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function [fractionBleached] = ComputePhotopigmentBleaching(irradiance,receptortype,units,source,initialFraction,timeUnits)
% [fractionBleached] = ComputePhtopigmentBleaching(irradiance,[units],[source],[initialFraction],[timeUnits])
%
% Compute fraction of photopigment bleached, given irradiance of light
% reaching the eye.
%
% There are two distinct uses, controlled by the value of initialFraction.
%
% Usage 1 - If initialFraction is not passed or is empty, the steady state
% fraction of pigment bleached is returned for each irradiance in the
% passed input irradiance.
%
% Usage 2 - If initialFraction is passed as a scalar, this is taken as the
% time zero fraction bleached, and inputirradiance is taken to be the time
% variying irradiance, with fractionBleached the time varying fraction
% bleached.
%
% When time varying signals are handled, the unit of time is as specified
% by the timeunits argument (default, msec).
%
% As far as I can tell, the fundemantal measurements of the half-bleach
% constant for cones were made by Rushton and Henry (1968, Vision Reserch,
% 8, 617-631). This fact I learned from CVRL
% (http://www.cvrl.org/database/text/intros/introbleaches.htm).
%
% I am pretty sure that the Rushton and Henry measurements were made for
% 560 nm light, and they give (see their Figure 2) a half-bleach constant
% of 4.3 log10 trolands (20,000 td). This number is also given in Boynton
% and Kaiser, Human Color Vision, 2nd edition, pp 211 and following.
%
% It's probably fine to compute bleaching for L and M cones given retinal
% illuminance in trolands, given that these are effects that matter over
% log10 units. But trolands are not going to help much for the S-cones.
% According to CVRL there aren't good measurements for the half-bleaching
% constant for S cones because putting enough short-wavelength light onto
% the retina to bleach the S cones is not good for the eyes.
%
% None-the-less, it seems nice to have this routine written so that it will
% return a number if you give it irradiance either in trolands or in
% isomerizations/cone-sec. For 560 nm light and the CIE 10 deg
% fundamentals, I compute that 1 td is 137 isomerizations/cone-sec for L
% cones and 110 isomerizations/cone-sec for M cones. Take the weighted
% average value of (2*L + 1*M) = 128 and multiply by (10.^4.3) to get a
% half-bleach constant in isomerizations/cone-sec of 2.55e+06 (6.4 log10
% isomerizations/cone-sec). [Computations done 6/2/14 using
% IsomerizationsInEyeDemo and setting the fundamentals to 'CIE10deg' and
% wavelength to 560 nm by hand in the code. These are for the 'Boynton'
% source.]
%
% [ASIDE: I used 10 deg fundamentals to compute the bleaching constant
% expressed in terms of isomerizations, because I figure that Rushton's
% measurements are based on a fairly large field. Because the macular
% pigment absorbs a fair amount of light, this matters. If I compute
% instead with 2-deg fundamentals, I get that 1 td is 23.7 L cone
% isomerizations/cone-sec and 19.5 M cone isomerizations/cone-sec. These
% two numbers are ballpark consistent with Rodiek page 475 who gives 18.3
% and 15.9 for a monochromatic 540 THz light (555 nm)].
%
% This routine will do the computation either on the basis of input in
% trolands or input in isomerization/cone-sec, using the appropirate
% constant as above. Note that the computation of isomerizations takes
% into account lens and macular pigment, while the troland value is the
% straight troland value. A second advantage of using units of
% isomerizations/cone-sec is that you can compute this for other regions of
% the visual field and presumably the numbers will be about right. You can
% also compute for S-cones on the assumption that the half-bleach constant
% is the same for S-cones as for L- and M- cones.
%
% As far as I can tell, the computations and analysis of bleaching do not
% take into account changes in isomerization rate that occur because of
% change in spectral sensitivity of cones with bleaching. That is, the
% measurements are simply of steady state pigment density and are modeled
% with a formula that assumes monochromatic light (see treatment in
% Boynton).
%
% irradiance -- retinal irradiance specified as determined by units. If
% initialFraction is empty, this is a single number and
% steady state bleaching fraction is returned. If
% initialFraction is a number, then this is a time series
% of irradiance versus time, and fraction bleached for the
% same times is returned.
%
% receptortype
% 'cones' -- computations for cones. [Default]
%
% units -- units of irradiacne
% 'trolands' input irradiance in trolands. Note that the computation
% only makes sense for L and M cones if this is the input.
% This is photopic trolands if receptor type is 'cones'.
% [Default]
% 'isomerizations' nominal isomerization rate in
% isomerizations/cone-sec, comptued taking into account
% pre-retinal absorption as well as nominal cone axial
% density. But not taking into account any pigment
% bleaching.
%
% source -- source of underlying data
% 'Boynton' Boynton and Kaiser, Human Color Vision, 2nd edition,
% pp. 211 and following. [Default]
%
% initialFraction -- fraction of input bleached at time zero. If
% empty, steady state fraction bleached is
% returned. Default is empty.
%
% timeUnits -- units for time
% 'msec' millseconds [Default]
%
% 05/23/14 dhb Wrote it.
% 05/26/14 dhb Clean up.
% 06/02/14 dhb Take isomerizations number based on 2:1 L:M assumed ratio.
% 12/18/18 dhb Modify header comments for possibility of passing time
% varying signal. This breaks old usage that allowed
% computing steady state bleaching for a set of vector
% inputs, but I think that is OK.
%% Specify receptor type
if (nargin < 2 || isempty(receptortype))
receptortype = 'cones';
end
%% Specify units
if (nargin < 3 || isempty(units))
units = 'trolands';
end
%% Specify source
if (nargin < 4 || isempty(source))
source = 'Boynton';
end
%% Specify initial fraction
if (nargin < 5 || isempty(initialFraction))
initialFraction = [];
end
%% Time units, relevant if initialFraction is not empty
if (nargin < 6 || isempty(timeUnits))
timeUnits = 'msec';
end
%% Do it
switch (receptortype)
case 'cones'
switch (source)
case 'Boynton'
switch (units)
case 'trolands'
Izero = 10^4.3;
case 'isomerizations'
Izero = 10^6.4;
otherwise
error('Unkown input units specified');
end
% Steady state calculation
if (isempty(initialFraction))
fractionBleached = (irradiance./(irradiance + Izero));
% Time varying calculation.
else
% Take timeunits into account.
switch (timeunits)
case 'msec'
disp('Need to implement time varying calculation');
otherwise
error('Unknown time units specified');
end
end
otherwise
error('Unknown source specified');
end
otherwise
error('Unknown receptor type specified');
end
%% Test code.
%
% If you run the lines below you should get a plot that
% looks like Figure 6.3 (p. 212) in Boyton and Kaiser
% (red curve in plot with blue overlay) plus a shifted
% copy in blue. The isomerizations are not matched to the
% trolands numbers.
TEST = 0;
if (TEST)
trolands = logspace(0,7,1000);
isomerizations = 128*trolands;
fractionBleached = ComputePhotopigmentBleaching(trolands,'cones','trolands','Boynton');
fractionBleached1 = ComputePhotopigmentBleaching(isomerizations,'cones','isomerizations','Boynton');
figure; clf; hold on;
plot(log10(trolands),fractionBleached,'r','LineWidth',6);
plot(log10(isomerizations),fractionBleached1,'b','LineWidth',6);
plot(log10(trolands),fractionBleached1,'b','LineWidth',2);
end
end
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