Abstract

We have developed diffraction phase and fluorescence (DPF) microscopy as a new technique for simultaneous quantitative phase imaging and epi-fluorescence investigation of live cells. The DPF instrument consists of an interference microscope, which is incorporated into a conventional inverted fluorescence microscope. The quantitative phase images are characterized by sub-nanometer optical path-length stability over periods from milliseconds to a cell lifetime. The potential of the technique for quantifying rapid nanoscale motions in live cells is demonstrated by experiments on red blood cells, while the composite phase-fluorescence imaging mode is exemplified with mitotic kidney cells.

© 2006 Optical Society of America

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    [CrossRef] [PubMed]
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    [CrossRef]
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    [CrossRef] [PubMed]
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    [CrossRef]
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    [CrossRef] [PubMed]
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    [CrossRef] [PubMed]
  19. G. Popescu, K. Badizadegan, R. R. Dasari and M. S. Feld, "Observation of dynamic subdomains in red blood cells," J. Biomed. Opt. Lett. 11, 040503 (2006).
    [CrossRef]
  20. T. Ikeda, G. Popescu, R. R. Dasari and M. S. Feld, "Hilbert phase microscopy for investigating fast dynamics in transparent systems," Opt. Lett. 30,1165-1168 (2005).
    [CrossRef] [PubMed]
  21. G. Popescu, T. Ikeda, C. A. Best, K. Badizadegan, R. R. Dasari and M. S. Feld, "Erythrocyte structure and dynamics quantified by Hilbert phase microscopy," J. Biomed. Opt. Lett. 10, 060503 (2005).
    [CrossRef]
  22. G. Popescu, T. Ikeda, R. R. Dasari and M. S. Feld, "Diffraction phase microscopy for quantifying cell structure and dynamics," Opt. Lett. 31, 775-777 (2006).
    [CrossRef] [PubMed]
  23. F. Brochard and J. F. Lennon, "Frequency spectrum of the flicker phenomenon in erythrocytes," J. Physique 36, 1035-1047 (1975).
    [CrossRef]
  24. G. Popescu, T. Ikeda, K. Goda, C. A. Best, M. Laposata, S. Manley, R. R. Dasari, K. Badizadegan and M. S. Feld, "Optical measurement of cell membrane tension," Phys. Rev. Lett., under review.
    [PubMed]

2006

G. Popescu, K. Badizadegan, R. R. Dasari and M. S. Feld, "Observation of dynamic subdomains in red blood cells," J. Biomed. Opt. Lett. 11, 040503 (2006).
[CrossRef]

G. Popescu, T. Ikeda, R. R. Dasari and M. S. Feld, "Diffraction phase microscopy for quantifying cell structure and dynamics," Opt. Lett. 31, 775-777 (2006).
[CrossRef] [PubMed]

2005

2004

2003

D. J. Stephens and V. J. Allan, "Light microscopy techniques for live cell imaging," Science 300, 82-86 (2003).
[CrossRef] [PubMed]

2001

C. Yang, A. Wax, M. S. Hahn, K. Badizadegan, R. R. Dasari and M. S. Feld, "Phase-referenced interferometer with subwavelength and subhertz sensitivity applied to the study of cell membrane dynamics," Opt Lett. 26, 1271-1273 (2001).
[CrossRef]

2000

C. H. Yang, A. Wax, I. Georgakoudi, E. B. Hanlon, K. Badizadegan, R. R. Dasari and M. S. Feld, "Interferometric phase-dispersion microscopy," Opt Lett. 25, 1526-1528 (2000).
[CrossRef]

B. E. Allman, P. J. McMahon, J. B. Tiller, K. A. Nugent, D. Paganin, A. Barty, I. McNulty, S. P. Frigo, Y. X. Wang and C. C. Retsch, "Noninterferometric quantitative phase imaging with soft x rays," J. Opt. Soc. Am. A-Opt. Image Sci. Vis. 17,1732-1743 (2000).
[CrossRef] [PubMed]

S. Bajt, A. Barty, K. A. Nugent, M. McCartney, M. Wall and D. Paganin, "Quantitative phase-sensitive imaging in a transmission electron microscope", Ultramicroscopy 83, 67-73 (2000).
[CrossRef] [PubMed]

1999

D. Zicha, E. Genot, G. A. Dunn and I. M. Kramer, "TGF beta 1 induces a cell-cycle-dependent increase in motility of epithelial cells," J. Cell Sci.,  112, 447-454 (1999).
[PubMed]

1998

D. Paganin and K. A. Nugent, "Noninterferometric phase imaging with partially coherent light," Phys. Rev. Lett. 80,2586-2589 (1998).
[CrossRef]

1997

G. A. Dunn, D. Zicha and P. E. Fraylich, "Rapid, microtubule-dependent fluctuations of the cell margin," J. Cell Sci. 110, 3091-3098 (1997).

1995

D. Zicha and G. A. Dunn, "An Image-Processing System For Cell Behavior Studies In Subconfluent Cultures," J. Microscopy,  179, 11-21 (1995).
[CrossRef]

1975

F. Brochard and J. F. Lennon, "Frequency spectrum of the flicker phenomenon in erythrocytes," J. Physique 36, 1035-1047 (1975).
[CrossRef]

Akkin, T.

Allan, V. J.

D. J. Stephens and V. J. Allan, "Light microscopy techniques for live cell imaging," Science 300, 82-86 (2003).
[CrossRef] [PubMed]

Allman, B. E.

B. E. Allman, P. J. McMahon, J. B. Tiller, K. A. Nugent, D. Paganin, A. Barty, I. McNulty, S. P. Frigo, Y. X. Wang and C. C. Retsch, "Noninterferometric quantitative phase imaging with soft x rays," J. Opt. Soc. Am. A-Opt. Image Sci. Vis. 17,1732-1743 (2000).
[CrossRef] [PubMed]

Badizadegan, K.

G. Popescu, K. Badizadegan, R. R. Dasari and M. S. Feld, "Observation of dynamic subdomains in red blood cells," J. Biomed. Opt. Lett. 11, 040503 (2006).
[CrossRef]

G. Popescu, T. Ikeda, C. A. Best, K. Badizadegan, R. R. Dasari and M. S. Feld, "Erythrocyte structure and dynamics quantified by Hilbert phase microscopy," J. Biomed. Opt. Lett. 10, 060503 (2005).
[CrossRef]

H. Iwai, C. Fang-Yen, G. Popescu, A. Wax, K. Badizadegan, R. R. Dasari and M. S. Feld, "Quantitative phase imaging using actively stabilized phase-shifting low-coherence interferometry," Opt. Lett. 29, 2399-2401 (2004).
[CrossRef] [PubMed]

G. Popescu, L. P. Deflores, J. C. Vaughan, K. Badizadegan, H. Iwai, R. R. Dasari and M. S. Feld, "Fourier phase microscopy for investigation of biological structures and dynamics," Opt. Lett. 29, 2503-2505 (2004).
[CrossRef] [PubMed]

C. Yang, A. Wax, M. S. Hahn, K. Badizadegan, R. R. Dasari and M. S. Feld, "Phase-referenced interferometer with subwavelength and subhertz sensitivity applied to the study of cell membrane dynamics," Opt Lett. 26, 1271-1273 (2001).
[CrossRef]

C. H. Yang, A. Wax, I. Georgakoudi, E. B. Hanlon, K. Badizadegan, R. R. Dasari and M. S. Feld, "Interferometric phase-dispersion microscopy," Opt Lett. 25, 1526-1528 (2000).
[CrossRef]

G. Popescu, T. Ikeda, K. Goda, C. A. Best, M. Laposata, S. Manley, R. R. Dasari, K. Badizadegan and M. S. Feld, "Optical measurement of cell membrane tension," Phys. Rev. Lett., under review.
[PubMed]

Bajt, S.

S. Bajt, A. Barty, K. A. Nugent, M. McCartney, M. Wall and D. Paganin, "Quantitative phase-sensitive imaging in a transmission electron microscope", Ultramicroscopy 83, 67-73 (2000).
[CrossRef] [PubMed]

Barty, A.

S. Bajt, A. Barty, K. A. Nugent, M. McCartney, M. Wall and D. Paganin, "Quantitative phase-sensitive imaging in a transmission electron microscope", Ultramicroscopy 83, 67-73 (2000).
[CrossRef] [PubMed]

B. E. Allman, P. J. McMahon, J. B. Tiller, K. A. Nugent, D. Paganin, A. Barty, I. McNulty, S. P. Frigo, Y. X. Wang and C. C. Retsch, "Noninterferometric quantitative phase imaging with soft x rays," J. Opt. Soc. Am. A-Opt. Image Sci. Vis. 17,1732-1743 (2000).
[CrossRef] [PubMed]

Best, C. A.

G. Popescu, T. Ikeda, C. A. Best, K. Badizadegan, R. R. Dasari and M. S. Feld, "Erythrocyte structure and dynamics quantified by Hilbert phase microscopy," J. Biomed. Opt. Lett. 10, 060503 (2005).
[CrossRef]

G. Popescu, T. Ikeda, K. Goda, C. A. Best, M. Laposata, S. Manley, R. R. Dasari, K. Badizadegan and M. S. Feld, "Optical measurement of cell membrane tension," Phys. Rev. Lett., under review.
[PubMed]

Brochard, F.

F. Brochard and J. F. Lennon, "Frequency spectrum of the flicker phenomenon in erythrocytes," J. Physique 36, 1035-1047 (1975).
[CrossRef]

Cense, B.

Choma, M. A.

Chu, M. C.

Colomb, T.

Creazzo, T. L.

Cuche, E.

Dasari, R. R.

G. Popescu, T. Ikeda, R. R. Dasari and M. S. Feld, "Diffraction phase microscopy for quantifying cell structure and dynamics," Opt. Lett. 31, 775-777 (2006).
[CrossRef] [PubMed]

G. Popescu, K. Badizadegan, R. R. Dasari and M. S. Feld, "Observation of dynamic subdomains in red blood cells," J. Biomed. Opt. Lett. 11, 040503 (2006).
[CrossRef]

T. Ikeda, G. Popescu, R. R. Dasari and M. S. Feld, "Hilbert phase microscopy for investigating fast dynamics in transparent systems," Opt. Lett. 30,1165-1168 (2005).
[CrossRef] [PubMed]

G. Popescu, T. Ikeda, C. A. Best, K. Badizadegan, R. R. Dasari and M. S. Feld, "Erythrocyte structure and dynamics quantified by Hilbert phase microscopy," J. Biomed. Opt. Lett. 10, 060503 (2005).
[CrossRef]

H. Iwai, C. Fang-Yen, G. Popescu, A. Wax, K. Badizadegan, R. R. Dasari and M. S. Feld, "Quantitative phase imaging using actively stabilized phase-shifting low-coherence interferometry," Opt. Lett. 29, 2399-2401 (2004).
[CrossRef] [PubMed]

G. Popescu, L. P. Deflores, J. C. Vaughan, K. Badizadegan, H. Iwai, R. R. Dasari and M. S. Feld, "Fourier phase microscopy for investigation of biological structures and dynamics," Opt. Lett. 29, 2503-2505 (2004).
[CrossRef] [PubMed]

C. Fang-Yen, M. C. Chu, H. S. Seung, R. R. Dasari and M. S. Feld, "Noncontact measurement of nerve displacement during action potential with a dual-beam low-coherence interferometer," Opt. Lett. 29, 2028-2030 (2004).
[CrossRef] [PubMed]

C. Yang, A. Wax, M. S. Hahn, K. Badizadegan, R. R. Dasari and M. S. Feld, "Phase-referenced interferometer with subwavelength and subhertz sensitivity applied to the study of cell membrane dynamics," Opt Lett. 26, 1271-1273 (2001).
[CrossRef]

C. H. Yang, A. Wax, I. Georgakoudi, E. B. Hanlon, K. Badizadegan, R. R. Dasari and M. S. Feld, "Interferometric phase-dispersion microscopy," Opt Lett. 25, 1526-1528 (2000).
[CrossRef]

G. Popescu, T. Ikeda, K. Goda, C. A. Best, M. Laposata, S. Manley, R. R. Dasari, K. Badizadegan and M. S. Feld, "Optical measurement of cell membrane tension," Phys. Rev. Lett., under review.
[PubMed]

Dave, D. P.

de Boer, J. E.

Deflores, L. P.

Depeursinge, C.

Diller, K. R.

Dunn, G. A.

D. Zicha, E. Genot, G. A. Dunn and I. M. Kramer, "TGF beta 1 induces a cell-cycle-dependent increase in motility of epithelial cells," J. Cell Sci.,  112, 447-454 (1999).
[PubMed]

G. A. Dunn, D. Zicha and P. E. Fraylich, "Rapid, microtubule-dependent fluctuations of the cell margin," J. Cell Sci. 110, 3091-3098 (1997).

D. Zicha and G. A. Dunn, "An Image-Processing System For Cell Behavior Studies In Subconfluent Cultures," J. Microscopy,  179, 11-21 (1995).
[CrossRef]

Ellerbee, A. K.

Emery, Y.

Fang-Yen, C.

Feld, M. S.

G. Popescu, T. Ikeda, R. R. Dasari and M. S. Feld, "Diffraction phase microscopy for quantifying cell structure and dynamics," Opt. Lett. 31, 775-777 (2006).
[CrossRef] [PubMed]

G. Popescu, K. Badizadegan, R. R. Dasari and M. S. Feld, "Observation of dynamic subdomains in red blood cells," J. Biomed. Opt. Lett. 11, 040503 (2006).
[CrossRef]

T. Ikeda, G. Popescu, R. R. Dasari and M. S. Feld, "Hilbert phase microscopy for investigating fast dynamics in transparent systems," Opt. Lett. 30,1165-1168 (2005).
[CrossRef] [PubMed]

G. Popescu, T. Ikeda, C. A. Best, K. Badizadegan, R. R. Dasari and M. S. Feld, "Erythrocyte structure and dynamics quantified by Hilbert phase microscopy," J. Biomed. Opt. Lett. 10, 060503 (2005).
[CrossRef]

G. Popescu, L. P. Deflores, J. C. Vaughan, K. Badizadegan, H. Iwai, R. R. Dasari and M. S. Feld, "Fourier phase microscopy for investigation of biological structures and dynamics," Opt. Lett. 29, 2503-2505 (2004).
[CrossRef] [PubMed]

H. Iwai, C. Fang-Yen, G. Popescu, A. Wax, K. Badizadegan, R. R. Dasari and M. S. Feld, "Quantitative phase imaging using actively stabilized phase-shifting low-coherence interferometry," Opt. Lett. 29, 2399-2401 (2004).
[CrossRef] [PubMed]

C. Fang-Yen, M. C. Chu, H. S. Seung, R. R. Dasari and M. S. Feld, "Noncontact measurement of nerve displacement during action potential with a dual-beam low-coherence interferometer," Opt. Lett. 29, 2028-2030 (2004).
[CrossRef] [PubMed]

C. Yang, A. Wax, M. S. Hahn, K. Badizadegan, R. R. Dasari and M. S. Feld, "Phase-referenced interferometer with subwavelength and subhertz sensitivity applied to the study of cell membrane dynamics," Opt Lett. 26, 1271-1273 (2001).
[CrossRef]

C. H. Yang, A. Wax, I. Georgakoudi, E. B. Hanlon, K. Badizadegan, R. R. Dasari and M. S. Feld, "Interferometric phase-dispersion microscopy," Opt Lett. 25, 1526-1528 (2000).
[CrossRef]

G. Popescu, T. Ikeda, K. Goda, C. A. Best, M. Laposata, S. Manley, R. R. Dasari, K. Badizadegan and M. S. Feld, "Optical measurement of cell membrane tension," Phys. Rev. Lett., under review.
[PubMed]

Fraylich, P. E.

G. A. Dunn, D. Zicha and P. E. Fraylich, "Rapid, microtubule-dependent fluctuations of the cell margin," J. Cell Sci. 110, 3091-3098 (1997).

Frigo, S. P.

B. E. Allman, P. J. McMahon, J. B. Tiller, K. A. Nugent, D. Paganin, A. Barty, I. McNulty, S. P. Frigo, Y. X. Wang and C. C. Retsch, "Noninterferometric quantitative phase imaging with soft x rays," J. Opt. Soc. Am. A-Opt. Image Sci. Vis. 17,1732-1743 (2000).
[CrossRef] [PubMed]

Genot, E.

D. Zicha, E. Genot, G. A. Dunn and I. M. Kramer, "TGF beta 1 induces a cell-cycle-dependent increase in motility of epithelial cells," J. Cell Sci.,  112, 447-454 (1999).
[PubMed]

Georgakoudi, I.

C. H. Yang, A. Wax, I. Georgakoudi, E. B. Hanlon, K. Badizadegan, R. R. Dasari and M. S. Feld, "Interferometric phase-dispersion microscopy," Opt Lett. 25, 1526-1528 (2000).
[CrossRef]

Goda, K.

G. Popescu, T. Ikeda, K. Goda, C. A. Best, M. Laposata, S. Manley, R. R. Dasari, K. Badizadegan and M. S. Feld, "Optical measurement of cell membrane tension," Phys. Rev. Lett., under review.
[PubMed]

Hahn, M. S.

C. Yang, A. Wax, M. S. Hahn, K. Badizadegan, R. R. Dasari and M. S. Feld, "Phase-referenced interferometer with subwavelength and subhertz sensitivity applied to the study of cell membrane dynamics," Opt Lett. 26, 1271-1273 (2001).
[CrossRef]

Hanlon, E. B.

C. H. Yang, A. Wax, I. Georgakoudi, E. B. Hanlon, K. Badizadegan, R. R. Dasari and M. S. Feld, "Interferometric phase-dispersion microscopy," Opt Lett. 25, 1526-1528 (2000).
[CrossRef]

Ikeda, T.

G. Popescu, T. Ikeda, R. R. Dasari and M. S. Feld, "Diffraction phase microscopy for quantifying cell structure and dynamics," Opt. Lett. 31, 775-777 (2006).
[CrossRef] [PubMed]

G. Popescu, T. Ikeda, C. A. Best, K. Badizadegan, R. R. Dasari and M. S. Feld, "Erythrocyte structure and dynamics quantified by Hilbert phase microscopy," J. Biomed. Opt. Lett. 10, 060503 (2005).
[CrossRef]

T. Ikeda, G. Popescu, R. R. Dasari and M. S. Feld, "Hilbert phase microscopy for investigating fast dynamics in transparent systems," Opt. Lett. 30,1165-1168 (2005).
[CrossRef] [PubMed]

G. Popescu, T. Ikeda, K. Goda, C. A. Best, M. Laposata, S. Manley, R. R. Dasari, K. Badizadegan and M. S. Feld, "Optical measurement of cell membrane tension," Phys. Rev. Lett., under review.
[PubMed]

Iwai, H.

Izatt, J. A.

Joo, C.

Kramer, I. M.

D. Zicha, E. Genot, G. A. Dunn and I. M. Kramer, "TGF beta 1 induces a cell-cycle-dependent increase in motility of epithelial cells," J. Cell Sci.,  112, 447-454 (1999).
[PubMed]

Laposata, M.

G. Popescu, T. Ikeda, K. Goda, C. A. Best, M. Laposata, S. Manley, R. R. Dasari, K. Badizadegan and M. S. Feld, "Optical measurement of cell membrane tension," Phys. Rev. Lett., under review.
[PubMed]

Lennon, J. F.

F. Brochard and J. F. Lennon, "Frequency spectrum of the flicker phenomenon in erythrocytes," J. Physique 36, 1035-1047 (1975).
[CrossRef]

Magistretti, P. J.

Manley, S.

G. Popescu, T. Ikeda, K. Goda, C. A. Best, M. Laposata, S. Manley, R. R. Dasari, K. Badizadegan and M. S. Feld, "Optical measurement of cell membrane tension," Phys. Rev. Lett., under review.
[PubMed]

Mann, C. J.

Marquet, P.

McCartney, M.

S. Bajt, A. Barty, K. A. Nugent, M. McCartney, M. Wall and D. Paganin, "Quantitative phase-sensitive imaging in a transmission electron microscope", Ultramicroscopy 83, 67-73 (2000).
[CrossRef] [PubMed]

McMahon, P. J.

B. E. Allman, P. J. McMahon, J. B. Tiller, K. A. Nugent, D. Paganin, A. Barty, I. McNulty, S. P. Frigo, Y. X. Wang and C. C. Retsch, "Noninterferometric quantitative phase imaging with soft x rays," J. Opt. Soc. Am. A-Opt. Image Sci. Vis. 17,1732-1743 (2000).
[CrossRef] [PubMed]

McNulty, I.

B. E. Allman, P. J. McMahon, J. B. Tiller, K. A. Nugent, D. Paganin, A. Barty, I. McNulty, S. P. Frigo, Y. X. Wang and C. C. Retsch, "Noninterferometric quantitative phase imaging with soft x rays," J. Opt. Soc. Am. A-Opt. Image Sci. Vis. 17,1732-1743 (2000).
[CrossRef] [PubMed]

Milner, T. E.

Nugent, K. A.

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B. E. Allman, P. J. McMahon, J. B. Tiller, K. A. Nugent, D. Paganin, A. Barty, I. McNulty, S. P. Frigo, Y. X. Wang and C. C. Retsch, "Noninterferometric quantitative phase imaging with soft x rays," J. Opt. Soc. Am. A-Opt. Image Sci. Vis. 17,1732-1743 (2000).
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B. E. Allman, P. J. McMahon, J. B. Tiller, K. A. Nugent, D. Paganin, A. Barty, I. McNulty, S. P. Frigo, Y. X. Wang and C. C. Retsch, "Noninterferometric quantitative phase imaging with soft x rays," J. Opt. Soc. Am. A-Opt. Image Sci. Vis. 17,1732-1743 (2000).
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B. E. Allman, P. J. McMahon, J. B. Tiller, K. A. Nugent, D. Paganin, A. Barty, I. McNulty, S. P. Frigo, Y. X. Wang and C. C. Retsch, "Noninterferometric quantitative phase imaging with soft x rays," J. Opt. Soc. Am. A-Opt. Image Sci. Vis. 17,1732-1743 (2000).
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M. A. Choma, A. K. Ellerbee, C. H. Yang, T. L. Creazzo and J. A. Izatt, "Spectral-domain phase microscopy," Opt. Lett. 30, 1162-1164 (2005).
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G. Popescu, K. Badizadegan, R. R. Dasari and M. S. Feld, "Observation of dynamic subdomains in red blood cells," J. Biomed. Opt. Lett. 11, 040503 (2006).
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G. Popescu, T. Ikeda, C. A. Best, K. Badizadegan, R. R. Dasari and M. S. Feld, "Erythrocyte structure and dynamics quantified by Hilbert phase microscopy," J. Biomed. Opt. Lett. 10, 060503 (2005).
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G. A. Dunn, D. Zicha and P. E. Fraylich, "Rapid, microtubule-dependent fluctuations of the cell margin," J. Cell Sci. 110, 3091-3098 (1997).

D. Zicha, E. Genot, G. A. Dunn and I. M. Kramer, "TGF beta 1 induces a cell-cycle-dependent increase in motility of epithelial cells," J. Cell Sci.,  112, 447-454 (1999).
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B. E. Allman, P. J. McMahon, J. B. Tiller, K. A. Nugent, D. Paganin, A. Barty, I. McNulty, S. P. Frigo, Y. X. Wang and C. C. Retsch, "Noninterferometric quantitative phase imaging with soft x rays," J. Opt. Soc. Am. A-Opt. Image Sci. Vis. 17,1732-1743 (2000).
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Opt. Lett.

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G. Popescu, T. Ikeda, K. Goda, C. A. Best, M. Laposata, S. Manley, R. R. Dasari, K. Badizadegan and M. S. Feld, "Optical measurement of cell membrane tension," Phys. Rev. Lett., under review.
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D. Paganin and K. A. Nugent, "Noninterferometric phase imaging with partially coherent light," Phys. Rev. Lett. 80,2586-2589 (1998).
[CrossRef]

Science

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Ultramicroscopy

S. Bajt, A. Barty, K. A. Nugent, M. McCartney, M. Wall and D. Paganin, "Quantitative phase-sensitive imaging in a transmission electron microscope", Ultramicroscopy 83, 67-73 (2000).
[CrossRef] [PubMed]

Supplementary Material (2)

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Figures (3)

Fig. 1.
Fig. 1.

DPF setup. F1, 2, filters; M1, 2, mirrors; L1-4 lenses (f1-4, respective focal lengths); G, grating; SF, spatial filter; IP1, 2, image planes; SF, spatial filter.

Fig. 2.
Fig. 2.

(a). (4 Mb) MOVIE: Thermal fluctuations of red blood cell membrane. (b) (4 Mb) MOVIE: Eosinophil chemotaxis and attack on red blood cell. Color bars represent optical phase in radians.

Fig. 3.
Fig. 3.

(a). Quantitative phase image of a kidney cell. The colorbar indicates optical phase shift in radians. (b) Fluorescence image of the same DNA-stained cell. (c) Overlaid images from (a) and (b): red- DPM image and blue- epi-fluorescence image. Colorbar indicates phase in radians.

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