Abstract

Changes in the sensitivity of the fovea of one of the eyes (the right eye) under influence of a glaring source of light shining into that eye can be demonstrated by comparing the visual impression obtained by that eye from a dimly illuminated surface with the impression obtained by the other eye (the left eye) from a similar surface. By adjusting the illumination of one of the surfaces the apparent brightnesses can be made to match exactly. In this way a quantitative method of measuring the sensitivity of the fovea under influence of various conditions of illumination is obtained. When a small area of the retina is illuminated the sensitivity of this area and also of the surroundings drops to a considerably lower level within 0.1 second (α-adaptation). The rate of recovery of the sensitivity after exposure is strongly dependent upon the time of exposure (β-adaptation). α-adaptation can neither be accounted for by assuming a strong adaptive influence of the stray light within the eye, nor by assuming a lateral diffusion of photosensitive substances or of photolytic products over the retina. The hypothesis is put forward that α-adaptation is of electrical origin. This hypothesis accounts for many of the phenomena observed with incandescent lamp light but not for those observed with strongly chromatic light. The only hypothesis presenting itself at present to account for these phenomena seems to be that α-adaptation has its seat in the retinal synapses.

© 1939 Optical Society of America

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References

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  1. E. Hering, “Grundzüge der Lehre vom Lichtsinn,” Graefe Saemisch Handbuch Ges. Aug. (1905).
  2. N. Inouye and S. Oinuma, “Untersuchung der Dunkeladaptation des einen Auges mit Hilfe des helladaptierten Andern,” Gr. Arch. Ophth. 79, 145–159 (1911).
    [CrossRef]
  3. R. Dittler and I. Koïke, “Ueber die Adaptationsfähigkeit der Fovea Centralis,” Zeits. Psych. Physiol. Sinnesorg,  46, 166–178 (1912).
  4. W. D. Wright, “The measurement and analysis of colour adaptation phenomena,” Proc. Roy. Soc. B115, 49–87 (1934).
    [CrossRef]
  5. W. D. Wright, “The foveal light adaptation process,” Proc. Roy. Soc. B122, 220–245 (1937).
    [CrossRef]
  6. G. von Békésy, “Ueber die Bestimmung des einem reinen Tonempfinden entsprechenden Erregungsgebietes der Basilarmembran vermittelst Ermüdungserscheinungen,” Physik. Zeits. 30, 115–125 (1929).
  7. L. S. Ornstein and J. F. Schouten, “De rol van electrische, photochemische en diffusieverschynselen by het zien.” Proc. K. Akad. Amsterdam 40, 376–382 (1937).
  8. J. F. Schouten, “Visueele meting van adaptatie en van de wederzydsche beinvloeding van netvlieselementen,” Ac. Thesis (Utrecht, 1937).
  9. J. F. Schouten, “Zur Analyse der Blendung,” Proc. K. Akad. W. Amsterdam 37, 506–516 (1934).
  10. L. L. Holladay, “The Fundamentals of Glare and Visibility,” J. Opt. Soc. Am. 12, 271–319 (1926); J. Opt. Soc. Am. 14, 1–15 (1927).
    [CrossRef]
  11. Y. le Grand, “Recherches sur la diffusion de la lumière dans l’oeil humain,” Rev. d’optique 16, 201–214241–266 (1937).
  12. F. K. Moss, “A Modified Broca Pupillometer,” J. Opt. Soc. Am. 22, 735–738 (1932).
    [CrossRef]
  13. J. von Kries, “Theoretische Studien über die Umstimmung des Sehorgans,” Festschr. Univ. Freiburg143–158 (1902).
  14. W. S. Stiles and B. H. Crawford, “The luminous efficiency of rays entering the eye pupil at different points,” Proc. Roy. Soc. B112, 428–450 (1933). W. S. Stiles, “The luminous efficiency of monochromatic rays entering the eye pupil at different points and a new colour effect,” Proc. Roy. Soc. B123, 90–118 (1937). B. H. Crawford, “The luminous efficiency of light entering the eye pupil at different points and its relation to brightness threshold measurements,” Proc. Roy. Soc. B124, 81–96 (1937). W. D. Wright and J. H. Nelson, “The relation between the apparent intensity of a beam of light and the angle at which the beam strikes the retina,” Proc. Phys. Soc. 48, 401–405 (1936).
    [CrossRef]
  15. J. F. Schouten, “Eine entoptische Methode zur Bestimmung der spektralen Durchlässigkeit der Augapfelwand,” Proc. K. Akad. Amsterdam 37, 516–520 (1934).
  16. A. Broca and D. Sulzer, “La sensation lumineuse en fonction du temps,” J. physiol. et pathol. gén. 4, 632–640 (1902).
  17. S. Hecht, “The dark adaptation of the human eye,” J. Gen. Physiol. 2, 499–517 (1920).S. Hecht, “The nature of foveal dark adaptation,” J. Gen. Physiol. 4, 113–139 (1922).
    [CrossRef]
  18. See e.g. R. Granit, “Die Elektrophysiologie der Netzhaut und des Sehnerven,” Acta Ophth. Suppl. VIII,  14, 1–98 (1936).
  19. F. W. Fröhlich, “Beiträge zur allgemeinen Physiologie der Sinnesorgane,” Zeits. Sinnesphysiol. 48, 28–164, 354–438 (1914).
  20. A. Kohlrausch, “Elektrische Erscheinungen am Auge,” Bethes Handb. Norm. Path. Physiol. 12, No. 2, II, 1394–1496 (1931).
  21. P. J. Bouma, “Phenomena of eye fatigue and after images with monochromatic light,” Physica 1, 429–436 (1934).
    [CrossRef]
  22. S. M. Newhall, “The Constancy of the Blue Arc Phenomenon,” J. Opt. Soc. Am. 27, 165–176 (1937).
    [CrossRef]
  23. A. Goldmann, “Lichtelektrische Untersuchungen an Farbstoffzellen,” Ann. d. Physik 4, 27, 449–536 (1908).
  24. W. Vanselow and S. E. Sheppard, “Photo-Voltaic cells with silver-silver bromide electrodes I,” J. Phys. Chem. 33, 331–353 (1929).
    [CrossRef]

1937 (4)

W. D. Wright, “The foveal light adaptation process,” Proc. Roy. Soc. B122, 220–245 (1937).
[CrossRef]

L. S. Ornstein and J. F. Schouten, “De rol van electrische, photochemische en diffusieverschynselen by het zien.” Proc. K. Akad. Amsterdam 40, 376–382 (1937).

Y. le Grand, “Recherches sur la diffusion de la lumière dans l’oeil humain,” Rev. d’optique 16, 201–214241–266 (1937).

S. M. Newhall, “The Constancy of the Blue Arc Phenomenon,” J. Opt. Soc. Am. 27, 165–176 (1937).
[CrossRef]

1936 (1)

See e.g. R. Granit, “Die Elektrophysiologie der Netzhaut und des Sehnerven,” Acta Ophth. Suppl. VIII,  14, 1–98 (1936).

1934 (4)

P. J. Bouma, “Phenomena of eye fatigue and after images with monochromatic light,” Physica 1, 429–436 (1934).
[CrossRef]

J. F. Schouten, “Eine entoptische Methode zur Bestimmung der spektralen Durchlässigkeit der Augapfelwand,” Proc. K. Akad. Amsterdam 37, 516–520 (1934).

J. F. Schouten, “Zur Analyse der Blendung,” Proc. K. Akad. W. Amsterdam 37, 506–516 (1934).

W. D. Wright, “The measurement and analysis of colour adaptation phenomena,” Proc. Roy. Soc. B115, 49–87 (1934).
[CrossRef]

1933 (1)

W. S. Stiles and B. H. Crawford, “The luminous efficiency of rays entering the eye pupil at different points,” Proc. Roy. Soc. B112, 428–450 (1933). W. S. Stiles, “The luminous efficiency of monochromatic rays entering the eye pupil at different points and a new colour effect,” Proc. Roy. Soc. B123, 90–118 (1937). B. H. Crawford, “The luminous efficiency of light entering the eye pupil at different points and its relation to brightness threshold measurements,” Proc. Roy. Soc. B124, 81–96 (1937). W. D. Wright and J. H. Nelson, “The relation between the apparent intensity of a beam of light and the angle at which the beam strikes the retina,” Proc. Phys. Soc. 48, 401–405 (1936).
[CrossRef]

1932 (1)

1931 (1)

A. Kohlrausch, “Elektrische Erscheinungen am Auge,” Bethes Handb. Norm. Path. Physiol. 12, No. 2, II, 1394–1496 (1931).

1929 (2)

G. von Békésy, “Ueber die Bestimmung des einem reinen Tonempfinden entsprechenden Erregungsgebietes der Basilarmembran vermittelst Ermüdungserscheinungen,” Physik. Zeits. 30, 115–125 (1929).

W. Vanselow and S. E. Sheppard, “Photo-Voltaic cells with silver-silver bromide electrodes I,” J. Phys. Chem. 33, 331–353 (1929).
[CrossRef]

1926 (1)

1920 (1)

S. Hecht, “The dark adaptation of the human eye,” J. Gen. Physiol. 2, 499–517 (1920).S. Hecht, “The nature of foveal dark adaptation,” J. Gen. Physiol. 4, 113–139 (1922).
[CrossRef]

1914 (1)

F. W. Fröhlich, “Beiträge zur allgemeinen Physiologie der Sinnesorgane,” Zeits. Sinnesphysiol. 48, 28–164, 354–438 (1914).

1912 (1)

R. Dittler and I. Koïke, “Ueber die Adaptationsfähigkeit der Fovea Centralis,” Zeits. Psych. Physiol. Sinnesorg,  46, 166–178 (1912).

1911 (1)

N. Inouye and S. Oinuma, “Untersuchung der Dunkeladaptation des einen Auges mit Hilfe des helladaptierten Andern,” Gr. Arch. Ophth. 79, 145–159 (1911).
[CrossRef]

1908 (1)

A. Goldmann, “Lichtelektrische Untersuchungen an Farbstoffzellen,” Ann. d. Physik 4, 27, 449–536 (1908).

1905 (1)

E. Hering, “Grundzüge der Lehre vom Lichtsinn,” Graefe Saemisch Handbuch Ges. Aug. (1905).

1902 (2)

J. von Kries, “Theoretische Studien über die Umstimmung des Sehorgans,” Festschr. Univ. Freiburg143–158 (1902).

A. Broca and D. Sulzer, “La sensation lumineuse en fonction du temps,” J. physiol. et pathol. gén. 4, 632–640 (1902).

Bouma, P. J.

P. J. Bouma, “Phenomena of eye fatigue and after images with monochromatic light,” Physica 1, 429–436 (1934).
[CrossRef]

Broca, A.

A. Broca and D. Sulzer, “La sensation lumineuse en fonction du temps,” J. physiol. et pathol. gén. 4, 632–640 (1902).

Crawford, B. H.

W. S. Stiles and B. H. Crawford, “The luminous efficiency of rays entering the eye pupil at different points,” Proc. Roy. Soc. B112, 428–450 (1933). W. S. Stiles, “The luminous efficiency of monochromatic rays entering the eye pupil at different points and a new colour effect,” Proc. Roy. Soc. B123, 90–118 (1937). B. H. Crawford, “The luminous efficiency of light entering the eye pupil at different points and its relation to brightness threshold measurements,” Proc. Roy. Soc. B124, 81–96 (1937). W. D. Wright and J. H. Nelson, “The relation between the apparent intensity of a beam of light and the angle at which the beam strikes the retina,” Proc. Phys. Soc. 48, 401–405 (1936).
[CrossRef]

Dittler, R.

R. Dittler and I. Koïke, “Ueber die Adaptationsfähigkeit der Fovea Centralis,” Zeits. Psych. Physiol. Sinnesorg,  46, 166–178 (1912).

Fröhlich, F. W.

F. W. Fröhlich, “Beiträge zur allgemeinen Physiologie der Sinnesorgane,” Zeits. Sinnesphysiol. 48, 28–164, 354–438 (1914).

Goldmann, A.

A. Goldmann, “Lichtelektrische Untersuchungen an Farbstoffzellen,” Ann. d. Physik 4, 27, 449–536 (1908).

Granit, R.

See e.g. R. Granit, “Die Elektrophysiologie der Netzhaut und des Sehnerven,” Acta Ophth. Suppl. VIII,  14, 1–98 (1936).

Hecht, S.

S. Hecht, “The dark adaptation of the human eye,” J. Gen. Physiol. 2, 499–517 (1920).S. Hecht, “The nature of foveal dark adaptation,” J. Gen. Physiol. 4, 113–139 (1922).
[CrossRef]

Hering, E.

E. Hering, “Grundzüge der Lehre vom Lichtsinn,” Graefe Saemisch Handbuch Ges. Aug. (1905).

Holladay, L. L.

Inouye, N.

N. Inouye and S. Oinuma, “Untersuchung der Dunkeladaptation des einen Auges mit Hilfe des helladaptierten Andern,” Gr. Arch. Ophth. 79, 145–159 (1911).
[CrossRef]

Kohlrausch, A.

A. Kohlrausch, “Elektrische Erscheinungen am Auge,” Bethes Handb. Norm. Path. Physiol. 12, No. 2, II, 1394–1496 (1931).

Koïke, I.

R. Dittler and I. Koïke, “Ueber die Adaptationsfähigkeit der Fovea Centralis,” Zeits. Psych. Physiol. Sinnesorg,  46, 166–178 (1912).

le Grand, Y.

Y. le Grand, “Recherches sur la diffusion de la lumière dans l’oeil humain,” Rev. d’optique 16, 201–214241–266 (1937).

Moss, F. K.

Newhall, S. M.

Oinuma, S.

N. Inouye and S. Oinuma, “Untersuchung der Dunkeladaptation des einen Auges mit Hilfe des helladaptierten Andern,” Gr. Arch. Ophth. 79, 145–159 (1911).
[CrossRef]

Ornstein, L. S.

L. S. Ornstein and J. F. Schouten, “De rol van electrische, photochemische en diffusieverschynselen by het zien.” Proc. K. Akad. Amsterdam 40, 376–382 (1937).

Schouten, J. F.

L. S. Ornstein and J. F. Schouten, “De rol van electrische, photochemische en diffusieverschynselen by het zien.” Proc. K. Akad. Amsterdam 40, 376–382 (1937).

J. F. Schouten, “Zur Analyse der Blendung,” Proc. K. Akad. W. Amsterdam 37, 506–516 (1934).

J. F. Schouten, “Eine entoptische Methode zur Bestimmung der spektralen Durchlässigkeit der Augapfelwand,” Proc. K. Akad. Amsterdam 37, 516–520 (1934).

J. F. Schouten, “Visueele meting van adaptatie en van de wederzydsche beinvloeding van netvlieselementen,” Ac. Thesis (Utrecht, 1937).

Sheppard, S. E.

W. Vanselow and S. E. Sheppard, “Photo-Voltaic cells with silver-silver bromide electrodes I,” J. Phys. Chem. 33, 331–353 (1929).
[CrossRef]

Stiles, W. S.

W. S. Stiles and B. H. Crawford, “The luminous efficiency of rays entering the eye pupil at different points,” Proc. Roy. Soc. B112, 428–450 (1933). W. S. Stiles, “The luminous efficiency of monochromatic rays entering the eye pupil at different points and a new colour effect,” Proc. Roy. Soc. B123, 90–118 (1937). B. H. Crawford, “The luminous efficiency of light entering the eye pupil at different points and its relation to brightness threshold measurements,” Proc. Roy. Soc. B124, 81–96 (1937). W. D. Wright and J. H. Nelson, “The relation between the apparent intensity of a beam of light and the angle at which the beam strikes the retina,” Proc. Phys. Soc. 48, 401–405 (1936).
[CrossRef]

Sulzer, D.

A. Broca and D. Sulzer, “La sensation lumineuse en fonction du temps,” J. physiol. et pathol. gén. 4, 632–640 (1902).

Vanselow, W.

W. Vanselow and S. E. Sheppard, “Photo-Voltaic cells with silver-silver bromide electrodes I,” J. Phys. Chem. 33, 331–353 (1929).
[CrossRef]

von Békésy, G.

G. von Békésy, “Ueber die Bestimmung des einem reinen Tonempfinden entsprechenden Erregungsgebietes der Basilarmembran vermittelst Ermüdungserscheinungen,” Physik. Zeits. 30, 115–125 (1929).

von Kries, J.

J. von Kries, “Theoretische Studien über die Umstimmung des Sehorgans,” Festschr. Univ. Freiburg143–158 (1902).

Wright, W. D.

W. D. Wright, “The foveal light adaptation process,” Proc. Roy. Soc. B122, 220–245 (1937).
[CrossRef]

W. D. Wright, “The measurement and analysis of colour adaptation phenomena,” Proc. Roy. Soc. B115, 49–87 (1934).
[CrossRef]

Acta Ophth. Suppl. VIII (1)

See e.g. R. Granit, “Die Elektrophysiologie der Netzhaut und des Sehnerven,” Acta Ophth. Suppl. VIII,  14, 1–98 (1936).

Ann. d. Physik (1)

A. Goldmann, “Lichtelektrische Untersuchungen an Farbstoffzellen,” Ann. d. Physik 4, 27, 449–536 (1908).

Bethes Handb. Norm. Path. Physiol. (1)

A. Kohlrausch, “Elektrische Erscheinungen am Auge,” Bethes Handb. Norm. Path. Physiol. 12, No. 2, II, 1394–1496 (1931).

Festschr. Univ. Freiburg (1)

J. von Kries, “Theoretische Studien über die Umstimmung des Sehorgans,” Festschr. Univ. Freiburg143–158 (1902).

Gr. Arch. Ophth. (1)

N. Inouye and S. Oinuma, “Untersuchung der Dunkeladaptation des einen Auges mit Hilfe des helladaptierten Andern,” Gr. Arch. Ophth. 79, 145–159 (1911).
[CrossRef]

Graefe Saemisch Handbuch Ges. Aug (1)

E. Hering, “Grundzüge der Lehre vom Lichtsinn,” Graefe Saemisch Handbuch Ges. Aug. (1905).

J. Gen. Physiol. (1)

S. Hecht, “The dark adaptation of the human eye,” J. Gen. Physiol. 2, 499–517 (1920).S. Hecht, “The nature of foveal dark adaptation,” J. Gen. Physiol. 4, 113–139 (1922).
[CrossRef]

J. Opt. Soc. Am. (3)

J. Phys. Chem. (1)

W. Vanselow and S. E. Sheppard, “Photo-Voltaic cells with silver-silver bromide electrodes I,” J. Phys. Chem. 33, 331–353 (1929).
[CrossRef]

J. physiol. et pathol. gén. (1)

A. Broca and D. Sulzer, “La sensation lumineuse en fonction du temps,” J. physiol. et pathol. gén. 4, 632–640 (1902).

Physica (1)

P. J. Bouma, “Phenomena of eye fatigue and after images with monochromatic light,” Physica 1, 429–436 (1934).
[CrossRef]

Physik. Zeits. (1)

G. von Békésy, “Ueber die Bestimmung des einem reinen Tonempfinden entsprechenden Erregungsgebietes der Basilarmembran vermittelst Ermüdungserscheinungen,” Physik. Zeits. 30, 115–125 (1929).

Proc. K. Akad. Amsterdam (2)

L. S. Ornstein and J. F. Schouten, “De rol van electrische, photochemische en diffusieverschynselen by het zien.” Proc. K. Akad. Amsterdam 40, 376–382 (1937).

J. F. Schouten, “Eine entoptische Methode zur Bestimmung der spektralen Durchlässigkeit der Augapfelwand,” Proc. K. Akad. Amsterdam 37, 516–520 (1934).

Proc. K. Akad. W. Amsterdam (1)

J. F. Schouten, “Zur Analyse der Blendung,” Proc. K. Akad. W. Amsterdam 37, 506–516 (1934).

Proc. Roy. Soc. (3)

W. D. Wright, “The measurement and analysis of colour adaptation phenomena,” Proc. Roy. Soc. B115, 49–87 (1934).
[CrossRef]

W. D. Wright, “The foveal light adaptation process,” Proc. Roy. Soc. B122, 220–245 (1937).
[CrossRef]

W. S. Stiles and B. H. Crawford, “The luminous efficiency of rays entering the eye pupil at different points,” Proc. Roy. Soc. B112, 428–450 (1933). W. S. Stiles, “The luminous efficiency of monochromatic rays entering the eye pupil at different points and a new colour effect,” Proc. Roy. Soc. B123, 90–118 (1937). B. H. Crawford, “The luminous efficiency of light entering the eye pupil at different points and its relation to brightness threshold measurements,” Proc. Roy. Soc. B124, 81–96 (1937). W. D. Wright and J. H. Nelson, “The relation between the apparent intensity of a beam of light and the angle at which the beam strikes the retina,” Proc. Phys. Soc. 48, 401–405 (1936).
[CrossRef]

Rev. d’optique (1)

Y. le Grand, “Recherches sur la diffusion de la lumière dans l’oeil humain,” Rev. d’optique 16, 201–214241–266 (1937).

Zeits. Psych. Physiol. Sinnesorg (1)

R. Dittler and I. Koïke, “Ueber die Adaptationsfähigkeit der Fovea Centralis,” Zeits. Psych. Physiol. Sinnesorg,  46, 166–178 (1912).

Zeits. Sinnesphysiol. (1)

F. W. Fröhlich, “Beiträge zur allgemeinen Physiologie der Sinnesorgane,” Zeits. Sinnesphysiol. 48, 28–164, 354–438 (1914).

Other (1)

J. F. Schouten, “Visueele meting van adaptatie en van de wederzydsche beinvloeding van netvlieselementen,” Ac. Thesis (Utrecht, 1937).

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Figures (12)

Fig. 1
Fig. 1

Schematic representation of the different ways in which the perception of light by the fovea may be affected by the presence of a glaring light source.

Fig. 2
Fig. 2

Sketch of the apparatus.

Fig. 3
Fig. 3

Relative position of the glaring light source L, the test object T and the object of comparison C. L and T are only visible to the right eye, whereas C is only visible to the left eye.

Fig. 4
Fig. 4

Curves showing the brightness of diascleral re-illumination as a function of the position of the light spot on the sclerotic. Arbitrary scale of brightness.

Fig. 5
Fig. 5

Values of the glare index V for different points of fixation. Each value of V is represented at the corresponding point of fixation.

Fig. 6
Fig. 6

The slit-wheel. The glare source L becomes visible through slit S1, whereas the test object T and the object of comparison C become visible through slit S2.

Fig. 7
Fig. 7

Curve showing dependence of the sensitivity upon the time as measured with the slit-wheel, during the first 0.2 sec. of exposure. Glare angle φ=6°. Illumination at the eye 180 lux. Each point gives the mean of four readings.

Fig. 8
Fig. 8

Curves showing the dependence of the sensitivity ρ on the glare angle φ for different values of the illumination at the eye E.

Fig. 9
Fig. 9

Curves showing the recovery of the sensitivity ρ after direct adaptation as a function of the time of recovery t, for different brightnesses of the glare source. B=100 corresponds with 900 candles/m2 (5600 photons). Time of exposure τ=60 sec.

Fig. 10
Fig. 10

Curves showing the recovery of the sensitivity ρ as a function of the time of recovery t, for different times of exposure τ. Upper set of curves: Direct adaptation. Brightness 710 candles/m2 (4600 photons). Middle set of curves: Indirect adaptation. Glare angle 3°. Illumination at the eye 130 lux. Lower set of curves: Diascleral adaptation. Illumination at the eye 2900 lux.

Fig. 11
Fig. 11

Verification of the empirical relation 4, with the values taken from Fig. 10 (upper set of curves). Log (1−ρ)/τ is plotted against log (τ+t). The straight line is the theoretical line for ρs=0.09.

Fig. 12
Fig. 12

Resemblance between the adaptive and the electrical properties of the eye. Left. Upper curve. The dependence of the brightness perception of an object of 5 candles/m2 upon the time. Measured with the slit-wheel. Lower curve. The dependence of the sensitivity upon the time (see Fig. 7). Right. Analysis of the action potential of the dark-adapted eye of a cat. (After Granit.)

Tables (1)

Tables Icon

Table I Values of P in Eq. (3). The experiments are taken, over a period of 5 months.

Equations (5)

Equations on this page are rendered with MathJax. Learn more.

ρ = E 1 / E 2
V = ( E 2 - E 1 ) / E 1 = 1 / ρ - 1
V = P ( E / φ ) .
1 - ρ = ( 1 - ρ s ) ( τ / ( τ + t ) ) ,
d ( 1 - ρ ) d t = - ( 1 - ρ ) 2 ( 1 - ρ s ) τ ,