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202                                   Algae: Anatomy, Biochemistry, and Biotechnology

                  with the molecular transparency of the atmosphere and the distance the light must travel through it;
                  therefore, it varies with altitude and meteorological conditions. In water, upwelling (bottom
                  reflected radiance) and downwelling (depth attenuated radiance) irradiance are usually quantified
                  using paired cosine collectors mounted on a frame that is lowered in the water column. Spherical
                  collectors are also mounted on a frame and lowered in the water to obtain light profiles. For photo-
                  physiological studies, spherical collectors provide a realistic measure of the total amount of PAR
                  available for photosynthesis in a water body.
                     Accurate measurement of photosynthetic radiation is essential for controlled environmental
                  research. The most common method of measuring PAR gives equal value, that is, equal energy
                  content, to all photons with wavelength between 400 and 700 nm and is referred to as the photo-
                  synthetic photon flux (PPF) measured in mmol m 22  sec 21 . The ideal PPF sensor would respond
                  equally to all photons between 400 and 700 nm. However, photosynthesis is driven also by
                  photons with wavelength below 400 and above 700 nm, and photons of different wavelength
                  induce inequal amount of photosynthesis. For these reasons, an accurate measurement of PAR
                  should follow the relative quantum efficiency (RQE) curve, which weights the photosynthetic
                  value of all photons with wavelength from 360 to 760 nm. The RQE resembles the absorption spec-
                  trum of the absorbing algal structures. A sensor that responds according to this curve measures yield
                  photon flux (YPF) in micromoles per square meter per second, the same unit of PPF.
                     Detector designed to measure YPF or PPF are commercially available. Both types consist of
                  an optical diffuser collector (planar or spherical) and multiple-spectral filters in front of a broad-
                  spectrum photovoltaic detector.
                     Planar detectors measure only unreflected sunlight. They should possess cosine corrected
                  response; as the light impinging on the measurement plane is proportional to the cosine of the
                  angle at which the light is incident, the response of the detector must also be proportional to the
                  cosine of the incidence angle [Equation (5.22)]. An integrating sphere measures both unreflected
                  and all the scattered sunlight; it needs no corrections because it is the ideal cosine detector, respond-
                  ing equally to light coming from all directions.
                     The peak responsivity and bandwidth of a detector can be controlled through the use of filters.
                  Filters can suppress out-of-band light but cannot boost signal, and restricts the spatial responsivity
                  of the detector. Filters operate by absorption or interference. Colored glass filters are doped with
                  materials that selectively absorb light by wavelength, and obey Lambert–Beer law. The peak trans-
                  mission is inherent to the additives, while bandwidth is dependent on thickness. Interference filters,
                  which select very narrow bandwidths, rely on thin layers of dielectric to cause constructive inter-
                  ference between the wavefronts of the incident white light. So by varying filter thickness, we can
                  selectively modify the spectral responsivity of a detector to match a particular function. Pass band
                  filters are often used to select a specific wavelength radiation. Any of these filter types can be com-
                  bined to form a composite filter that matches a particular photobiological process. Multiple filters
                  cemented in layers give a net transmission equal to the product of the individual transmissions. The
                  detector’s overall spectral sensitivity is equal to the product of the responsivity of the detector and
                  the transmission of the filters. Given a desired overall sensitivity and known detector responsivity,
                  we can solve for the ideal filter transmission curve.
                     Different measurement geometries necessitate specialized input optics. Radiance and lumi-
                  nance measurements require a narrow viewing angle (,48) in order to satisfy the conditions under-
                  lying the measurement units. Power measurements, on the other hand, require a uniform response
                  to radiation regardless of input angle to capture all the light.


                  PHOTOSYNTHESIS–IRRADIANCE RESPONSE CURVE (P VERSUS E CURVE)
                  The P versus E curve, or light-response curve, shows how photosynthetic rate varies with increas-
                  ing irradiance. The rate of photosynthesis can be measured as net O 2 evolution. This curve
                  describes a complex multi-step process that depends on several limiting factors. Three regions
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