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Thesis-PDF - IAP/TU Wien

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Figure 4.7: Scheme of the photoreceptor photocycle (intermediate states<br />

omitted). Upon absorption of light with a wavelength of around 365 nm the<br />

rhodopsin-like protein changes into the excited state, then upon a second<br />

absorption at a wavelength of around 436 nm it falls back again into the<br />

ground state. This last step from the excited state to the ground state is<br />

almost exclusively triggered by light. It occurs only once in a thousand years<br />

through thermal deactivation at ambient temperature.<br />

Sensitivity<br />

For detecting the direction of the light of a specific range a photoreceptor demands<br />

a high packing density of chromophore molecules organized in a lattice<br />

structure, with high absorption cross section (high absorption probability) of the<br />

chromophore and very low dark noise 10 . For transmitting the detected signal a<br />

photoreceptor must generate an electrical potential difference, or an electrical current.<br />

This is exactly what Euglena’s photoreceptor is designed for.<br />

One of the most investigated photoreception systems is that of Chlamydomonas.<br />

It consist of a patch of rhodopsin-like proteins in the plasma membrane.<br />

The packing density of these molecules appears to be about 20 − 30, 000/µ 2 m of<br />

membrane, with a molar absorption coefficient of 40 − 60, 000 M −1 ∗ cm −1 and a<br />

dark noise of almost zero. The number of embedded molecules per µ 2 m of membrane,<br />

the absorption, the absorption cross section, and the dark noise are at the<br />

best of theoretical limits. Nevertheless, the fraction of photons absorbed from a<br />

single layer of these molecules is less than 0.05%.<br />

An estimate of how many photons this simplest but real photoreceptive system<br />

can absorb is effectuated here. On a sunny day about 10 18 photons ∗m −2 ∗ s −1 per<br />

nanometer wavelength are incident. On a cloudy day this lowers to 10 17 photons<br />

∗m −2 ∗ s −1 per nanometer wavelength. Through absorption of the water column<br />

this number is lowered to 10 17 and 10 16 respectively. Thus a photoreceptor of a<br />

cross section of about 1µm 2 can catch at most 10 5 photons, 10 7 in its 100 nm<br />

absorption band. Since only about 0.05% of these are effectively catched by the<br />

protein monolayer, the maximum number lowers to 5 ∗ 10 3 . Now during revolving<br />

motion of the algae it is illuminated for about 400 ms and shaded by the stigma<br />

10 Dark noise is inherent to such a receptor and independent of light level. It arises from the<br />

random thermal motion of the molecules.<br />

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