Daphnia collects more food at low particle concentrations by increasing its maximum filtering rate either by enlarging the area of its filter screens or by increasing its appendage beat rate (ABR). Various species of Daphnia grown at low levels of food phenotypically enlarge their filter screens. Models of the flow across a filter predict that the energy required to obtain a certain increase in filtering rate is a linear function of the screen area but increases proportionally to the square of ABR. A daphniid should be able to increase its filtering rate without expending more energy if it enlarges the filter and sirnultaneously reduces ABR. To test the model predictions, we measured morphometric parameters (areas, OF en space, intersetular distances) of the filters and ABR for Daphnia adapted to high food and to low food. Daphniids adapted to low food not only have larger filter screens, they also have finer meshes. In Daphnia adapted to low food, mean ABR is slightly reduced and there is a negative relationship between filter-screen area and ABR in the low-food treatment. Gains in filtering rate are higher and reductions in ABR are lower than predicted by the equilibrium model, hence Daphnia optimizes input rather than minimizing energy expenditures for filtering.Since Koza and KoEinek (1985) reported an increase in the size of filtering screens on the 3rd and 4th limbs of Daphnia at low concentrations of food, more evidence has accumulated that indicates that this is a common phenomenon in the genus (e.g. Pop 199 1; Stuchlik 199 1). Lampert (1994) showed experimentally that daphniids grown at very low concentrations of food can have filter screen areas twice as large as those grown at high levels of food. He interpreted this response as a phenotypic adaptation to a low-food environment.At low concentrations of food, higher filtering rates result in higher feeding rates (McMahon and Rigler 1965). Increased feeding rates result in lower threshold food concentrations for growth (Lampert 1977), hence very small differences in filtering rates have a pronounced effect on the animal's energy budget when food is scarce (Lampert 1984).If daphniids phenotypically adjust their filter-screen areas under low-food conditions to gather more food or decrease the areas at high levels of food instead of maintaining large filters all the time, one might expect that maintenance of large filter bears some costs that can be saved by having smaller filters. To increase filtering rate, a daphniid would have two options, increase either the pumping rate (appendage beat rate) or the efficiency of the filtering structures (e.g. filter-screen size). The first option would increase the flow rate across the filter meshes while the second option would have no effect on the
AcknowledgmentsWe thank James F. Haney and John J. Sasner for allowing us to copy their video system for appendage beat rate measurements. Maren Volquardscn and Heinke Claussen measured filter screen areas of daphniids used for beat rate analysis, and Nancy Zeh...