- Molecular, Cellular and Tissue Engineering
- Sc.D. Physiology Harvard 1990
- M.S. Biomedical Huazhong University of Science & Technology 1984
- B.S. Biomechanics Huazhong University of Science & Technology 1982
- Professor, Department of Mechanical Science and Engineering, UIUC, March 1, 2006-date
- Senior Lecturer, Department of Physiology and Cell Biology, Harvard School of Public Health, July 2005-February 2006.
- Associate Professor, Department of Physiology and Cell Biology, Harvard School of Public Health, 2000-2005
- Adjunct Vice Dean, School of Life Sciences, Huazhong University of Science and Technology, 1998-2006
- Assistant Professor, Department of Physiology and Cell Biology, Harvard School of Public Health, 1994-2000
- Visiting Professor, Huazhong University of Science and Technology, 1994-date
- Visiting Professor, University of Paris XII, 1994
- Research Associate, Department of Physiology, Harvard School of Public Health, 1990-1994
- Research Fellow, Department of Physiology, Harvard School of Public Health, 1986-1990
- Visiting Fellow, Department of Physiology, Harvard School of Public Health, 1986-1987
- Visiting Fellow, Department of Biophysiology and Biochemistry, National Institutes of Health, 1984-1986
- Research Assistant, Department of Bioengineering, Huazhong University of Science & Technology, 1983-1984
For more information
- Molecular, Cellular and Tissue Engineering
Professor Wang uses advanced research techniques to gain a fundamental understanding of cell mechanics, including cytoskeletal biomechanics and control of cell form and function; bio-imaging of cytoskeletal structures and stress distribution in living cells; mechanotransduction, nuclear deformation and gene expression; and mechanical biotechnologies and their applications to cells, tissues, and organisms.
He has developed a technology called intracellular stress tomography, which he uses to address fundamental questions about stress propogation and distribution in living cells. He has also developed three-dimensional magnetic twisting cytometry technology and used it to quantify mechanical anisotropy in living cells.
Professor Wang was the first researcher to provide direct evidence that transmembrane adhesion molecule integrins mediate the transmission of force across the cell surface to the cytoskeleton. This fundamental finding opened the field of cell mechanics to the study of combinating biochemistry and biomechanics at cellular and subcellular levels. Using a technique called magnetic twisting cytometry to deliver small forces to specific sites on the cell membrane, he also demonstrated that cytoskeleton tension plays a dominant role in dictating cell shear stiffness and thus cell shape stability. He also showed that these localized forces cause cytoplasmic and nuclear deformation in remote parts of the cell. Recent collaborative work with colleagues at Harvard and Boston universities resulted in a fundamental finding that cells can adjust their internal response to external forces depending on how fast or slow these forces are applied. His research group currently has active collaborations with five different laboratories on campus, including the Bioengineering Department, the Beckman Institute, the Department of Cell and Developmental Biology, the Department of Chemical and Biomolecular Engineering, and the College of Veterinary Medicine.
Selected Articles in Journals
- Tabdili, H., M. D. Langer, Q. Shi, Y. C. Poh, N. Wang, and D. Leckband, "Cadherin-Dependent Mechanotransduction Depends on Ligand Identity but not Affinity," J Cell Sci. 2012 Jun 20. [Epub ahead of print]. PMID: 22718345.
- Liu, J., T. T. Youhua, Z. H. Huafeng, Y. Zhang, P. Xu, C. J. Junwei, Y. C. Poh, K. Tang, N. Wang*, and B. Huang*, "Soft Fibrin Gels Promote Selection and Growth of Tumorigenic Cells," Nature Materials. 11(8):734-41, 2012. PMID:22751180. PMC3405191. * Co-corresponding senior authors.
- Minary-Jolandan, M., A. Tajik, N. Wang, and M. F. Yu, "Intrinsically High-Q Dynamic AFM Imaging in Liquid with a Significantly Extended Needle Tip," Nanotechnology 23:235704, 2012. PMID: 22595833 NIHMS379051.
- Poh, Y. C., S. P. Shevtsov, F. Chowdhury, D. C. Wu, S. Na, M. Dundr, and N. Wang, "Dynamic Force-induced Direct Dissociation of Protein Complexes in a Nuclear Body in Living Cells," Nature Communications. 3: 866 doi:10.1038/ncomms1873. 2012. PMID: 22643893.
- Leckband, D. E., Q. le Duc, N. Wang, and J. de Rooij, "Mechanotransduction at Cadherin-mediated Adhesions," Curr. Opin. Cell Biol., Oct;23(5):523-30, 2011. Epub 2011 Sep 2. PMID: 2189033.
- Li, D., J. Zhou, F. Chowdhury, J. Cheng, N. Wang, and F. Wang, "The Role of Mechanical Factors in Fate Decisions of Stem Cells," Regenerative Medicine, Mar;6(2):229-40, 2011. PMID: 21391856 PMC3128460.
- Stamenović, D. and N. Wang, "Stress Transmission within the Cell," Comprehensive Physiology, Volume 1, Issue 1, January 2011. 10.1002/cphy.c100019.
- Uda, Y., Y. C. Poh, F. Chowdhury, D. C. Wu, T. S. Tanaka, M. Sato, and N. Wang, "Force via Integrins but not E-cadherin Decreases Oct3/4 Expression in Embryonic Stem Cells," Biochem Biophys Research Commun, Nov 18;415(2):396-400, 2011. PMID: 22037576. PMC3221912.
- Chowdhury F., Y. Li, Y.C. Poh, T. Tamaki, N. Wang*, and T. S. Tanaka*, "Soft Substrates Promote Homogeneous Self-renewal of Embryonic Stem Cells via Downregulating Cell-matrix Tractions," PLoS ONE, 5(12): e15655, 2010. doi:10.1371/journal.pone.0015655. PMID: 21179449. PMC3001487. *Co-corresponding senior authors.
- Yum, K., N. Wang*, and M. F. Yu*, "Electrochemically-controlled Deconjugation and Delivery Single Quantum Dots into the Nucleus of Living Cells," Small,6(19):2109-13, 2010. *Co-Corresponding senior authors.
- Shin, M. E., Y. He, D. Li, S. Na, F. Chowdhury, Y. C. Poh, O. Collin, P. Su, P. de Lanerolle, M. A. Schwartz, N. Wang, and F. Wang, "Spatiotemporal Organization, Regulation and Functions of Tractions during Neutrophil Chemotaxis," Blood, July 8, 2010. [Epub ahead of print] PMID: 20616216.
- le Duc, Q., Q. Shi, I. Blonk, A. Sonnenberg, N. Wang*, D. Leckband*, and J. de Rooij*, "Vinculin Potentiates E-cadherin Mechanosensing and is Recruited to Actin-anchored Sites within Adherens Junctions in a MyosinII Dependent Manner," J Cell Biol., 189:1107-1115, June 28, 2010. (This paper is commented and highlighted by M. Smutny, AS Yap. Neighborly relations: cadherins and mechanotransduction. J Cell Biol.189:1075-1077, 2010. Published June 28, 2010) *Co-Corresponding senior authors.
- Poh, Y.C., F. Chowdhury, T. S. Tanaka, and N. Wang, "Embryonic Stem Cells do not Stiffen on Rigid Substrates," Biophys. J. 99:L01-L03, 2010. PMID: 20643049.
- Wang, N., "Structural Basis of Stress Concentration in the Cytoskeleton," Molecular Cell Biomechanics, 7:33-44, 2010. PMID: 20806722.
- Yum, K., S. Na, K. Xiang, N. Wang, and M. F. Yu, "Mechanochemical Delivery and Dynamic Tracking of Single Fluorescent Quantum Dots in Living Cells," Nano Letters, 9(5):2193-8, May 2009. PMID: 19366190.
- !Chowdhury, F., S. Na, D. Li, Y.C. Poh, T. Tanaka, F. Wang, and N. Wang, "Material Properties of the Cell Dictate Stress-induced Spreading and Differentiation in Embryonic Stem Cells," Nat. Materials, Jan; 9(1):82-8, 2010. Publication online 18 October, 2009. DOI: 10.1038/NMAT2563. PMID: 19838182 PMC2833279 (Commented in News and Views of the same issue of Nat Mater. by Holle and Engler, “Cell rheology: Stressed-out stem cells” 9:4-6, 2010).
- Poh, Y. C., S. Na, F. Chowdhury, M. Ouyoung, Y. Wang, and N. Wang, "Rapid Activation of Rac GTPase in Living Cells by Force is Independent of Src," PLoS ONE, 4(11):e7886, Nov 18, 2009. PMID: 19924282.
- Kasza K. E., F. Nakamura, S. Hu, P. Kollmannsberger, B. Fabry, T. P. Stossel, N. Wang, and D. A. Weitz, “Filamin A is Essential for Active Cell Stiffening But Not Passive Stiffening under External Force,” Biophys. J., 96(10):4326-35, May 20, 2009. PMID: 19450503.
- Kim, T. J., J. Seong, M. Ouyang, J. Sun, S. Lu, J. P. Hong, N. Wang, and Y. Wang, “Substrate Rigidity Regulates Ca(2+) Oscillation via RhoA Pathway In Stem Cells,” J Cell Physiol, 218, 285-293, 2009. PMID: 1884423.
- Na, S., F. Chowdhury, B. Tay, M. Ouyang, M. Gregor, Y. Wang, G. Wiche, and N. Wang, “Plectin Contributes to Mechanical Properties of Living Cells,” Am J Physiol Cell Physiol, 296(4): C868-77, 2009. Feb 25, 2009. [Epub ahead of print] PMID: 19244477
- Chowdhury F., S. Na, O. Collin, B. Tay, F. Li, T. Tanaka, D.E. Leckband, and N. Wang, “Is cell rheology governed by nonequilibrium to equilibrium transition of noncovalent bonds?,” Biophysical Journal 95, 719-27, 2008. Epub 2008 Oct 3. PMID: 18835892.
- Collin, O., S. Na, F. Chowdhury, M. Hong, M. Shin, F. Wang, and N. Wang, “Self-organized Podosomes are Dynamic Mechanosensors," Current Biology, 8, 1288-1294, 2008.
- Na, S. and N. Wang, “Application of Fluorescence Resonance Energy Transfer and Magnetic Twisting Cyometry to Quantify Mechano-Chemical Signaling Activities in a Living Cell,” Science Signaling, 1, 11, 2008 (Cover highlight).
- Na, S., O. Collin, F. Chowdhury, B. Tay, M. Ouyang, Y. Wang, and N. Wang, “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, 2008. This paper is highlighted in Editor’s Choice Section in Science Signaling, the weekly journal and knowledge environment from the publishers of Science magazine, A. M. VanHook, “Faster than a Speeding Molecule,” Science Signaling, 1, ec175, 2008.
- Stamenovic, D., N. Rosenblatt, M. Montoya-Zavala, M. D. Matthews, S. Hu, B. Suki, N. Wang, and D. E. Ingber, “Rheological Behavior of Living Cells is Timescale Dependent,” Biophysical Journal, 93, L39-L41, 2007.
- Bursac, P., B. Fabry, X. Trepat, G. Lenormand, J. P. Butler, N. Wang, J. J. Fredberg, and S. S. An, “Cytoskeleton dynamics: Fluctuations within the Network,” Biochemical and Biophysical Research Communication, 355, 324-330, 2007.
- Rosenblatt, N., S. Hu, B. Suki, N. Wang, and D. Stamenovic, ”Contributions of the Active and Passive Components of the Cytoskeletal Prestress to Stiffening of Airway Smooth Muscle Cells,” Annual Biomedical Engineering, 35, 224-34, 2007.
- Hu, S. and N. Wang, “Control of Stress Propagation in the Cytoplasm by Prestress and Loading Frequency,” Molecular Cell Biomechanics, 3, 49-60, 2006.
- An, S. S., B. Fabry, X. Trepat, N. Wang, and J. J. Fredberg, “Do Biophysical Properties of the Airway Smooth Muscle Cell in Culture Predict Airway Hyperresponsiveness in Vivo?” American Journal of Physiology Lung Cellular Molecular Physiology, 35, 55-64, 2006.
- An, S. S., C. M. Pennella, A. Gonnabathula, J. Chen, N. Wang, M. Gaestel, P. M. Hassoun, J. J. Fredberg, and U. S. Kayyali, “Hypoxia Alters Distinct Biomechanical Properties of Endothelial Cells Via p38 MAPK and Rho Kinase-dependent Pathways,” American Journal of Physiology Cell Physiology, 289, C521-C530, 2005.
- Tolić-Norrelykke, I. M. and N. Wang, “Traction in Smooth Muscle Cells Varies with Cell Spreading,” Journal of Biomechanics, 38, 1405-1412, 2005.
- Hu, S., J. Chen, J. P. Butler, and N. Wang, “Prestress Mediates Force Propagation into the Nucleus,” Biochemical and Biophysical Research Communications, 329, 423-428, 2005.
- Wang, N. and Z. Suo, “Long-distance Propagation of Forces in a Cell,” Biochemical and Biophysical Research Communications, 328, 1133-1138, 2005.
- Rosenblatt, N., S. Hu, J. Chen, N. Wang, and D. Stamenovi, “Distending Stress of the Cytoskeleton is a Key Determinant of Cell Rheological Behavior,” Biochemical and Biophysical Research Communications, 321, 617-622, 2004.
- ! Hu, S., L. Eberhard, J. Chen, J. C. Love, J. P. Butler, J. J. Fredberg, G. M. Whitesides, and N. Wang, “Mechanical Anisotropy of Adherent Cells Probed by a Three Dimensional Magnetic Twisting Device,” American Journal of Physiology Cell Physiology, 287, C1184-C1191, 2004.
- Pui-de-Morales, M., E. Millet, B. Fabry, D. Navajas, N. Wang, J. P. Butler, and J. J. Fredberg, ”Cytoskeletal Mechanics in the Adherent Human Airway Smooth Muscle Cell: Probe Specificity and Scaling of Protein-Protein Dynamics,” American Journal of Physiology Cell Physiology, 287, C643-C654, 2004.
- Moy, A. B., K. Blackwell, N. Wang, K. Haxhinasto, M. K. Kasiske, D. M. Shasby, J. Bodmer, G. Reyes, and A. English, “Phorbol Ester-mediated Pulmonary Artery Endothelial Barrier Dysfunction Through Regulation of Actin Cytoskeletal Mechanics,” American Journal of Physiology Lung Cellular Molecular Physiology, 287, L153-L167, 2004.
- Stamenović, D., B. Suki, B. Fabry, N. Wang, J. J. Fredberg, and J. E. Buy, “Rheology of Airway Smooth Muscle Cells is Associated with Cytoskeletal Contractile Stress,” Journal of Applied Physiology, 96, 1600-1605, 2004.
- Hu, S., J. Chen, and N. Wang, “Cell Spreading Controls Balance of Prestress by Microtubules and Extracellular Matrix,” Frontiers in Bioscience, 9, 2177-2182, 2004.
- Polte, T. R., G. S. Eichler, N. Wang, and D. E. Ingber, “Extracellular Matrix Controls Myosin Light Chain Phosphorylation and Cell Contractility Through Modulation of Cell Shape and Cytoskeletal Prestress,” American Journal of Physiology Cell, 286, C518-28, 2004.
- Tolić-Norrelykke, I. M. and N. Wang, “Traction, Trypsin, and Tensegrity,” Internet Electron Journal of Molecular Design, 2, 642-652, 2003.
- Hu, S., J. Chen, B. Fabry, Y. Numaguchi, A. Gouldstone, D. E. Ingber, J. J. Fredberg, J. P. Butler, and N. Wang, “Intracellular Stress Tomography Reveals Stress Focusing and Structural Anisotropy in the Cytoskeleton of Living Cells,” American Journal of Physiology Cell, 285, C1082-C1090, 2003.
- Numaguchi, Y., S. Huang, T. R. Polte, G. S. Eichler, N. Wang, and D. E. Ingber, ”Caldesmon-dependent Switching Between Capillary Endothelial Cell Growth and Apoptosis Through Modulation of Cell Shape and Contractility,” Angiogenesis, 6, 55-64, 2003.
- Wang, N. and D. Stamenvoic, “Mechanics of Vimentin Intermediate Filaments,” Journal of Muscle Research and Cell Motility, 23, 535-540, 2002.
- Stamenović, D., S. M. Mijailovich, I. M. Toli_-Norrelykke, and N. Wang, “Experimental Tests of the Cellular Tensegrity Hypothesis,” Biorheology, 40, 221-225, 2002.
- Tolić-Nørrelykke, I.M., J. P. Butler, J. Chen, and N. Wang, “Spatial and Temporal Traction Response in Human Airway Smooth Muscle Cells,” Am. J. Physiol. Cell., 283, C1254-C1266, 2002.
- Stamenović, D., Z. Liang, J. Chen, and N. Wang, “The Effect of the Cytoskeletal Prestress on the Mechanical Impedance of Cultured Airway Smooth Muscle Cells,” Journal of Applied Physiology, 92, 1443-1450, 2002.
- Parker, K. K., A. L. Brock, C. Brangwynne, R. Mannix, N. Wang, E. Ostuni, N. A. Geisse, J. C. Adams, G. M. Whiteside, and D. E. Ingber, “Directional Control of Lamellipodia Extension by Constraining Cell Shape and Orienting Cell Tractional Forces,” Federation of American Societies for Experimental Biology Journal, 16, 1195-1204, 2002.
- Wang, N., E. Ostuni, G. M. Whitesides, and D. E. Ingber, “Micropatterning Tracitonal Forces in Living Cells,” Cell Motility Cytoskeleton, 52, 97-106, 2002.
- Stamenović, D., S. M. Mijailovich, I. M. Tolić-Norrelykke, J. Chen, and N. Wang, “Cell Prestress. II. Contribution of Microtubules,” American Journal of Physiology Cell, 282, C617-C624, 2002.
- Wang, N., I. M. Tolić-Norrelykke, J. Chen, S. M. Mijailovich, J. P. Butler, J. J. Fredberg, and D. Stamenović, “Cell Prestress. I. Stiffness and Prestress are Closely Associated in Adherent Contractile Cells,” American Journal of Physiology Cell, 282, C606-C616, 2002.
- Chen, J., B. Fabry, E. L. Schiffrin, and N. Wang, “Twisting Integrin Receptors Increases Endothelin-1 Gene Expression,” Am. J. Physiol. Cell, 280, C1475-C1484, 2001.
- Wang, N. and D. Stamenović, “Contribution of Intermediate Filaments to Cell Stiffness, Stiffening, and Growth,” American Journal of Physiology Cell, 279, C188-C194, 2000.
- Cai, S., L. Pestic-Dragovich, M. E. O’Donnell, N. Wang, D. Ingber, E. Elson, and P. de Lanerolle, “Regulation of Cytoskeletal Mechanics and Cell Growth by Myosin Light Chain Phosphorylation,” American Journal Physiology Cell, 275, C1349-C1356, 1998.
- Eckes, B., D. Dogic, E. Colucci-Guyon, N. Wang, A. Maniotis, D. Ingber, A. Merckling, F. Langa, M. Aumailley, A. Delouvee, V. Koteliansky, C. Babinet, and T. Krieg, “Impaired Mechanical Stability, Migration, and Contractile Capacity in Vimentin-deficient Fibroblasts,” Journal of Cell Science, 111, 1897-1907, 1998.
- Wang, N., “Mechanical Interactions Among Cytoskeletal Filaments,” Hypertension, 32, 162-165, 1998.
- Pourati, J., A. Maniotis, D. Spiegel, J. L. Schaffer, J. P. Butler, J. J. Fredberg, D. E. Ingber, D. Stamenović, and N. Wang, “Is Cytoskeletal Tension a Major Determinant of Cell Deformability in Adherent Endothelial Cells?” American Journal of Physiology Cell, 274, C1283-C1289, 1998.
- Lee, K.-M., K. Y. Tsai, N. Wang, and D. E. Ingber, “Extracellular Matrix and Pulmonary Hypertension: Control of Vascular Smooth Muscle Cell Contratcility,” American Journal of Physiology Heart Circulatory Physiology, 274, H76-H82, 1998.
- Goldmann, W. H., R. Glaneder, M. Ludwig, W. Xu, E. Adamson, N. Wang, and R. M. Ezzell, “Differences in Elasticity of Vinculin-deficient F9 Cells Measured by Magnetometry and Atomic Force Microscopy,” Experimental Cell Research, 239, 235-242, 1998.
- Planus, E., G. Barlovatz-Meimon, R. A. Rogers, S. Bonavaud, D. E. Ingber, and N. Wang, “Binding of Urokinase to Plasminogen Activator Inhibitor Type-1 Mediates Cell Adhesion and Spreading, “ Journal of Cell Science, 110, 1091-1098, 1997.
- Ezzell, R. M., W. H. Goldman, N. Wang, N. Parasharama, and D. E. Ingber, “Vinculin is Required for Cell Spreading and Transmembrane Mechanical Coupling of Integrins to the Cytoskeleton,” Experimental Cell Research, 231, 14-26, 1997.
- Potard, U. S. B., J. P. Butler, and N. Wang, “Cytoskeletal Mechanics in Confluent Epithelial Cells Probed Through Integrins and E-cadherins,” American Journal of Physiology Cell Physiology, 272, C1654-C1663, 1997.
- Tagawa, H., N. Wang, T. Narishige, D. E. Ingber, M. Zile, and G. Cooper, “Cytoskeletal Mechanics in Pressure Overload Cardiac Hypertrophy,” Circulation Research, 80, 281-289, 1997.
- Hubmayr, R. D., S. Shore, J. J. Fredberg, E. Planus, R. A. Panettieri, Jr., W. Moller, J. Heyder, and N. Wang, “Pharmacologic Activation Changes Stiffness of Cultured Human Airway Smooth Muscle Cells,” American Journal of Physiology Cell, 271, C1660-C1668, 1996.
- Stamenović, D., J. J. Fredberg, N. Wang, P. Butler, and D. E. Ingber, ”A Microstructural Approach to Cytoskeletal Mechanics,” Journal of Theoretical Biology, 1181, 125-136, 1996.
- Yoshida, M., W. F. Westlin, N. Wang, D. E. Ingber, A. Rosenzweig, N. Resnick, and M. A. Gimbrone, Jr., “Leukocyte Adhesion to Vascular Endothelium Induces E-selectin Association with the Actin-cytoskeleton,” Journal of Cell Biology, 133, 445-455, 1996.
- Brown, R. E., C. E. Kovacs, J. P. Butler, N. Wang, J. L. Lehr, and R. B. Banzett, “The Avian Lung: Is There an Aerodynamic Expiratory Valve?” Journal of Experimental Biology, 198, 2349-2357, 1995.
- Ingber, D. E., D. Prusty, Z. Sun, H. Betensky, and N. Wang, ”Cell Shape, Cytoskeletal Mechanics, and Cell Cycle Control in Angiogenesis,” Journal of Biomechanics, 28, 1471-1484, 1995.
- Wang, N. and D. E. Ingber, “Probing Transmembrane Mechanical Coupling and Cytomechanics Using Magnetic Twisting Cytometry,” Biochemistry Cell Biology, 73, 327-335, 1995.
- Wang, N., E. Planus, M. Pouchelet, J. J. Fredberg, and G. Barlovatz-Meimon, ”Urokinase Receptor Mediates Mechanical Force Transfer Across the Cell Surface,” American Journal of Physiology Cell Physiology, 268, C1062-C1066, 1995.
- Wang, N. and D. E. Ingber, “Control of Cytoskeletal Mechanics by Extracellular Matrix, Cell Shape, and Mechanical Tension,” Biophysical Journal, 66, 1281-1289, 1994.
- Wang, N., J. P. Butler, and D. E. Ingber, “Mechanotransduction Across the Cell Surface and Through the Cytoskeleton,” Science, 260, 1124-1127, 1993.
- Wang, N., R. B. Banzett, C. S. Nations, and F. A. Jenkins, Jr., “An Aerodynamic Valve in the Avian Primary Bronchus,” Journal of Experimental Zoology, 262, 441-445, 1992.
- Banzett, R. B., C. S. Nations, N. Wang, J. P. Butler, and J. L. Lehr, “Mechanical Independence of Wingbeat and Breathing in Starlings,” Respiration Physiology, 89, 27-36, 1992.
- Banzett, R. B., C. S. Nations, N. Wang, J. J. Fredberg, and J. P. Butler, “Pressure Profiles Show Features Essential to Aerodynamic Valving in Geese,” Respiration Physiology, 84, 295-309, 1991.
- Wang, N., J. P. Butler, and R. B. Banzett, “Gas Exchange Across Avian Egg Shells Oscillates in Phase with Heartbeat,” Journal of Applied Physiology, 69, 1549-1552, 1990.
- Wang, N., R. B. Banzett, J. P. Butler, and J. J. Fredberg, “Bird Lung Models Show that Connective Inertia Effects Inspiratory Aerodynamic Valving,” Respiration Physiology, 73, 111-124, 1988.