%PDF-1.6
%
1 0 obj
<>
endobj
2 0 obj
<>stream
Antal Martinecz, Mihoko Niitsuma
Fractional integral-like processing in retinal cones reduces noise and improves adaptation
www.plosone.org
www.plosone.org
endstream
endobj
3 0 obj
<>/XObject<>>>/CropBox[0 0 612 792]/MediaBox[0 0 612 792]/Parent 16 0 R/Annots 17 0 R/Contents 18 0 R/TrimBox[0 0 612 792]>>
endobj
17 0 obj
[19 0 R 20 0 R 21 0 R 22 0 R 23 0 R 24 0 R 25 0 R 26 0 R 27 0 R 28 0 R 29 0 R 30 0 R 31 0 R 32 0 R 33 0 R 34 0 R 35 0 R 36 0 R 37 0 R 38 0 R 39 0 R 40 0 R 41 0 R 42 0 R 43 0 R 44 0 R]
endobj
19 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref014)>>
endobj
20 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref015)>>
endobj
21 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref017)>>
endobj
22 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref018)>>
endobj
23 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref019)>>
endobj
24 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref020)>>
endobj
25 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref021)>>
endobj
26 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref022)>>
endobj
27 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref023)>>
endobj
28 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref025)>>
endobj
29 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref015)>>
endobj
30 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref026)>>
endobj
31 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref029)>>
endobj
32 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref015)>>
endobj
33 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref015)>>
endobj
34 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref017)>>
endobj
35 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.e001)>>
endobj
36 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.e004)>>
endobj
37 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.e005)>>
endobj
38 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref010)>>
endobj
39 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref030)>>
endobj
40 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref033)>>
endobj
41 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref015)>>
endobj
42 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref030)>>
endobj
43 0 obj
<>/Border[0 0 0]/Dest(Rpone.0205099.ref032)>>
endobj
44 0 obj
<>/Border[0 0 0]/A 45 0 R>>
endobj
45 0 obj
<>
endobj
18 0 obj
[46 0 R 47 0 R 48 0 R 49 0 R 50 0 R 51 0 R 52 0 R 53 0 R 54 0 R 55 0 R 56 0 R 57 0 R 58 0 R 59 0 R 60 0 R 61 0 R 62 0 R 63 0 R 64 0 R 65 0 R 66 0 R 67 0 R 68 0 R 69 0 R]
endobj
46 0 obj
<>stream
q
0.83 0.64 0.02 0 k
203.4142 628.3276 m
212.9386 628.3276 l
h
f*
477.7512 615.3449 m
487.2756 615.3449 l
h
f*
492.4346 615.3449 m
501.9591 615.3449 l
h
f*
554.1165 576.3402 m
563.6409 576.3402 l
h
f*
340.9512 563.3008 m
350.4756 563.3008 l
h
f*
541.5874 563.3008 m
551.0551 563.3008 l
h
f*
346.6205 550.3181 m
356.0882 550.3181 l
h
f*
499.4646 550.3181 m
508.989 550.3181 l
h
f*
351.4394 524.2961 m
360.9638 524.2961 l
h
f*
366.1228 524.2961 m
375.6472 524.2961 l
h
f*
439.0299 485.348 m
448.4976 485.348 l
h
f*
399.515 247.1811 m
409.0394 247.1811 l
h
f*
414.1984 247.1811 m
423.7228 247.1811 l
h
f*
271.5024 221.1591 m
281.0268 221.1591 l
h
f*
431.8866 195.1937 m
441.3543 195.1937 l
h
f*
446.5701 195.1937 m
456.0945 195.1937 l
h
f*
295.937 182.1543 m
305.0646 182.1543 l
h
f*
310.2236 182.1543 m
320.2583 182.1543 l
h
f*
489.9402 169.1717 m
511.8803 169.1717 l
h
f*
520.8945 169.1717 m
530.3622 169.1717 l
h
f*
534.8409 169.1717 m
544.3654 169.1717 l
h
f*
549.5244 169.1717 m
559.0488 169.1717 l
h
f*
438.5197 156.189 m
448.0441 156.189 l
h
f*
358.3559 143.1496 m
367.8803 143.1496 l
h
f*
373.0394 143.1496 m
382.5638 143.1496 l
h
f*
0 g
1 j
1 J
0 w
10 0 0 10 200.0125 707.4141 cm
BT
/F7 1 Tf
1 TL
-0.004 Tc
0 0 Td
(activity;)Tj
3.3391 0 Td
(however,)Tj
3.8381 0 Td
(the)Tj
1.474 0 Td
(time)Tj
2.024 0 Td
(required)Tj
3.6396 0 Td
(to)Tj
1.0262 0 Td
(complete)Tj
3.8777 0 Td
(each)Tj
2.0296 0 Td
(step)Tj
1.8199 0 Td
(progressively)Tj
5.4424 0 Td
(increases,)Tj
4.0592 0 Td
(resulting)Tj
3.7304 0 Td
(in)Tj
-36.3002 -1.3039 Td
(the)Tj
1.4683 0 Td
(temporary)Tj
4.4163 0 Td
(accumulation)Tj
5.6806 0 Td
(of)Tj
1.0148 0 Td
(partially)Tj
3.4866 0 Td
(deactivated)Tj
4.7338 0 Td
(rhodopsin)Tj
4.354 0 Td
(molecules)Tj
4.1953 0 Td
(that)Tj
1.7801 0 Td
(still)Tj
1.6384 0 Td
(show)Tj
2.3018 0 Td
(some)Tj
-35.07 -1.2982 Td
(residual)Tj
3.3788 0 Td
(activity.)Tj
3.3392 0 Td
(It)Tj
0.8617 0 Td
(is)Tj
0.8448 0 Td
(currently)Tj
3.8891 0 Td
(unclear)Tj
3.1918 0 Td
(whether)Tj
3.4809 0 Td
(\(and)Tj
2.0693 0 Td
(how\))Tj
2.279 0 Td
(these)Tj
2.2507 0 Td
(residual)Tj
3.3789 0 Td
(activities)Tj
3.7133 0 Td
(affect)Tj
2.3755 0 Td
(the)Tj
-35.053 -1.2983 Td
(signal)Tj
2.5398 0 Td
(produced)Tj
4.0478 0 Td
(by)Tj
1.1792 0 Td
(the)Tj
1.4684 0 Td
(cell.)Tj
-8.039 -1.2983 Td
(During)Tj
3.1407 0 Td
(a)Tj
0.652 0 Td
(previous)Tj
3.651 0 Td
(conference,)Tj
4.8245 0 Td
(we)Tj
1.3266 0 Td
(reported)Tj
3.6624 0 Td
(\(as)Tj
1.3549 0 Td
(preliminary)Tj
4.938 0 Td
(results\))Tj
3.1521 0 Td
(that)Tj
1.7801 0 Td
(the)Tj
1.4683 0 Td
(structure)Tj
3.8211 0 Td
(of)Tj
1.0148 0 Td
(this)Tj
-35.9827 -1.3039 Td
(process)Tj
3.1804 0 Td
(has)Tj
1.5477 0 Td
(the)Tj
1.4683 0 Td
(potential)Tj
3.7588 0 Td
(to)Tj
1.0261 0 Td
(approximate)Tj
5.2724 0 Td
(the)Tj
1.474 0 Td
(mathematical)Tj
5.6409 0 Td
(operations)Tj
4.4561 0 Td
(of)Tj
1.0148 0 Td
(fractional)Tj
4.0365 0 Td
(integration.)Tj
-32.876 -1.2982 Td
([)Tj
0.83 0.64 0.02 0 k
(14)Tj
0 g
(].)Tj
2.0749 0 Td
(Furthermore,)Tj
5.6126 0 Td
(we)Tj
1.3322 0 Td
(have)Tj
2.058 0 Td
(also)Tj
1.7688 0 Td
(shown)Tj
2.8459 0 Td
(that)Tj
1.7802 0 Td
(the)Tj
1.4683 0 Td
(phosphorylation)Tj
6.8145 0 Td
(process)Tj
3.1804 0 Td
(can)Tj
1.6271 0 Td
(approximate)Tj
5.2781 0 Td
(this)Tj
-35.841 -1.2983 Td
(kind)Tj
2.0579 0 Td
(of)Tj
1.0148 0 Td
(behaviour,)Tj
4.4674 0 Td
(based)Tj
2.4718 0 Td
(on)Tj
1.2755 0 Td
(the)Tj
1.4684 0 Td
(commonly)Tj
4.5467 0 Td
(used)Tj
2.058 0 Td
(models)Tj
3.1067 0 Td
(of)Tj
1.0148 0 Td
(the)Tj
1.4683 0 Td
(cones)Tj
2.4832 0 Td
([)Tj
0.83 0.64 0.02 0 k
(15)Tj
0 g
()Tj
0.83 0.64 0.02 0 k
(17)Tj
0 g
(].)Tj
-26.2373 -1.3039 Td
(Fractional)Tj
4.2803 0 Td
(integrals)Tj
3.6339 0 Td
(generalise)Tj
4.184 0 Td
(traditional)Tj
4.422 0 Td
(Riemann)Tj
3.8834 0 Td
(integrals)Tj
3.634 0 Td
(by)Tj
1.1792 0 Td
(allowing)Tj
3.6227 0 Td
(integration)Tj
4.6488 0 Td
(of)Tj
-34.6845 -1.2983 Td
(non-integer)Tj
4.9719 0 Td
(times)Tj
2.3867 0 Td
(\(e.g.)Tj
1.8992 0 Td
(half-integrals\).)Tj
6.0491 0 Td
(Fractional)Tj
4.2803 0 Td
(calculus,)Tj
3.634 0 Td
(which)Tj
2.6476 0 Td
(also)Tj
1.7688 0 Td
(encompasses)Tj
5.4028 0 Td
(fractional)Tj
-33.0404 -1.2983 Td
(integrals,)Tj
3.8607 0 Td
(has)Tj
1.5421 0 Td
(many)Tj
2.4774 0 Td
(interesting)Tj
4.4901 0 Td
(real-world)Tj
4.3823 0 Td
(applications)Tj
5.04 0 Td
(in)Tj
1.0374 0 Td
(various)Tj
3.1408 0 Td
(fields,)Tj
2.5625 0 Td
(such)Tj
2.058 0 Td
(as)Tj
1.0148 0 Td
(robotics)Tj
3.4639 0 Td
([)Tj
0.83 0.64 0.02 0 k
(18)Tj
0 g
(],)Tj
-35.07 -1.3039 Td
(modelling)Tj
4.2689 0 Td
(ground)Tj
3.1691 0 Td
(water)Tj
2.4321 0 Td
(pollution)Tj
3.8835 0 Td
([)Tj
0.83 0.64 0.02 0 k
(19)Tj
0 g
(],)Tj
2.0749 0 Td
(modelling)Tj
4.2746 0 Td
(drug)Tj
2.109 0 Td
(diffusion)Tj
3.8041 0 Td
(in)Tj
1.0374 0 Td
(the)Tj
1.4684 0 Td
(human)Tj
3.0784 0 Td
(body)Tj
2.2167 0 Td
([)Tj
0.83 0.64 0.02 0 k
(20)Tj
0 g
(],)Tj
-33.8171 -1.2982 Td
(modelling)Tj
4.2689 0 Td
(the)Tj
1.4683 0 Td
(dynamics)Tj
4.0592 0 Td
(of)Tj
1.0148 0 Td
(neurons)Tj
3.5093 0 Td
([)Tj
0.83 0.64 0.02 0 k
(21)Tj
0 g
(],)Tj
2.0749 0 Td
(and)Tj
1.7292 0 Td
(modelling)Tj
4.2746 0 Td
(protein)Tj
3.1521 0 Td
(dynamics)Tj
4.0535 0 Td
([)Tj
0.83 0.64 0.02 0 k
(22)Tj
0 g
(].)Tj
2.075 0 Td
(Moreover,)Tj
-31.6798 -1.2983 Td
(fractional)Tj
4.0365 0 Td
(calculus)Tj
3.4072 0 Td
(has)Tj
1.5477 0 Td
(been)Tj
2.1203 0 Td
(gaining)Tj
3.2145 0 Td
(traction)Tj
3.3675 0 Td
(in,)Tj
1.2642 0 Td
(and)Tj
1.7292 0 Td
(proved)Tj
3.0217 0 Td
(to)Tj
1.0261 0 Td
(be)Tj
1.1509 0 Td
(a)Tj
0.6519 0 Td
(useful)Tj
2.5966 0 Td
(tool)Tj
1.7744 0 Td
(for,)Tj
1.6158 0 Td
(our)Tj
1.6214 0 Td
(topic)Tj
2.2337 0 Td
(of)Tj
-36.3796 -1.3039 Td
(interest:)Tj
3.4355 0 Td
(image)Tj
2.6306 0 Td
(and)Tj
1.7291 0 Td
(signal)Tj
2.5398 0 Td
(processing)Tj
4.4674 0 Td
([)Tj
0.83 0.64 0.02 0 k
(23)Tj
0 g
()Tj
0.83 0.64 0.02 0 k
(25)Tj
0 g
(].)Tj
-13.6062 -1.2983 Td
(In)Tj
1.1111 0 Td
(this)Tj
1.6725 0 Td
(paper,)Tj
2.7099 0 Td
(we)Tj
1.3266 0 Td
(investigate)Tj
4.4617 0 Td
(whether)Tj
3.4752 0 Td
(the)Tj
1.474 0 Td
(multi-stage)Tj
4.7112 0 Td
(deactivation)Tj
5.1023 0 Td
(process)Tj
3.1861 0 Td
(of)Tj
1.0148 0 Td
(rhodopsin)Tj
4.354 0 Td
(and)Tj
-35.7956 -1.2982 Td
(related)Tj
2.9423 0 Td
(residual)Tj
3.3789 0 Td
(activities)Tj
3.7133 0 Td
(offer)Tj
2.0977 0 Td
(any)Tj
1.661 0 Td
(signal)Tj
2.5399 0 Td
(processing)Tj
4.4673 0 Td
(benefits,)Tj
3.5603 0 Td
(and)Tj
1.7292 0 Td
(how)Tj
1.9388 0 Td
(it)Tj
0.7881 0 Td
(affects)Tj
2.7326 0 Td
(signals)Tj
2.9026 0 Td
(in)Tj
1.0375 0 Td
(gen-)Tj
-35.4895 -1.2983 Td
(eral.)Tj
1.9218 0 Td
(In)Tj
1.1055 0 Td
(our)Tj
1.6271 0 Td
(previous)Tj
3.651 0 Td
(work,)Tj
2.5002 0 Td
(we)Tj
1.3322 0 Td
(used)Tj
2.058 0 Td
(the)Tj
1.4683 0 Td
(model)Tj
2.7439 0 Td
(presented)Tj
4.1159 0 Td
(in)Tj
1.0375 0 Td
([)Tj
0.83 0.64 0.02 0 k
(15)Tj
0 g
(])Tj
1.8482 0 Td
(to)Tj
1.0261 0 Td
(show)Tj
2.3017 0 Td
(that)Tj
1.7745 0 Td
(this)Tj
1.6781 0 Td
(process)Tj
3.1804 0 Td
(has)Tj
-35.3704 -1.3039 Td
(the)Tj
1.4683 0 Td
(potential)Tj
3.7587 0 Td
(to)Tj
1.0261 0 Td
(approximate)Tj
5.2781 0 Td
(fractional)Tj
4.0365 0 Td
(integral-like)Tj
5.057 0 Td
(behaviour.)Tj
4.4674 0 Td
(To)Tj
1.3379 0 Td
(investigate)Tj
4.4617 0 Td
(the)Tj
1.4684 0 Td
(effect)Tj
2.364 0 Td
(of)Tj
1.0148 0 Td
(the)Tj
-35.7389 -1.2983 Td
(deactivation)Tj
5.1023 0 Td
(process,)Tj
3.4072 0 Td
(we)Tj
1.3323 0 Td
(have)Tj
2.0579 0 Td
(expanded)Tj
4.0876 0 Td
(this)Tj
1.6724 0 Td
(model)Tj
2.7439 0 Td
(with)Tj
2.0012 0 Td
(the)Tj
1.4684 0 Td
(activity)Tj
3.1181 0 Td
(of)Tj
1.0148 0 Td
(arrestin)Tj
3.2938 0 Td
(bound)Tj
2.8403 0 Td
(rhodop-)Tj
-34.1402 -1.2983 Td
(sin,)Tj
1.6214 0 Td
(as)Tj
1.0148 0 Td
(it)Tj
0.7823 0 Td
(was)Tj
1.7008 0 Td
(not)Tj
1.576 0 Td
(previously)Tj
4.3484 0 Td
(included.)Tj
-9.8475 -1.3039 Td
(In)Tj
1.1111 0 Td
(addition,)Tj
3.8211 0 Td
(we)Tj
1.3266 0 Td
(show)Tj
2.3017 0 Td
(that)Tj
1.7802 0 Td
(the)Tj
1.4683 0 Td
(activity)Tj
3.1181 0 Td
(of)Tj
1.0148 0 Td
(rhodopsin)Tj
4.354 0 Td
(still)Tj
1.6384 0 Td
(approximates)Tj
5.6353 0 Td
(fractional)Tj
4.0365 0 Td
(integration)Tj
-32.8023 -1.2982 Td
(after)Tj
2.0352 0 Td
(the)Tj
1.4683 0 Td
(addition)Tj
3.6 0 Td
(of)Tj
1.0148 0 Td
(the)Tj
1.4684 0 Td
(arrestin)Tj
3.2938 0 Td
(binding)Tj
3.3505 0 Td
(process)Tj
3.1862 0 Td
(to)Tj
1.0261 0 Td
(the)Tj
1.4683 0 Td
(cone)Tj
2.1203 0 Td
(model.)Tj
2.9707 0 Td
(Furthermore,)Tj
5.6126 0 Td
(the)Tj
1.474 0 Td
(addition)Tj
-34.0892 -1.2983 Td
(of)Tj
1.0147 0 Td
(the)Tj
1.4684 0 Td
(arrestin)Tj
3.2938 0 Td
(binding)Tj
3.3506 0 Td
(process)Tj
3.1861 0 Td
(model)Tj
2.7439 0 Td
(expands)Tj
3.4979 0 Td
(the)Tj
1.4684 0 Td
(frequency)Tj
4.1895 0 Td
(range)Tj
2.4605 0 Td
(of)Tj
1.0148 0 Td
(the)Tj
1.4683 0 Td
(approximation.)Tj
6.4006 0 Td
(Our)Tj
-35.5575 -1.3039 Td
(main)Tj
2.2903 0 Td
(purpose)Tj
3.4413 0 Td
(for)Tj
1.3889 0 Td
(including)Tj
4.0422 0 Td
(these)Tj
2.2507 0 Td
(results)Tj
2.812 0 Td
(is)Tj
0.8447 0 Td
(to)Tj
1.0261 0 Td
(demonstrate)Tj
5.2327 0 Td
(that)Tj
1.7802 0 Td
(residual)Tj
3.3789 0 Td
(activities)Tj
3.7133 0 Td
(can)Tj
1.6271 0 Td
(accumu-)Tj
-33.8284 -1.2983 Td
(late)Tj
1.6214 0 Td
(in)Tj
1.0374 0 Td
(signalling)Tj
4.0649 0 Td
(processes;)Tj
4.1952 0 Td
(therefore,)Tj
4.0819 0 Td
(they)Tj
1.9219 0 Td
(should)Tj
2.9196 0 Td
(not)Tj
1.5704 0 Td
(be)Tj
1.1509 0 Td
(neglected.)Tj
4.2293 0 Td
(Finally,)Tj
3.1691 0 Td
(as)Tj
1.0148 0 Td
(the)Tj
1.4683 0 Td
(activity)Tj
3.1181 0 Td
(itself)Tj
-35.5632 -1.2983 Td
(can)Tj
1.627 0 Td
(be)Tj
1.1509 0 Td
(described)Tj
4.0478 0 Td
(as)Tj
1.0148 0 Td
(fractional)Tj
4.0422 0 Td
(integration,)Tj
4.8699 0 Td
(its)Tj
1.1509 0 Td
(effects)Tj
2.7212 0 Td
(can)Tj
1.6271 0 Td
(be)Tj
1.1508 0 Td
(predicted)Tj
4.0025 0 Td
(without)Tj
3.3279 0 Td
(explicitly)Tj
3.8324 0 Td
(model-)Tj
-34.5654 -1.2982 Td
(ling)Tj
1.7461 0 Td
(the)Tj
1.4683 0 Td
(process.)Tj
ET
Q
q
1 j
1 J
0 w
11.9999 0 0 11.9999 200.0125 309.2598 cm
BT
/F2 1 Tf
1 TL
-0.0033 Tc
0 0 Td
(Materials)Tj
4.5071 0 Td
(and)Tj
1.989 0 Td
(methods)Tj
/F11 1 Tf
-6.4961 -1.5023 Td
(Mathematical)Tj
5.9858 0 Td
(model)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 200.0125 274.2236 cm
BT
/F7 1 Tf
1 TL
-0.004 Tc
0 0 Td
(Active)Tj
2.7722 0 Td
(rhodopsin)Tj
4.354 0 Td
(is)Tj
0.8447 0 Td
(constantly)Tj
4.3427 0 Td
(deactivated)Tj
4.7338 0 Td
(by)Tj
1.1792 0 Td
(the)Tj
1.4683 0 Td
(following)Tj
3.9799 0 Td
(process:)Tj
3.4129 0 Td
(first,)Tj
2.0295 0 Td
(rhodopsin)Tj
4.354 0 Td
(is)Tj
0.8504 0 Td
(phos-)Tj
-34.3216 -1.2982 Td
(phorylated)Tj
4.524 0 Td
(5-7)Tj
1.525 0 Td
(times)Tj
2.3868 0 Td
(in)Tj
1.0375 0 Td
(rapid)Tj
2.3357 0 Td
(succession;)Tj
4.6488 0 Td
(following)Tj
3.9798 0 Td
(this,)Tj
1.8992 0 Td
(it)Tj
0.788 0 Td
(is)Tj
0.8448 0 Td
(inhibited)Tj
3.8551 0 Td
(by)Tj
1.1848 0 Td
(arrestin)Tj
3.2939 0 Td
(before)Tj
2.7382 0 Td
(finally)Tj
-35.0416 -1.2983 Td
(decaying)Tj
3.7643 0 Td
(into)Tj
1.8369 0 Td
(opsins)Tj
2.7836 0 Td
(within)Tj
2.8119 0 Td
(the)Tj
1.474 0 Td
(next)Tj
1.9559 0 Td
(few)Tj
1.6158 0 Td
(seconds)Tj
3.3675 0 Td
([)Tj
0.83 0.64 0.02 0 k
(26)Tj
0 g
()Tj
0.83 0.64 0.02 0 k
(29)Tj
0 g
(].)Tj
3.549 0 Td
(Phosphorylation)Tj
6.8541 0 Td
(rates)Tj
2.1033 0 Td
(exponentially)Tj
-32.1163 -1.2982 Td
(decrease)Tj
3.6056 0 Td
(with)Tj
1.9956 0 Td
(each)Tj
2.0296 0 Td
(successive)Tj
4.2462 0 Td
(phosphorylation:)Tj
/F12 1 Tf
7.0413 0 Td
(\r)Tj
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 393.7322 233.5748 cm
BT
/F13 1 Tf
1 TL
-0.0053 Tc
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 397.9275 235.2755 cm
BT
/F14 1 Tf
1 TL
0 0 Td
()Tj
/F12 1 Tf
-0.004 Tc
0.9921 0 Td
(\r)Tj
/F14 1 Tf
0 Tc
0.6746 0 Td
()Tj
/F7 1 Tf
-0.004 Tc
0.4876 0 Td
(0.9)Tj
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 431.2629 239.6976 cm
BT
/F13 1 Tf
1 TL
-0.0053 Tc
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 433.2472 235.2755 cm
BT
/F7 1 Tf
1 TL
-0.004 Tc
0 0 Td
(,)Tj
0.4478 0 Td
(where)Tj
/F12 1 Tf
2.6476 0 Td
(\r)Tj
/F7 1 Tf
(,)Tj
0.9014 0 Td
(is)Tj
0.8447 0 Td
(the)Tj
1.474 0 Td
(rate)Tj
1.7461 0 Td
(of)Tj
1.0148 0 Td
(the)Tj
1.4684 0 Td
(first)Tj
-33.8681 -1.3039 Td
(phosphorylation)Tj
6.8088 0 Td
([)Tj
0.83 0.64 0.02 0 k
(15)Tj
0 g
(].)Tj
2.0806 0 Td
(However,)Tj
4.0762 0 Td
(the)Tj
1.4684 0 Td
(rate)Tj
1.7461 0 Td
(of)Tj
1.0148 0 Td
(arrestin)Tj
3.2938 0 Td
(binding)Tj
3.3561 0 Td
(linearly)Tj
3.2088 0 Td
(increases)Tj
3.8324 0 Td
(with)Tj
2.0013 0 Td
(each)Tj
2.0239 0 Td
(phos-)Tj
-34.9112 -1.2982 Td
(phorylation:)Tj
/F12 1 Tf
5.125 0 Td
()Tj
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 256.0818 207.6094 cm
BT
/F13 1 Tf
1 TL
-0.0053 Tc
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 260.2771 209.2535 cm
BT
/F14 1 Tf
1 TL
0 0 Td
()Tj
/F13 1 Tf
-0.004 Tc
0.9921 0 Td
[()]TJ
/F14 1 Tf
0 Tc
0.4875 0 Td
()Tj
/F7 1 Tf
-0.004 Tc
0.4933 0 Td
(0.5,)Tj
1.6214 0 Td
(where)Tj
/F12 1 Tf
2.6475 0 Td
()Tj
ET
Q
q
1 j
1 J
0 w
7.4999 0 0 6.6659 327.5716 207.6094 cm
BT
/F13 1 Tf
1 TL
-0.0053 Tc
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
10 0 0 10 329.5559 209.2535 cm
BT
/F7 1 Tf
1 TL
-0.004 Tc
0 0 Td
(,)Tj
0.4478 0 Td
(is)Tj
0.8447 0 Td
(the)Tj
1.474 0 Td
(rate)Tj
1.7462 0 Td
(of)Tj
1.0148 0 Td
(arrestin)Tj
3.2938 0 Td
(binding)Tj
3.3505 0 Td
(to)Tj
1.0262 0 Td
(rhodopsin)Tj
4.354 0 Td
(phosphorylated)Tj
/F13 1 Tf
6.4402 0 Td
[()]TJ
/F7 1 Tf
-36.9464 -1.2982 Td
(times.)Tj
2.6135 0 Td
(This)Tj
1.9899 0 Td
(process)Tj
3.1805 0 Td
(is)Tj
0.8504 0 Td
(described)Tj
4.0535 0 Td
(and)Tj
1.729 0 Td
(modelled)Tj
3.9402 0 Td
(in)Tj
1.0374 0 Td
(detail)Tj
2.4151 0 Td
(in)Tj
1.0375 0 Td
([)Tj
0.83 0.64 0.02 0 k
(15)Tj
0 g
()Tj
0.83 0.64 0.02 0 k
(17)Tj
0 g
(],)Tj
3.5433 0 Td
(which)Tj
2.6532 0 Td
(we)Tj
1.3266 0 Td
(used)Tj
2.0579 0 Td
(as)Tj
1.0148 0 Td
(a)Tj
0.652 0 Td
(founda-)Tj
-34.0948 -1.3039 Td
(tion)Tj
1.8425 0 Td
(for)Tj
1.3833 0 Td
(our)Tj
1.6271 0 Td
(model)Tj
2.7439 0 Td
(\(Eqs)Tj
0.83 0.64 0.02 0 k
1.9956 0 Td
(1a)Tj
0 g
()Tj
0.83 0.64 0.02 0 k
(1d)Tj
0 g
(\).)Tj
3.2144 0 Td
(We)Tj
1.61 0 Td
(extended)Tj
3.8608 0 Td
(the)Tj
1.4683 0 Td
(model)Tj
2.7439 0 Td
(by)Tj
1.1792 0 Td
(adding)Tj
2.9934 0 Td
(the)Tj
1.4683 0 Td
(stage)Tj
2.2054 0 Td
(where)Tj
2.6475 0 Td
(rhodopsin)Tj
-32.9836 -1.2983 Td
(is)Tj
0.8447 0 Td
(inhibited)Tj
3.8551 0 Td
(by)Tj
1.1849 0 Td
(arrestin,)Tj
3.5206 0 Td
(and)Tj
1.7291 0 Td
(retains)Tj
2.9253 0 Td
(only)Tj
1.9729 0 Td
(a)Tj
0.6576 0 Td
(fraction)Tj
3.3562 0 Td
(of)Tj
1.0148 0 Td
(its)Tj
1.1452 0 Td
(original)Tj
3.3278 0 Td
(activity)Tj
3.1181 0 Td
(\()Tj
0.83 0.64 0.02 0 k
(Eq)Tj
1.6328 0 Td
(1e)Tj
0 g
(\))Tj
1.4626 0 Td
([)Tj
0.83 0.64 0.02 0 k
(10)Tj
0 g
(,)Tj
0.83 0.64 0.02 0 k
1.7348 0 Td
(30)Tj
0 g
()Tj
0.83 0.64 0.02 0 k
(33)Tj
0 g
(].)Tj
-33.4825 -1.2983 Td
(With)Tj
2.279 0 Td
(each)Tj
2.0296 0 Td
(phosphorylation,)Tj
7.0356 0 Td
(rhodopsin)Tj
4.3539 0 Td
(is)Tj
0.8504 0 Td
(inhibited)Tj
3.8494 0 Td
(by)Tj
1.1849 0 Td
(50%)Tj
1.9275 0 Td
([)Tj
0.83 0.64 0.02 0 k
(15)Tj
0 g
(],)Tj
2.075 0 Td
(and)Tj
1.7347 0 Td
(the)Tj
1.4684 0 Td
(binding)Tj
3.3505 0 Td
(of)Tj
1.0148 0 Td
(arrestin)Tj
-33.1537 -1.3039 Td
(further)Tj
3.0274 0 Td
(inhibits)Tj
3.2655 0 Td
(activity)Tj
3.1181 0 Td
(by)Tj
/F13 1 Tf
1.1792 0 Td
[()]TJ
/F7 1 Tf
0.7086 0 Td
(=)Tj
0.7994 0 Td
(5090%)Tj
3.3958 0 Td
([)Tj
0.83 0.64 0.02 0 k
(30)Tj
0 g
()Tj
0.83 0.64 0.02 0 k
(32)Tj
0 g
(].)Tj
-14.2978 -1.2982 Td
(The)Tj
1.7858 0 Td
(equations)Tj
4.0932 0 Td
(of)Tj
1.0148 0 Td
(the)Tj
1.4684 0 Td
(model)Tj
2.7439 0 Td
(for)Tj
1.3889 0 Td
(6)Tj
0.6916 0 Td
(phosphorylations)Tj
7.1773 0 Td
(are)Tj
1.4456 0 Td
(as)Tj
1.0148 0 Td
(follows:)Tj
/F15 1 Tf
0 Tc
-10.7432 -1.8312 Td
(_)Tj
/F13 1 Tf
-0.085 -0.017 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 336.359 110.2677 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 349.9086 112.7622 cm
BT
/F14 1 Tf
1 TL
0 0 Td
()Tj
/F16 1 Tf
1.7688 0 Td
[()1()19()11()]TJ
1.8311 0 Td
[()]TJ
/F14 1 Tf
()Tj
/F13 1 Tf
[()]TJ
/F14 1 Tf
1.0318 0 Td
()Tj
0.601 0 Td
( )Tj
/F17 1 Tf
0.9978 0 Td
(g)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 416.5795 110.2677 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 420.0944 112.7622 cm
BT
/F13 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 423.7228 110.2677 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 559.1622 112.7055 cm
BT
/F14 1 Tf
1 TL
0 0 Td
()Tj
/F7 1 Tf
(1a)Tj
/F14 1 Tf
()Tj
/F15 1 Tf
-23.2496 -2.6928 Td
(_)Tj
/F13 1 Tf
-0.085 -0.0171 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 330.2929 83.1118 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 343.7858 85.6062 cm
BT
/F14 1 Tf
1 TL
0 0 Td
()Tj
/F17 1 Tf
1.7688 0 Td
(g)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 365.8393 83.1118 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 369.3543 85.6062 cm
BT
/F13 1 Tf
1 TL
0 0 Td
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 373.0393 83.1118 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(0)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 378.7086 85.6062 cm
BT
/F14 1 Tf
1 TL
0 0 Td
( )Tj
0.9977 0 Td
()Tj
/F17 1 Tf
(g)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 396.907 83.1118 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 402.633 85.6062 cm
BT
/F14 1 Tf
1 TL
0 0 Td
()Tj
/F17 1 Tf
0.9921 0 Td
(b)Tj
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 418.1102 83.1118 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 421.5685 85.6062 cm
BT
/F14 1 Tf
1 TL
0 0 Td
()Tj
/F13 1 Tf
[()]TJ
ET
Q
q
1 j
1 J
0 w
5.9998 0 0 5.9998 429.1086 83.1118 cm
BT
/F15 1 Tf
1 TL
0 0 Td
(1)Tj
ET
Q
q
1 j
1 J
0 w
10 0 0 10 558.4251 85.5496 cm
BT
/F14 1 Tf
1 TL
0 0 Td
()Tj
/F7 1 Tf
(1b)Tj
/F14 1 Tf
()Tj
ET
Q
q
1 j
1 J
0 w
576 737.1 m
36 737.1 l
36 737.6 l
576 737.6 l
f*
36 741.2598 107.1496 23.6976 re
W* n
q
107.0929 0 0 23.6409 36 741.3165 cm
q
/I1 Do
Q
Q
Q
q
0 0 612 792 re
W* n
1 j
1 J
0 w
7.9999 0 0 7.9999 403.1433 745.7952 cm
BT
/F0 1 Tf
1 TL
-0.005 Tc
0 0 Td
(Signal)Tj
2.9409 0 Td
(processing)Tj
4.9678 0 Td
(of)Tj
1.0417 0 Td
(rhodops)Tj
3.5433 0 Td
(in)Tj
0.978 0 Td
(in)Tj
0.978 0 Td
(vertebrate)Tj
4.6488 0 Td
(vision)Tj
ET
Q
q
1 j
1 J
0 w
36 48.0002 m
576 48.0002 l
576 47.5002 l
36 47.5002 l
f*
0.83 0.64 0.02 0 k
81.5811 34.9228 m
237.8835 34.9228 l
h
f*
0 g
7.9999 0 0 7.9999 36 36 cm
BT
/F0 1 Tf
1 TL
-0.005 Tc
0 0 Td
(PLOS)Tj
2.8559 0 Td
(ONE)Tj
2.3669 0 Td
(|)Tj
0.83 0.64 0.02 0 k
0.4748 0 Td
(https://doi.or)Tj
5.4142 0 Td
(g/10.137)Tj
3.8198 0 Td
(1/journal.po)Tj
5.1236 0 Td
(ne.02050)Tj
4.089 0 Td
(99)Tj
0 g
2.0906 0 Td
(October)Tj
3.7276 0 Td
(4,)Tj
1.0417 0 Td
(2018)Tj
34.1365 0 Td
(2)Tj
0.7654 0 Td
(/)Tj
ET
endstream
endobj
47 0 obj
<>stream
endstream
endobj
48 0 obj
<>stream
endstream
endobj
49 0 obj
<>stream
endstream
endobj
50 0 obj
<>stream
endstream
endobj
51 0 obj
<>stream
endstream
endobj
52 0 obj
<>stream
endstream
endobj
53 0 obj
<>stream
endstream
endobj
54 0 obj
<>stream
endstream
endobj
55 0 obj
<>stream
endstream
endobj
56 0 obj
<>stream
endstream
endobj
57 0 obj
<>stream
endstream
endobj
58 0 obj
<>stream
endstream
endobj
59 0 obj
<>stream
endstream
endobj
60 0 obj
<>stream
endstream
endobj
61 0 obj
<>stream
endstream
endobj
62 0 obj
<>stream
endstream
endobj
63 0 obj
<>stream
endstream
endobj
64 0 obj
<>stream
endstream
endobj
65 0 obj
<>stream
endstream
endobj
66 0 obj
<>stream
endstream
endobj
67 0 obj
<>stream
endstream
endobj
68 0 obj
<>stream
endstream
endobj
69 0 obj
<>stream
BT
66.402 0 Td
(11)Tj
ET
Q
endstream
endobj
70 0 obj
<>/XObject<>>>/CropBox[0 0 612 792]/MediaBox[0 0 612 792]/Parent 16 0 R/Annots 75 0 R/Contents 76 0 R/TrimBox[0 0 612 792]>>
endobj
74 0 obj
<>stream
xie~kgs֭>NիOiAi (*(P "†R e%b2d
Iΐ(<#S&$S2YOut#c;/EE~Ķg =z͛Ao@ @ ̜9sɃ
_OO__Þ={dccc: #'" )`}+a@ @ 4H;x[Z@ @ EޑǧJ1/E_ +WP|tȑ" %id]`Eݖ@ @ J =8MjI 42.|Mȣ ` MQ{i>>@ @ 74 8=MxMJu@ E@QqR?r{*E @ ;%̷ދIwa 'y(NZheU{
Ԍ Gsfw0iǧ;@ Ȃ:89NG}D @ 2.u&_Oc7 @9Syf\wlB @ (;H',- @6FxL+8>ҏ7o&F @ #u&(>*NmS@ ι9846 'PP/羺