Abstract

M-cone onsets appear dimmer than the background and elicit electroretinograms (ERGs) resembling the light offset response. We sought a corresponding anomalous pupillary light reflex (PLR) using a 4-primary ganzfeld as stimulator and pupillometer. Increments and decrements of white light were compared with M- and L-cone onsets and offsets using silent substitution. Luminance bias (LB) could be added to or subtracted from the cone-isolating stimuli. There was a normal PLR to L-cone increments, but the pupil constricted mainly to M-cone decrements. Changing LB produced a neutral point where on and off responses were balanced. The results reflect ERG and psychophysical studies. This observation may be linked to the antagonistic nature of the M-cone input to cone opponent mechanisms.

© 2018 Optical Society of America

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    [Crossref]
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    [Crossref]
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    [Crossref]
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    [Crossref]
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    [Crossref]
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    [Crossref]
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    [Crossref]
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    [Crossref]
  27. C. McMahon, J. Carroll, S. Awua, J. Neitz, and M. Neitz, “The L:M cone ratio in males of African descent with normal color vision,” J. Vis. 8(2), 5 (2008).
    [Crossref]
  28. H. Devries, “Luminosity curve of trichromats,” Nature 157, 736–737 (1946).
    [Crossref]
  29. C. R. Ingling, “Spectral sensitivity of cone opponent colour channels,” Vis. Res. 17, 1083–1089 (1977).
    [Crossref]
  30. M. V. Danilova, C. H. Chan, and J. D. Mollon, “Can spatial resolution reveal individual differences in the L:M cone ratio?” Vis. Res. 78, 26–38 (2013).
    [Crossref]
  31. J. M. Kraft and J. S. Werner, “Spectral efficiency across the life span: flicker photometry and brightness matching,” J. Opt. Soc. Am. A 11, 1213–1221 (1994).
    [Crossref]
  32. D. J. Calkins, S. J. Schein, Y. Tsukamoto, and P. Sterling, “M-cones and L-cones in macaque fovea connect to midget ganglion cells by different numbers of excitation synapses,” Nature 371, 70–72 (1994).
    [Crossref]
  33. R. C. Reid and R. M. Shapley, “Space and time maps of cone photoreceptor signals in macaque lateral geniculate nucleus,” J. Neurosci. 22, 6158–6175 (2002).
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    [Crossref]
  35. T. Woelders, T. Leenheers, M. C. M. Gordijn, R. A. Hut, D. G. M. Beersma, and E. J. Wams, “Melanopsin- and L-cone–induced pupil constriction is inhibited by S- and M-cones in humans,” Proc. Natl. Acad. Sci. USA 115, 792–797 (2018).
    [Crossref]

2018 (1)

T. Woelders, T. Leenheers, M. C. M. Gordijn, R. A. Hut, D. G. M. Beersma, and E. J. Wams, “Melanopsin- and L-cone–induced pupil constriction is inhibited by S- and M-cones in humans,” Proc. Natl. Acad. Sci. USA 115, 792–797 (2018).
[Crossref]

2016 (2)

J. Maguire, N. R. A. Parry, J. Kremers, D. Kommanapalli, I. J. Murray, and D. J. McKeefry, “Rod electroretinograms elicited by silent substitution stimuli from the light-adapted human eye,” Transl. Vis. Sci. Technol. 5, 13 (2016).
[Crossref]

N. R. A. Parry, D. J. McKeefry, J. Kremers, and I. J. Murray, “A dim view of M-cone onsets,” J. Opt. Soc. Am. A 33, A207–A213 (2016).
[Crossref]

2015 (1)

J. C. Park and J. J. McAnany, “Effect of stimulus size and luminance on the rod-, cone-, and melanopsin-mediated pupillary light reflex,” J. Vis. 15(3), 13 (2015).
[Crossref]

2014 (2)

M. Spitschan, S. Jain, D. H. Brainard, and G. K. Aguirre, “Opponent melanopsin and S-cone signals in the human pupillary light response,” Proc. Natl. Acad. Sci. USA 111, 15568–15572 (2014).
[Crossref]

D. McKeefry, J. Kremers, D. Kommanapalli, N. K. Challa, I. J. Murray, J. Maguire, and N. R. A. Parry, “Incremental and decremental L-and M-cone-driven ERG responses: I. Square-wave pulse stimulation,” J. Opt. Soc. Am. A 31, A159–A169 (2014).
[Crossref]

2013 (1)

M. V. Danilova, C. H. Chan, and J. D. Mollon, “Can spatial resolution reveal individual differences in the L:M cone ratio?” Vis. Res. 78, 26–38 (2013).
[Crossref]

2012 (2)

E. Petersen, A. Kijlstra, and M. Stanford, “Epidemiology of ocular toxoplasmosis,” Ocular Immunol. Inflamm. 20, 68–75 (2012).

A. B. Watson and J. I. Yellott, “A unified formula for light-adapted pupil size,” J. Vis. 12(10), 12 (2012).
[Crossref]

2010 (1)

B. B. Lee, P. R. Martin, and U. Grunert, “Retinal connectivity and primate vision,” Prog. Retin. Eye Res. 29, 622–639 (2010).
[Crossref]

2008 (3)

C. McMahon, J. Carroll, S. Awua, J. Neitz, and M. Neitz, “The L:M cone ratio in males of African descent with normal color vision,” J. Vis. 8(2), 5 (2008).
[Crossref]

A. D. Guler, J. L. Ecker, G. S. Lall, S. Haq, C. M. Altimus, H. W. Liao, A. R. Barnard, H. Cahill, T. C. Badea, H. Q. Zhao, M. W. Hankins, D. M. Berson, R. J. Lucas, K. W. Yau, and S. Hattar, “Melanopsin cells are the principal conduits for rod-cone input to non-image-forming vision,” Nature 453, 102–105 (2008).
[Crossref]

M. M. Bradley, L. Miccoli, M. A. Escrig, and P. J. Lang, “The pupil as a measure of emotional arousal and autonomic activation,” Psychophysiology 45, 602–607 (2008).
[Crossref]

2007 (1)

P. D. R. Gamlin, D. H. McDougal, J. Pokorny, V. C. Smith, K. W. Yau, and D. M. Dacey, “Human and macaque pupil responses driven by melanopsin-containing retinal ganglion cells,” Vis. Res. 47, 946–954 (2007).
[Crossref]

2005 (1)

D. M. Dacey, H. W. Liao, B. B. Peterson, F. R. Robinson, V. C. Smith, J. Pokorny, K. W. Yau, and P. D. Gamlin, “Melanopsin-expressing ganglion cells in primate retina signal colour and irradiance and project to the LGN,” Nature 433, 749–754 (2005).
[Crossref]

2004 (1)

J. L. Barbur, S. Moro, J. A. Harlow, B. L. Lam, and M. Liu, “Comparison of pupil responses to luminance and colour in severe optic neuritis,” Clin. Neurophysiol. 115, 2650–2658 (2004).
[Crossref]

2003 (1)

R. J. Lucas, S. Hattar, M. Takao, D. M. Berson, R. G. Foster, and K. W. Yau, “Diminished pupillary light reflex at high irradiances in melanopsin-knockout mice,” Science 299, 245–247 (2003).
[Crossref]

2002 (2)

R. C. Reid and R. M. Shapley, “Space and time maps of cone photoreceptor signals in macaque lateral geniculate nucleus,” J. Neurosci. 22, 6158–6175 (2002).

J. Carroll, J. Neitz, and M. Neitz, “Estimates of L:M cone ratio from ERG flicker photometry and genetics,” J. Vis. 2(8), 531–542 (2002).
[Crossref]

2000 (1)

A. Stockman and L. T. Sharpe, “The spectral sensitivities of the middle- and long-wavelength-sensitive cones derived from measurements in observers of known genotype,” Vis. Res. 40, 1711–1737 (2000).
[Crossref]

1999 (1)

L. Weiskrantz, A. Cowey, and J. L. Barbur, “Differential pupillary constriction and awareness in the absence of striate cortex,” Brain 122, 1533–1538 (1999).
[Crossref]

1998 (2)

J. L. Barbur, J. Wolf, and P. Lennie, “Visual processing levels revealed by response latencies to changes in different visual attributes,” Proc. R. Soc. B 265, 2321–2325 (1998).
[Crossref]

P. D. R. Gamlin, H. Y. Zhang, A. Harlow, and J. L. Barbur, “Pupil responses to stimulus color, structure and light flux increments in the rhesus monkey,” Vis. Res. 38, 3353–3358 (1998).
[Crossref]

1995 (1)

P. A. Stanley and A. K. Davies, “The effect of field of view size on steady-state pupil diameter,” Ophthalmic Physiol. Opt. 15, 601–603 (1995).
[Crossref]

1994 (3)

B. Winn, D. Whitaker, D. B. Elliott, and N. J. Phillips, “Factors affecting light-adapted pupil size in normal human subjects,” Invest. Ophthalmol. Visual Sci. 35, 1132–1137 (1994).

J. M. Kraft and J. S. Werner, “Spectral efficiency across the life span: flicker photometry and brightness matching,” J. Opt. Soc. Am. A 11, 1213–1221 (1994).
[Crossref]

D. J. Calkins, S. J. Schein, Y. Tsukamoto, and P. Sterling, “M-cones and L-cones in macaque fovea connect to midget ganglion cells by different numbers of excitation synapses,” Nature 371, 70–72 (1994).
[Crossref]

1988 (1)

P. K. Kaiser, “Sensation luminance: a new name to distinguish CIE luminance from luminance dependent on an individual’s spectral sensitivity,” Vis. Res. 28, 455–456 (1988).
[Crossref]

1977 (2)

C. R. Ingling, “Spectral sensitivity of cone opponent colour channels,” Vis. Res. 17, 1083–1089 (1977).
[Crossref]

L. A. Benevento, M. Rezak, and R. Santosanderson, “Autoradiographic study of projections of pretectum in rhesus monkey (macaca mulata): evidence for sensorimotor links to thalamus and oculomotor nuclei,” Brain Res. 127, 197–218 (1977).
[Crossref]

1962 (1)

H. Bouma, “Size of the static pupil as a function of wavelength and luminosity of the light incident on the human eye,” Nature 193, 690–691 (1962).
[Crossref]

1957 (1)

F. W. Campbell, “Refractive errors of the human eye,” Nature 179, 705 (1957).
[Crossref]

1946 (1)

H. Devries, “Luminosity curve of trichromats,” Nature 157, 736–737 (1946).
[Crossref]

1936 (1)

B. H. Crawford, “The dependence of pupil size upon external light stimulus under static and variable conditions,” Proc. R. Soc. Ser. B 121, 376–395 (1936).
[Crossref]

Aguirre, G. K.

M. Spitschan, S. Jain, D. H. Brainard, and G. K. Aguirre, “Opponent melanopsin and S-cone signals in the human pupillary light response,” Proc. Natl. Acad. Sci. USA 111, 15568–15572 (2014).
[Crossref]

Altimus, C. M.

A. D. Guler, J. L. Ecker, G. S. Lall, S. Haq, C. M. Altimus, H. W. Liao, A. R. Barnard, H. Cahill, T. C. Badea, H. Q. Zhao, M. W. Hankins, D. M. Berson, R. J. Lucas, K. W. Yau, and S. Hattar, “Melanopsin cells are the principal conduits for rod-cone input to non-image-forming vision,” Nature 453, 102–105 (2008).
[Crossref]

Awua, S.

C. McMahon, J. Carroll, S. Awua, J. Neitz, and M. Neitz, “The L:M cone ratio in males of African descent with normal color vision,” J. Vis. 8(2), 5 (2008).
[Crossref]

Badea, T. C.

A. D. Guler, J. L. Ecker, G. S. Lall, S. Haq, C. M. Altimus, H. W. Liao, A. R. Barnard, H. Cahill, T. C. Badea, H. Q. Zhao, M. W. Hankins, D. M. Berson, R. J. Lucas, K. W. Yau, and S. Hattar, “Melanopsin cells are the principal conduits for rod-cone input to non-image-forming vision,” Nature 453, 102–105 (2008).
[Crossref]

Barbur, J. L.

J. L. Barbur, S. Moro, J. A. Harlow, B. L. Lam, and M. Liu, “Comparison of pupil responses to luminance and colour in severe optic neuritis,” Clin. Neurophysiol. 115, 2650–2658 (2004).
[Crossref]

L. Weiskrantz, A. Cowey, and J. L. Barbur, “Differential pupillary constriction and awareness in the absence of striate cortex,” Brain 122, 1533–1538 (1999).
[Crossref]

J. L. Barbur, J. Wolf, and P. Lennie, “Visual processing levels revealed by response latencies to changes in different visual attributes,” Proc. R. Soc. B 265, 2321–2325 (1998).
[Crossref]

P. D. R. Gamlin, H. Y. Zhang, A. Harlow, and J. L. Barbur, “Pupil responses to stimulus color, structure and light flux increments in the rhesus monkey,” Vis. Res. 38, 3353–3358 (1998).
[Crossref]

Barnard, A. R.

A. D. Guler, J. L. Ecker, G. S. Lall, S. Haq, C. M. Altimus, H. W. Liao, A. R. Barnard, H. Cahill, T. C. Badea, H. Q. Zhao, M. W. Hankins, D. M. Berson, R. J. Lucas, K. W. Yau, and S. Hattar, “Melanopsin cells are the principal conduits for rod-cone input to non-image-forming vision,” Nature 453, 102–105 (2008).
[Crossref]

Beersma, D. G. M.

T. Woelders, T. Leenheers, M. C. M. Gordijn, R. A. Hut, D. G. M. Beersma, and E. J. Wams, “Melanopsin- and L-cone–induced pupil constriction is inhibited by S- and M-cones in humans,” Proc. Natl. Acad. Sci. USA 115, 792–797 (2018).
[Crossref]

Benevento, L. A.

L. A. Benevento, M. Rezak, and R. Santosanderson, “Autoradiographic study of projections of pretectum in rhesus monkey (macaca mulata): evidence for sensorimotor links to thalamus and oculomotor nuclei,” Brain Res. 127, 197–218 (1977).
[Crossref]

Berson, D. M.

A. D. Guler, J. L. Ecker, G. S. Lall, S. Haq, C. M. Altimus, H. W. Liao, A. R. Barnard, H. Cahill, T. C. Badea, H. Q. Zhao, M. W. Hankins, D. M. Berson, R. J. Lucas, K. W. Yau, and S. Hattar, “Melanopsin cells are the principal conduits for rod-cone input to non-image-forming vision,” Nature 453, 102–105 (2008).
[Crossref]

R. J. Lucas, S. Hattar, M. Takao, D. M. Berson, R. G. Foster, and K. W. Yau, “Diminished pupillary light reflex at high irradiances in melanopsin-knockout mice,” Science 299, 245–247 (2003).
[Crossref]

Bouma, H.

H. Bouma, “Size of the static pupil as a function of wavelength and luminosity of the light incident on the human eye,” Nature 193, 690–691 (1962).
[Crossref]

Bradley, M. M.

M. M. Bradley, L. Miccoli, M. A. Escrig, and P. J. Lang, “The pupil as a measure of emotional arousal and autonomic activation,” Psychophysiology 45, 602–607 (2008).
[Crossref]

Brainard, D. H.

M. Spitschan, S. Jain, D. H. Brainard, and G. K. Aguirre, “Opponent melanopsin and S-cone signals in the human pupillary light response,” Proc. Natl. Acad. Sci. USA 111, 15568–15572 (2014).
[Crossref]

Cahill, H.

A. D. Guler, J. L. Ecker, G. S. Lall, S. Haq, C. M. Altimus, H. W. Liao, A. R. Barnard, H. Cahill, T. C. Badea, H. Q. Zhao, M. W. Hankins, D. M. Berson, R. J. Lucas, K. W. Yau, and S. Hattar, “Melanopsin cells are the principal conduits for rod-cone input to non-image-forming vision,” Nature 453, 102–105 (2008).
[Crossref]

Calkins, D. J.

D. J. Calkins, S. J. Schein, Y. Tsukamoto, and P. Sterling, “M-cones and L-cones in macaque fovea connect to midget ganglion cells by different numbers of excitation synapses,” Nature 371, 70–72 (1994).
[Crossref]

Campbell, F. W.

F. W. Campbell, “Refractive errors of the human eye,” Nature 179, 705 (1957).
[Crossref]

Carroll, J.

C. McMahon, J. Carroll, S. Awua, J. Neitz, and M. Neitz, “The L:M cone ratio in males of African descent with normal color vision,” J. Vis. 8(2), 5 (2008).
[Crossref]

J. Carroll, J. Neitz, and M. Neitz, “Estimates of L:M cone ratio from ERG flicker photometry and genetics,” J. Vis. 2(8), 531–542 (2002).
[Crossref]

Challa, N. K.

Chan, C. H.

M. V. Danilova, C. H. Chan, and J. D. Mollon, “Can spatial resolution reveal individual differences in the L:M cone ratio?” Vis. Res. 78, 26–38 (2013).
[Crossref]

Cowey, A.

L. Weiskrantz, A. Cowey, and J. L. Barbur, “Differential pupillary constriction and awareness in the absence of striate cortex,” Brain 122, 1533–1538 (1999).
[Crossref]

Crawford, B. H.

B. H. Crawford, “The dependence of pupil size upon external light stimulus under static and variable conditions,” Proc. R. Soc. Ser. B 121, 376–395 (1936).
[Crossref]

Dacey, D. M.

P. D. R. Gamlin, D. H. McDougal, J. Pokorny, V. C. Smith, K. W. Yau, and D. M. Dacey, “Human and macaque pupil responses driven by melanopsin-containing retinal ganglion cells,” Vis. Res. 47, 946–954 (2007).
[Crossref]

D. M. Dacey, H. W. Liao, B. B. Peterson, F. R. Robinson, V. C. Smith, J. Pokorny, K. W. Yau, and P. D. Gamlin, “Melanopsin-expressing ganglion cells in primate retina signal colour and irradiance and project to the LGN,” Nature 433, 749–754 (2005).
[Crossref]

Danilova, M. V.

M. V. Danilova, C. H. Chan, and J. D. Mollon, “Can spatial resolution reveal individual differences in the L:M cone ratio?” Vis. Res. 78, 26–38 (2013).
[Crossref]

Davies, A. K.

P. A. Stanley and A. K. Davies, “The effect of field of view size on steady-state pupil diameter,” Ophthalmic Physiol. Opt. 15, 601–603 (1995).
[Crossref]

Devries, H.

H. Devries, “Luminosity curve of trichromats,” Nature 157, 736–737 (1946).
[Crossref]

Ecker, J. L.

A. D. Guler, J. L. Ecker, G. S. Lall, S. Haq, C. M. Altimus, H. W. Liao, A. R. Barnard, H. Cahill, T. C. Badea, H. Q. Zhao, M. W. Hankins, D. M. Berson, R. J. Lucas, K. W. Yau, and S. Hattar, “Melanopsin cells are the principal conduits for rod-cone input to non-image-forming vision,” Nature 453, 102–105 (2008).
[Crossref]

Elliott, D. B.

B. Winn, D. Whitaker, D. B. Elliott, and N. J. Phillips, “Factors affecting light-adapted pupil size in normal human subjects,” Invest. Ophthalmol. Visual Sci. 35, 1132–1137 (1994).

Escrig, M. A.

M. M. Bradley, L. Miccoli, M. A. Escrig, and P. J. Lang, “The pupil as a measure of emotional arousal and autonomic activation,” Psychophysiology 45, 602–607 (2008).
[Crossref]

Foster, R. G.

R. J. Lucas, S. Hattar, M. Takao, D. M. Berson, R. G. Foster, and K. W. Yau, “Diminished pupillary light reflex at high irradiances in melanopsin-knockout mice,” Science 299, 245–247 (2003).
[Crossref]

Gamlin, P. D.

D. M. Dacey, H. W. Liao, B. B. Peterson, F. R. Robinson, V. C. Smith, J. Pokorny, K. W. Yau, and P. D. Gamlin, “Melanopsin-expressing ganglion cells in primate retina signal colour and irradiance and project to the LGN,” Nature 433, 749–754 (2005).
[Crossref]

Gamlin, P. D. R.

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J. L. Barbur, S. Moro, J. A. Harlow, B. L. Lam, and M. Liu, “Comparison of pupil responses to luminance and colour in severe optic neuritis,” Clin. Neurophysiol. 115, 2650–2658 (2004).
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R. J. Lucas, S. Hattar, M. Takao, D. M. Berson, R. G. Foster, and K. W. Yau, “Diminished pupillary light reflex at high irradiances in melanopsin-knockout mice,” Science 299, 245–247 (2003).
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J. Maguire, N. R. A. Parry, J. Kremers, D. Kommanapalli, I. J. Murray, and D. J. McKeefry, “Rod electroretinograms elicited by silent substitution stimuli from the light-adapted human eye,” Transl. Vis. Sci. Technol. 5, 13 (2016).
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D. McKeefry, J. Kremers, D. Kommanapalli, N. K. Challa, I. J. Murray, J. Maguire, and N. R. A. Parry, “Incremental and decremental L-and M-cone-driven ERG responses: I. Square-wave pulse stimulation,” J. Opt. Soc. Am. A 31, A159–A169 (2014).
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B. B. Lee, P. R. Martin, and U. Grunert, “Retinal connectivity and primate vision,” Prog. Retin. Eye Res. 29, 622–639 (2010).
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J. C. Park and J. J. McAnany, “Effect of stimulus size and luminance on the rod-, cone-, and melanopsin-mediated pupillary light reflex,” J. Vis. 15(3), 13 (2015).
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P. D. R. Gamlin, D. H. McDougal, J. Pokorny, V. C. Smith, K. W. Yau, and D. M. Dacey, “Human and macaque pupil responses driven by melanopsin-containing retinal ganglion cells,” Vis. Res. 47, 946–954 (2007).
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McKeefry, D. J.

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Neitz, J.

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J. C. Park and J. J. McAnany, “Effect of stimulus size and luminance on the rod-, cone-, and melanopsin-mediated pupillary light reflex,” J. Vis. 15(3), 13 (2015).
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D. M. Dacey, H. W. Liao, B. B. Peterson, F. R. Robinson, V. C. Smith, J. Pokorny, K. W. Yau, and P. D. Gamlin, “Melanopsin-expressing ganglion cells in primate retina signal colour and irradiance and project to the LGN,” Nature 433, 749–754 (2005).
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P. D. R. Gamlin, D. H. McDougal, J. Pokorny, V. C. Smith, K. W. Yau, and D. M. Dacey, “Human and macaque pupil responses driven by melanopsin-containing retinal ganglion cells,” Vis. Res. 47, 946–954 (2007).
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D. M. Dacey, H. W. Liao, B. B. Peterson, F. R. Robinson, V. C. Smith, J. Pokorny, K. W. Yau, and P. D. Gamlin, “Melanopsin-expressing ganglion cells in primate retina signal colour and irradiance and project to the LGN,” Nature 433, 749–754 (2005).
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D. M. Dacey, H. W. Liao, B. B. Peterson, F. R. Robinson, V. C. Smith, J. Pokorny, K. W. Yau, and P. D. Gamlin, “Melanopsin-expressing ganglion cells in primate retina signal colour and irradiance and project to the LGN,” Nature 433, 749–754 (2005).
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L. A. Benevento, M. Rezak, and R. Santosanderson, “Autoradiographic study of projections of pretectum in rhesus monkey (macaca mulata): evidence for sensorimotor links to thalamus and oculomotor nuclei,” Brain Res. 127, 197–218 (1977).
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D. M. Dacey, H. W. Liao, B. B. Peterson, F. R. Robinson, V. C. Smith, J. Pokorny, K. W. Yau, and P. D. Gamlin, “Melanopsin-expressing ganglion cells in primate retina signal colour and irradiance and project to the LGN,” Nature 433, 749–754 (2005).
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M. Spitschan, S. Jain, D. H. Brainard, and G. K. Aguirre, “Opponent melanopsin and S-cone signals in the human pupillary light response,” Proc. Natl. Acad. Sci. USA 111, 15568–15572 (2014).
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E. Petersen, A. Kijlstra, and M. Stanford, “Epidemiology of ocular toxoplasmosis,” Ocular Immunol. Inflamm. 20, 68–75 (2012).

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R. J. Lucas, S. Hattar, M. Takao, D. M. Berson, R. G. Foster, and K. W. Yau, “Diminished pupillary light reflex at high irradiances in melanopsin-knockout mice,” Science 299, 245–247 (2003).
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D. J. Calkins, S. J. Schein, Y. Tsukamoto, and P. Sterling, “M-cones and L-cones in macaque fovea connect to midget ganglion cells by different numbers of excitation synapses,” Nature 371, 70–72 (1994).
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B. Winn, D. Whitaker, D. B. Elliott, and N. J. Phillips, “Factors affecting light-adapted pupil size in normal human subjects,” Invest. Ophthalmol. Visual Sci. 35, 1132–1137 (1994).

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T. Woelders, T. Leenheers, M. C. M. Gordijn, R. A. Hut, D. G. M. Beersma, and E. J. Wams, “Melanopsin- and L-cone–induced pupil constriction is inhibited by S- and M-cones in humans,” Proc. Natl. Acad. Sci. USA 115, 792–797 (2018).
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Brain (1)

L. Weiskrantz, A. Cowey, and J. L. Barbur, “Differential pupillary constriction and awareness in the absence of striate cortex,” Brain 122, 1533–1538 (1999).
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Brain Res. (1)

L. A. Benevento, M. Rezak, and R. Santosanderson, “Autoradiographic study of projections of pretectum in rhesus monkey (macaca mulata): evidence for sensorimotor links to thalamus and oculomotor nuclei,” Brain Res. 127, 197–218 (1977).
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Clin. Neurophysiol. (1)

J. L. Barbur, S. Moro, J. A. Harlow, B. L. Lam, and M. Liu, “Comparison of pupil responses to luminance and colour in severe optic neuritis,” Clin. Neurophysiol. 115, 2650–2658 (2004).
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R. C. Reid and R. M. Shapley, “Space and time maps of cone photoreceptor signals in macaque lateral geniculate nucleus,” J. Neurosci. 22, 6158–6175 (2002).

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D. M. Dacey, H. W. Liao, B. B. Peterson, F. R. Robinson, V. C. Smith, J. Pokorny, K. W. Yau, and P. D. Gamlin, “Melanopsin-expressing ganglion cells in primate retina signal colour and irradiance and project to the LGN,” Nature 433, 749–754 (2005).
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Ocular Immunol. Inflamm. (1)

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Figures (3)

Fig. 1.
Fig. 1. (a) Bold line shows the PLR to a 1000 ms 18.87    cd · m 2 stimulus with L, M, S, and rod contrast of 0.11 ( background = 17    cd · m 2 ). The same stimulus was also presented on a series of lower luminance backgrounds (resulting in increased Weber contrast), as depicted by the fine lines. Subject IJM. (b) Expanded view of the C = 0.11 run. Average of 30 repeats with 95% confidence intervals.
Fig. 2.
Fig. 2. Top group shows averaged PLRs to white, L-cone, and M-cone modulated stimuli with duration 1000 ms and ISI 2000 ms, as depicted by the timing bars above. (a)–(c) show different subjects. For each subject the left group shows increment responses and the right group decrements. The lower groups provide a direct comparison between L on and M off responses (left column for each subject) and M on and L off responses (right column).
Fig. 3.
Fig. 3. (a) PLR to cone-isolating increments of 1000 ms with a 2000 ms offset. Subject NRAP. Left column shows responses to L-isolating stimuli and right column to M-isolating stimuli. Each trace is to a different LB, with LB = 1 represented in bold. Above this there was an additional luminance increment and, below it, a luminance decrement. A second bold line shows the LB where there was a balance point between the pupil responding to the onset and to the offset. (b) and (c) show similar data for subjects ER and IJM.

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