epithelial-cells. J Cell Sci 108:127-142
41. Fanning AS (2001) Organization and regulation of the tight junction by the actin-myosin cytoskeleton. In: Cereijido M, Anderson J (eds) Tight Junctions, 2nd edn. CRC Press, Boca Raton
42. Madara JL (1987) Intestinal absorptive cell tight junctions are linked to cytoskeleton. Am J Physiol C 253:171-175
43. Kovbasnjuk ON, Szmulowicz U, Spring KR (1998) Regulation of the MDCK cell tight junction. J Membr Biol 161:93-104
44. Yap AS, Stevenson BR, Abel KC, et al (1995) Microtubule integrity is necessary for the epithelial barrier function of cultured thyroid-cell monolayers. Exp Cell Res 218:540-550
45. Salomon D, Saurat JH, Meda P (1988) Cell-To-cell communication within intact human-skin. J Clin Invest 82:248-254
46. Hall JE, Zampighi GA, Davis RM (1993) Progress in Cell Research, Volume 3: Gap Junctions. Elsevier, Amsterdam
47. Ingber DE (1997) Tensegrity: The architectural basis of cellular mechanotransduction. Annu Rev Physiol 59:575-599
48. Zamir E, Geiger B (2001) Components of cell-matrix adhesions. J Cell Sci 114:3577-3579
49. Shemesh T, Geiger B, Bershadsky AD, et al (2005) Focal adhesions as mechanosensors: A physical mechanism. Proc Natl Acad Sci U S A 102:12383-12388
50. Bershadsky AD, Balaban NQ, Geiger B (2003) Adhesion-Dependent cell mechanosensitivity. Annu Rev Cell Dev Biol 19:677-695
51. Adams CL, Nelson WJ (1998) Cytomechanics of cadherin-mediated cell-cell adhesion. Curr Opin Cell Biol 10:572-577
52. Geiger B, Volk T, Volberg T (1985) Molecular heterogeneity of adherens junctions. J Cell Biol 101:1523-1531
53. Garrod DR, Merritt AJ, Nie ZX (2002) Desmosomal adhesion: Structural basis, molecular mechanism and regulation (Review). Mol Membr Biol 19:81-94
54. Kowalczyk AP, Bornslaeger EA, Norvell SM, et al (1999) Desmosomes: Intercellular adhesive junctions specialized for attachment of intermediate filaments. Int Rev Cytol 185:237-302
55. Smith EA, Fuchs E (1998) Defining the interactions between intermediate filaments and desmosomes. J Cell Biol 141:1229-1241
56. Bornslaeger EA, Corcoran CM, Stappenbeck TS, et al (1996) Breaking the connection: Displacement of the desmosomal plaque protein desmoplakin from cell-cell interfaces disrupts anchorage of intermediate filament bundles and alters intercellular junction assembly. J Cell Biol 134:985-1001
57. Kouklis PD, Hutton E, Fuchs E (1994) Making a connection: Direct binding between keratin intermediate filaments and desmosomal proteins. J Cell Biol 127:1049-1060
58. Jones JCR, Hopkinson SB, Goldfinger LE (1998) Structure and assembly of hemidesmosomes. Bioessays 20:488-494
59. Borradori L, Sonnenberg A (1996) Hemidesmosomes: Roles in adhesion, signaling and human diseases. Curr Opin Cell Biol 8:647-656
60. Dowling J, Yu QC, Fuchs E (1996) Beta-4 integrin is required for epidermal cell adhesion, organization, and survival. Mol Biol Cell 7:3401
61. Karakesisoglou I, Yang YM, Fuchs E (2000) An epidermal plakin that integrates actin and microtubule networks at cellular junctions. J Cell Biol 149:195-208
62. Jefferson JJ, Leung CL, Liem RKH, (2004) Plakins: Goliaths that link cell junctions and the cytoskeleton. Nat Rev Mol Cell Biol 5:542-553
63. Kaverina I, Rottner K, Small JV (1998) Targeting, capture, and stabilization of microtubules at early focal adhesions. J Cell Biol 142:181-190
64. Stehbens SJ, Paterson AD, Crampton MS, et al (2006) Dynamic microtubules regulate the local concentration of E-cadherin at cell-cell contacts. J Cell Sci 119:1801-1811
65. Ingber DE (2003) Tensegrity II. How structural networks influence cellular information processing networks. J Cell Sci 116:1397-1408
66. Lee TYJ, Gotlieb AI (2003) Microfilaments and microtubules maintain endothelial integrity. Microsc Res Tech 60:115-125
67. Rodriguez OC, Schaefer AW, Mandato CA, et al (2003) Conserved microtubule-Actin interactions in cell movement and morphogenesis. Nat Cell Biol 5:599-609
68. Horwitz AR, Parsons JT (1999) Cell biology-cell migration-movin' on. Science 286:1102-1103
69. Ingber DE (2003) Tensegrity I. Cell structure and hierarchical systems biology. J Cell Sci 116:1157-1173
70. Zamansky GB, Nguyen U, Chou IN (1991) An immunofluorescence study of the calcium-induced coordinated reorganization of microfilaments, keratin intermediate filaments, and microtubules in cultured human epidermal keratinocytes. J Invest Dermatol 97:985-994
71. Pasdar M, Li Z, Krzeminski KA (1992) Desmosome assembly in MDCK epithelial-cells does not require the presence of functional microtubules. Cell Motil Cytoskeleton 23:201-212
72. Zamir E, Geiger B (2001) Molecular complexity and dynamics of cell-matrix adhesions. J Cell Sci 114:3583-3590
73. Kaiser HW, Ness W, Offers M, et al (1993) Talin-Adherens junction protein is localized at the epidermal-Dermal interface in skin. J Invest Dermatol 101:789-793
74. Jones PH, Harper S, Watt FM (1995) Stem-cell patterning and fate in human epidermis. Cell 80:83-93
75. Chapman SJ, Eady RAJ (1985) Blistering in keratinocyte cultures: A regular phenomenon associated with differentiation. Eur J Cell Biol 39:352-359
76. Mann PR, Constable H (1977) Induction of basal lamina formation in epidermal cell cultures in vitro. Br J Dermatol 96:421-426
77. Burgeson RE, Christiano AM (1997) The dermal-epidermal junction. Curr Opin Cell Biol 9:651-658
78. Timpl R, Brown JC (1996) Supramolecular assembly of basement membranes. Bioessays 18:123-132
79. Keene DR, Sakai LY, Lunstrum GP, et al (1987) Type-VII collagen forms an extended network of anchoring fibrils. J Cell Biol 104:611-621
80. Watt FM, Mattey DL, Garrod DR (1984) Calcium-induced reorganization of desmosomal components in cultured human keratinocytes. J Cell Biol 99:2211-2215
81. Jones JCR, Goldman RD (1985) Intermediate filaments and the initiation of desmosome assembly. J Cell Biol 101:506-517
82. Vasioukhin V, Bauer C, Yin M, et al (2000) Directed actin polymerization is the driving force for epithelial cell-cell adhesion. Cell 100:209-219
83. Okeefe EJ, Briggaman RA, Herman B (1987) Calcium-induced assembly of adherens junctions in keratinocytes. J Cell Biol 105:807-817
84. Kaiser HW, Ness W, Jungblut I, et al (1993) Adherens junctions- Demonstration in human epidermis. J Invest Dermatol 100:180-185
85. Haftek M, Hansen MU, Kaiser HW, et al (1996) Interkeratinocyte adherens junctions: Immunocytochemical visualization of cell-cell junctional structures, distinct from desmosomes, in human epidermis. J Invest Dermatol 106:498-504
86. Blaschuk OW, Rowlands TM (2002) Plasma membrane components of adherens junctions (Review). Mol Membr Biol 19:75-80
87. Jensen PJ, Wheelock MJ (1996) The relationships among adhesion, stratification and differentiation in keratinocytes. Cell Death Differ 3:357-371
88. Fuchs E, Byrne C (1994) The epidermis: Rising to the surface. Curr Opin Genet Dev 4:725-736
89. Reichert U, Michel S, Schmidt R (1993) The cornified envelope: A key structure of terminally differentiating keratinocytes. In: Darmon M, Blumenberg M (eds) Molecular Biology of the Skin: The Keratinocyte. Academic Press, San Diego
90. Fuchs E (1990) Epidermal differentiation. Curr Opin Cell Biol 2:1028-1035
91. Green KJ, Gaudry CA (2000) Are desmosomes more than tethers for intermediate filaments? Nat Rev Mol Cell Biol 1:208-216
92. Smith FJD, Irvine AD, Terron-Kwiatkowski A, et al (2006) Loss-of-function mutations in the gene encoding filaggrin cause ichthyosis vulgaris. Nat Genet 38:337-342
93. Farquhar MG, Palade GE (1963) Junctional complexes in various epithelia. J Cell Biol 17:375-412
94. Lane EB, McLean WHI (2004) Keratins and skin disorders. J Pathol 204:355-366
95. McGrath JA (2005) Inherited disorders of desmosomes. Australas J Dermatol 46:221-229
96. Sitaru C, Zillikens D (2005) Mechanisms of blister induction by autoantibodies. Exp Dermatol 14:861-875
97. Omary MB, Coulombe PA, McLean WHI (2004) Mechanisms of disease: Intermediate filament proteins and their associated diseases. New Engl J Med 351:2087-2100
98. McGrath JA (2001) Keratinocyte adhesion and the missing link: From Dowling-Meara to Hay-Wells-St John's Hospital Dermatological Society Annual Oration 2000. Clin Exp Dermatol 26:296-304
99. Fassihi H, Wong T, Wessagowit V, et al (2006) Target proteins in inherited and acquired blistering skin disorders. Clin Exp Dermatol 31:252-259
100. Smith FJD, Eady RAJ, Leigh IM, et al (1996) Plectin deficiency results in muscular dystrophy with epidermolysis bullosa. Nat Genet 13:450-457
101. Vidal F, Aberdam D, Miquel C, et al (1995) Integrin beta-4 mutations associated with junctional epidermolysis-bullosa with pyloric atresia. Nat Genet 10:229-234
102. Uitto J, Richard G (2004) Progress in epidermolysis bullosa: Genetic classification and clinical implications. Am J Med Genet C 131:61-74
103. Vassar R, Coulombe PA, Degenstein L, et al (1991) Mutant keratin expression in transgenic mice causes marked abnormalities resembling a human genetic skin-Disease. Cell 64:365-380
104. Bergman R, Harel A, Sprecher E (2007) Dyskeratosis as a histologic feature in epidermolysis bullosa simplex-Dowling Meara. J Am Acad Dermatol 57:463-466
105. Coulombe PA, Hutton ME, Vassar R, et al (1991) A function for keratins and a common thread among different types of epidermolysis bullosa simplex diseases. J Cell Biol 115:1661-1674
106. Zillikens D (1999) Acquired skin disease of hemidesmosomes. J Dermatol Sci 20:134-154
107. Armstrong DK, McKenna KE, Purkis PE, et al (1999) Haploinsufficiency of desmoplakin causes a striate subtype of palmoplantar keratoderma. Hum Mol Genet 8:143-148
108. Reis A, Hennies HC, Langbein L, et al (1994) Keratin-9 gene-mutations in epidermolytic palmoplantar keratoderma (EPPK). Nat Genet 6:174-179
109. Payne AS, Hanakawa Y, Amagai M, et al (2004) Desmosomes and disease: Pemphigus and bullous impetigo. Curr Opin Cell Biol 16:536-543
110. Habif TP (2004) Clinical Dermatology: A Color Guide to Diagnosis and Therapy, 4th edn. Mosby, Edinburgh
111. Evangelista F, Dasher DA, Diaz LA, et al (2008) E-cadherin is an additional immunological target for pemphigus autoantibodies. J Invest Dermatol 128:1710-1718
112. Bystryn J-C, Grando SA (2006) A novel explanation for acantholysis in pemphigus vulgaris: The basal cell shrinkage hypothesis. J Am Acad Dermatol 54:513-516
113. Grando SA, Bystryn J-C, Chernyavsky AI, et al (2009) Apoptolysis: A novel mechanism of skin blistering in pemphigus vulgaris linking the apoptotic pathways to basal cell shrinkage and suprabasal acantholysis. Exp Dermatol 18:764-770
114. Ladhani S (2001) Recent developments in staphylococcal scalded skin syndrome. Clin Microbiol Infect 7:301-307
115. Stanley JR, Amagai M (2006) Pemphigus, bullous impetigo, and the staphylococcal scaldedskin syndrome. New Engl J Med 355:1800-1810
116. Eliades A (2010) Dermatology Image Atlas. http://dermatlas.org. Accessed June 2010
117. Cohen B (2010) Dermatology Image Atlas. http://dermatlas. org. Accessed June 2010
118. Hosler G (2010) Dermatology Image Atlas. http://dermatlas. org. Accessed June 2010
119. Galou M, Gao J, Humbert J, et al (1997) The importance of intermediate filaments in the adaptation of tissues to mechanical stress: Evidence from gene knockout studies. Biol Cell 89:85-97
120. Andra K, Lassmann H, Bittner R, et al (1997) Targeted inactivation of plectin reveals essential function in maintaining the integrity of skin, muscle, and heart cytoarchitecture. Gene Dev 11:3143-3156
121. Andra K, Kornacker I, Jorgl A, et al (2003) (2003) Plectin-isoform- specific rescue of hemidesmosomal defects in plectin (-/-) keratinocytes. J Invest Dermatol 120:189-197
122. Vasioukhin V, Bowers E, Bauer C, et al (2001) Desmoplakin is essential in epidermal sheet formation. Nat Cell Biol 3:1076-1085
123. Amagai M, Tsunoda K, Suzuki H, et al (2000) Use of autoantigen knockout mice in developing an active autoimmune disease model for pemphigus. J Clin Invest 105:625-631
124. Hashimoto T (2003) Recent advances in the study of the pathophysiology of pemphigus. Arch Dermatol Res 295:S2-S11
125. Adelmann HB (1966) Marcello Malpighi and the Evolution of Embryology. Cornell University Press, New York
126. Vogel HG (2002) Mechanical properties of human skin: Animal models. In: Elsner P, Berardesca E, Wilhelm K-P, et al (eds) Bioengineering of the Skin: Skin Biomechanics. CRC Press, Boca Raton
127. Lanir Y, Fung YCB (1974) Two-Dimensional mechanical properties of rabbit skin-II. Experimental results. J Biomech 7:171-182
128. Manschot JFM, Brakkee AJM (1986) The measurement and modelling of mechanical properties of human skin in vivo-I. The measurement. J Biomech 19:511-515
129. Manschot JFM, Brakkee AJM (1986) The measurement and modelling of mechanical properties of human skin in vivo-II. The model. J Biomech 19:517-521
130. Lanir Y, Fung YCB (1974) Two dimensional mechanical properties of rabbit skin-I. Experimental system. J Biomech 7:29-34
131. Mansour JM, Davis BR, Srour M, et al (1993) A method for obtaining repeatable measurements of the tensile properties of skin at low strain. J Biomech 26:211-216
132. Daly CH (1982) Biomechanical properties of dermis. J Invest Dermatol 79:17s-20s.
133. Koutroupi KS, Barbenel JC (1990) Mechanical and failure behavior of the stratum corneum. J Biomech 23:281-287
134. Wu KS, van Osdol WW, Dauskardt RH (2006) Mechanical properties of human stratum corneum: Effects of temperature, hydration, and chemical treatment. Biomaterials 27:785-795
135. Wildnauer RH, Bothwell JW, Douglass AB (1971) Stratum corneum biomechanical properties. I. Influence of relative humidity on normal and extracted human stratum corneum. J Invest Dermatol 56:72-78
136. Lafrance H, Guillot M, Germain L, et al (1995) A method for the evaluation of tensile properties of skin equivalents. Med Eng Phys 17:537-543
137. Lafrance H, Yahia LH, Germain L, et al (1995) Study of the tensile properties of living skin equivalents. Biomed Mater Eng 5:195-208
138. Chistolini P, De Angelis G, De Luca M, et al (1999) Analysis of the mechanical properties of in vitro reconstructed epidermis: Preliminary results. Med Biol Eng Comput 37:670-672
139. Hendriks FM, Brokken D, Oomens CWJ, et al (2004) Influence of hydration and experimental length scale on the mechanical response of human skin in vivo using optical coherence tomography. Skin Res Technol 10:231-241
140. Hendriks FM, Brokken D, Oomens CWJ, et al (2006) The relative contributions of different skin layers to the mechanical behavior of human skin in vivo using suction experiments. Med Eng Phys 28:259-266
141. Dobrev HP (2002) Mechanical properties in other dermatological diseases. In: Elsner P, Berardesca E, Wilhelm K-P, et al (eds) Bioengineering of the Skin: Skin Biomechanics. CRC Press, Boca Raton
142. Bronneberg D, Bouten CVC, Oomens CWJ, et al (2006) An in vitro model system to study the damaging effects of prolonged mechanical loading of the epidermis. Ann Biomed Eng 34:506-514
143. Wolfram MA, Wolejsza NF, Laden K (1972) Biomechanical properties of delipidized stratum corneum. J Invest Dermatol 59:421-426
144. Wilhelmi BJ, Blackwell SJ, Phillips LG (1999) Langer's lines: To use or not to use. Plast Reconstr Surg 104:208-214
145. Tomson AM, Scholma J, Blaauw EH (1995) The effect of detachment of cultured epithelial sheets on cellular ultrastructure and organization. Epithelial Cell Biol 4:43-51
146. Humphrey JD (2003) Continuum biomechanics of soft biological tissues. Proc Roy Soc Lond Ser A 459:3-46
147. Vogel V, Sheetz M (2006) Local force and geometry sensing regulate cell functions. Nat Rev Mol Cell Biol 7:265-275
148. Ingber DE (2003) Mechanobiology and diseases of mechanotransduction. Ann Med 35:564-577
149. Zhu C, Bao G, Wang N (2000) Cell mechanics: Mechanical response, cell adhesion, and molecular deformation. Annu Rev Biomed Eng 2:189-226
150. Stamenovic D, Ingber DE (2002) Models of cytoskeletal mechanics of adherent cells. Biomech Model Mechanobiol 1:95-108
151. Humphrey JD (2002) On mechanical modeling of dynamic changes in the structure and properties of adherent cells. Math Mech Solids 7:521-539
152. Hochmuth RM (2000) Micropipette aspiration of living cells. J Biomech 33:15-22
153. Wang N, Butler JP, Ingber DE (1993) Mechanotransduction across the cell surface and through the cytoskeleton. Science 260:1124-1127
154. Wang N, Ingber DE (1995) Probing transmembrane mechanical coupling and cytomechanics using magnetic twisting cytometry. Biochem Cell Biol 73:327-335
155. Bausch AR, Moller W, Sackmann E (1999) Measurement of local viscoelasticity and forces in living cells by magnetic tweezers. Biophys J 76:573-579
156. Bausch AR, Ziemann F, Boulbitch AA, et al (1998) Local measurements of viscoelastic parameters of adherent cell surfaces by magnetic bead microrheometry. Biophys J 75:2038-2049
157. Matthews BD, Overby DR, Alenghat FJ, et al (2004) Mechanical properties of individual focal adhesions probed with a magnetic microneedle. Biochem Biophys Res Commun 313:758-764
158. Saif MTA, Sager CR, Coyer S (2003) Functionalized biomicroelectromechanical systems sensors for force response study at local adhesion sites of single living cells on substrates. Ann Biomed Eng 31:950-961
159. Yang S, Saif T (2005) Reversible and repeatable linear local cell force response under large stretches. Exp Cell Res 305:42-50
160. Tan JL, Tien J, Pirone DM, et al (2003) Cells lying on a bed of microneedles: An approach to isolate mechanical force. Proc Natl Acad Sci U S A 100:1484-1489
161. Balaban NQ, Schwarz US, Riveline D, et al (2001) Force and focal adhesion assembly: A close relationship studied using elastic micropatterned substrates. Nat Cell Biol 3:466-472
162. Thoumine O, Ott A, Cardoso O, et al (1999) Microplates: A new tool for manipulation and mechanical perturbation of individual cells. J Biochem Biophys Meth 39:47-62
163. Suresh S, Spatz J, Mills JP, et al (2005) Connections between single-cell biomechanics and human disease states: Gastrointestinal cancer and malaria. Acta Biomater 1:15-30
164. Guck J, Ananthakrishnan R, Mahmood H, et al (2001) The optical stretcher: A novel laser tool to micromanipulate cells. Biophys J 81:767-784
165. Radmacher M (1997) Measuring the elastic properties of biological samples with the AFM. IEEE Eng Med Biol 16:47-57
166. Bowen WR, Hilal N, Lovitt RW, et al (1998) Direct measurement of the force of adhesion of a single biological cell using an atomic force microscope. Colloid Surf A 136:231-234
167. Hoh JH, Schoenenberger CA (1994) Surface morphology and mechanical properties of MDCK monolayers by atomic force microscopy. J Cell Sci 107:1105-1114
168. Waschke J, Bruggeman P, Baumgartner W, et al (2005) Pemphigus foliaceus IgG causes dissociation of desmoglein 1-containing junctions without blocking desmoglein 1 transinteraction. J Clin Invest 115:3157-3165
169. Waschke J, Menendez-Castro C, Bruggeman P, et al (2007) Imaging and force spectroscopy on desmoglein 1 using atomic force microscopy reveal multivalent Ca2+-Dependent, lowaffinity trans-interaction. J Memb Sci 216:83-92
170. Heupel W-M, Zillikens D, Drenckhahn D, et al (2008) Pemphigus vulgaris IgG directly inhibit desmoglein 3-mediated transinteraction. J Immunol 181:1825-1834
171. Heupel W-M, Engerer P, Schmidt E, et al (2009) Pemphigus vulgaris IgG cause loss of desmoglein-mediated adhesion and keratinocyte dissociation independent of epidermal growth factor receptor. Am J Pathol 174:475-485
172. Spindler V, Heupel W-M, Efthymiadis A, et al (2009) Desmocollin 3-mediated binding is crucial for keratinocyte cohesion and is impaired in pemphigus. J Biol Chem 284:30556-30564
173. Fabry B, Maksym GN, Butler JP, et al (2001) Scaling the microrheology of living cells. Phys Rev Lett 87:148102
174. Wang N, Tolic-Norrelykke IM, Chen JX, et al (2002) Cell prestress. I. Stiffness and prestress are closely associated in adherent contractile cells. Am J Physiol Cell Physiol 282: C606-C616
175. Hennings H, Holbrook KA (1983) Calcium regulation of cell-cell contact and differentiation of epidermal cells in culture-An ultrastructural-study. Exp Cell Res 143:127-142
176. Green KJ, Geiger B, Jones JCR, et al (1987) The relationship between intermediate filaments and microfilaments before and during the formation of desmosomes and adherens-Type junctions in mouse epidermal keratinocytes. J Cell Biol 104:1389-1402
177. Brown TD (2000) Techniques for mechanical stimulation of cells in vitro: A review. J Biomech 33:3-14
178. Selby JC, Shannon MA (2008) A method to fabricate mesoscopic freestanding polydimethylsiloxane membranes used to probe the rheology of an epithelial sheet. J Biochem Biophys Meth 70:932-944
179. Selby JC, Shannon MA (2007) Apparatus for measuring the finite load-Deformation behavior of a sheet of epithelial cells cultured on a mesoscopic freestanding elastomer membrane. Rev Sci Instrum 78:094301
180. Selby JC, Shannon MA (2007) Mechanical response of a living human epidermal keratinocyte sheet as measured in a composite diaphragm inflation experiment. Biorheology 44:319-348
181. Yano S, Komine M, Fujimoto M, et al (2004) Mechanical stretching in vitro regulates signal transduction pathways and cellular proliferation in human epidermal keratinocytes. J Invest Dermatol 122:783-790
182. Kippenberger S, Bern A, Loitsch S, et al (2000) Signaling of mechanical stretch in human keratinocytes via MAP kinases. J Invest Dermatol 114:408-412
183. Russell D, Andrews PA, James J, et al (2004) Mechanical stress induces profound remodelling of keratin filaments and cell junctions in epidermolysis bullosa simplex keratinocytes. J Cell Sci 117:5233-5243
184. Fudge D, Russell D, Beriault D, et al (2008) The intermediate filament network in cultured human keratinocytes is remarkably extensible and resilient. PLoS ONE doi:10.1371/journal. pone.0002327
185. Janmey PA, Euteneuer U, Traub P, et al (1991) Viscoelastic properties of vimentin compared with other filamentous biopolymer networks. J Cell Biol 113:155-160
186. Janmey PA (1991) A torsion pendulum for measurement of the viscoelasticity of biopolymers and its application to actin networks. J Biochem Biophys Meth 22:41-53
187. Xu JY, Tseng Y, Wirtz D (2000) Strain hardening of actin filament networks-regulation by the dynamic cross-linking protein alpha-Actinin. J Biol Chem 275:35886-35892
188. Xu JY, Palmer A, Wirtz D (1998) Rheology and microrheology of semiflexible polymer solutions: Actin filament networks. Macromolecules 31:6486-6492
189. Tseng Y, Schafer BW, Almo SC, et al (2002) Functional synergy of actin filament crosslinking proteins. J Biol Chem 277:25609-25616
190. Ma LL, Xu JY, Coulombe PA, et al (1999) Keratin filament suspensions show unique micromechanical properties. J Biol Chem 274:19145-19151
191. Yamada S, Wirtz D, Coulombe PA (2003) The mechanical properties of simple epithelial keratins 8 and 18: Discriminating between interfacial and bulk elasticities. J Struct Biol 143:45-55
192. Yamada S, Wirtz D, Coulombe PA (2002) Pairwise assembly determines the intrinsic potential for self-organization and mechanical properties of keratin filaments. Mol Biol Cell 13:382-391
193. Ma LL, Yamada S, Wirtz D, et al (2001) A "hot-spot" mutation alters the mechanical properties of keratin filament networks. Nat Cell Biol 3:503-506
194. Bousquet O, Ma LL, Yamada S, et al (2001) The nonhelical tail domain of keratin 14 promotes filament bundling and enhances the mechanical properties of keratin intermediate filaments in vitro. J Cell Biol 155:747-753
195. Liu XM, Pollack GH (2002) Mechanics of F-Actin characterized with microfabricated cantilevers. Biophys J 83:2705-2715
196. Kis A, Kasas S, Babic B, et al (2002) Nanomechanics of microtubules. Phys Rev Lett 89:248101
197. Mucke N, Kreplak L, Kirmse R, et al (2004) Assessing the flexibility of intermediate filaments by atomic force microscopy. J Mol Biol 335:1241-1250
198. Guzman C, Jeney S, Kreplak L, et al (2006) Exploring the mechanical properties of single vimentin intermediate filaments by atomic force microscopy. J Mol Biol 360:623-630
199. Kiss B, Karsai A, Kellermayer MSZ (2006) Nanomechanical properties of desmin intermediate filaments. J Struct Biol 155:327-339
200. Kreplak L, Bar H, Leterrier JF, et al (2005) Exploring the mechanical behavior of single intermediate filaments. J Mol Biol 354:569-577
201. Fudge DS, Gardner KH, Forsyth VT, et al (2003) The mechanical properties of hydrated intermediate filaments: Insights from hagfish slime threads. Biophys J 85:2015-2027
202. Kreplak L, Fudge D (2006) Biomechanical properties of intermediate filaments: From tissues to single filaments and back. Bioessays 29:26-35
203. Qin Z, Buehler MJ, Kreplak L (2010) A multi-scale approach to understand the mechanobiology of intermediate filaments. J Biomech 43:15-22
204. Herrmann H, Bar H, Kreplak L, et al (2007) Intermediate filaments: From cell architecture to nanomechanics. Nat Rev Mol Cell Biol 8:562-573
205. Leckband D, Israelachvili J (2001) Intermolecular forces in biology. Q Rev Biophys 34:105-267
206. Florin EL, Moy VT, Gaub HE (1994) Adhesion forces between individual ligand-receptor pairs. Science 264:415-417
207. McGuiggan PM, Israelachvili JN (1990) Adhesion and short-range forces between surfaces 2. Effects of surface lattice mismatch. J Mater Res 5:2232-2243
208. Israelachvili JN, McGuiggan PM (1990) Adhesion and short-range forces between surfaces 1. New apparatus for surface force measurements. J Mater Res 5:2223-2231
209. Helm CA, Knoll W, Israelachvili JN (1991) Measurement of ligand receptor interactions. Proc Natl Acad Sci U S A 88:8169-8173
210. Svoboda K, Block SM (1994) Biological applications of optical forces. Annu Rev Biophys Biomol Struct 23:247-285
211. Evans E, Ritchie K, Merkel R (1995) Sensitive force technique to probe molecular adhesion and structural linkages at biological interfaces. Biophys J 68:2580-2587
212. Hanley W, McCarty O, Jadhav S, et al (2003) Single molecule characterization of P-selectin/ ligand binding. J Biol Chem 278:10556-10561
213. Leckband D, Prakasam A (2006) Mechanism and dynamics of cadherin adhesion. Annu Rev Biomed Eng 8:259-287
214. Sivasankar S, Gumbiner B, Leckband D (2001) Direct measurements of multiple adhesive alignments and unbinding trajectories between cadherin extracellular domains. Biophys J 80:1758-1768
215. Leckband D, Sivasankar S (2000) Mechanism of hemophilic cadherin adhesion. Curr Opin Cell Biol 12:587-592
216. Sivasankar S, Brieher W, Lavrik N, et al (1999) Direct molecular force measurements of multiple adhesive interactions between cadherin ectodomains. Proc Natl Acad Sci U S A 96:11820-11824
217. Panorchan P, Thompson MS, Davis KJ, et al (2006) Single-molecule analysis of cadherinmediated cell-cell adhesion. J Cell Sci 119:66-74
218. Leckband D (2004) Molecular mechanisms of cell adhesion: New perspectives from surface force measurements. J Adhes 80:409-432
219. Bayas MV, Leung A, Evans E, et al (2006) Lifetime measurements reveal kinetic differences between homophilic cadherin bonds. Biophys J 90:1385-1395
220. Fung CKM, Seiffert-Sinha K, Lai KWC, et al (2010) Investigation of human keratinocyte cell adhesion using atomic force microscopy. Nanomedicine: NMB 6:191-200
221. Elias PM, Ahn SK, Brown BE, et al (2002) Origin of the epidermal calcium gradient: Regulation by barrier status and role of active vs. passive mechanisms. J Invest Dermatol 119:1269-1274.