Measurements of Phytoplankton Absorption Other Than Per Unit of Chlorophyll a

Phytoplankton plays a critical role in determining light fields of the world’s oceans, primarily through absorption of light by photosynthetic pigments (see Chapters 1 to 5). Consequently there has been considerable interest from optical researchers in determining phytoplankton absorption. Conversely, from the biological point of view, this absorption assumes paramount importance because it is the sole source of energy for photosynthesis and thus should be central to direct estimates of primary production. There are two logical parts in determining this effect of phytoplankton and in estimating primary production. One is the estimation of abundance, and the other is estimation of specific effect or specific production rate. The earliest estimates of phytoplankton abundance were based on cell counts. From the time of Francis A. Richards’ Ph.D. dissertation, however, measurement of chlorophyll a concentration per unit of water volume, because of its relative ease, has assumed a central role in abundance estimation. Physiological studies and technological advances in optical instrumentation over the last decade lead me to question whether the continued use of chlorophyll a concentration to estimate phytoplankton abundance was wise either from the viewpoint of narrowing confidence intervals on estimates of absorption and production or from the viewpoint of mechanistic understanding of the processes involved. The measurement of chlorophyll a has become such a routine tool of biological oceanography, however, that the reasons for my heresy require elaboration. Some of the reasons are not too subtle. Chlorophyll a exists with other photosynthetic pigments in organized arrays associated with photosynthetic membranes. The function of these arrays is to harvest photons and transfer their energy to the specialized reaction center complexes that mediate photochemistry (see Chapter 9). The size of the arrays or packages and the ratio of chlorophyll a molecules to other light-harvesting pigments within the packages vary with phytoplankton cell size, total irradiance and its spectral distribution, as well as with other environmental parameters. It is well known that dark-adapted (= light-limited) cells increase their complements of photopigments. This plasticity in pigment packaging is evidenced in the variability of chlorophyll a-specific absorption coefficients. Simple optical models based only on chlorophyll a concentrations cannot be accurate or precise unless the effects of pigment packaging are considered.