Responses of lotic periphyton to pulses of phosphorus: P-flux controlled growth rate


  • Reproduced with the permission of the Minister of Environment.

Max L. Bothwell, Environment Canada, Pacific Biological Station, 3190 Hammond Bay Road, Nanaimo, BC V9T 6N7, Canada. E-mail:


1. The response of stream periphyton to the addition of limiting nutrients has been the focus of many studies. However, the influence of pulsed nutrient additions has not previously been examined. This study investigates the biomass accrual and physiological responses of phosphorus-limited lotic periphyton to hourly phosphate fluxes.

2. Two pulsing experiments were conducted: (i) a variable flux trial that compared variable hourly P-fluxes, delivered either continuously at different concentrations or at the same concentration but in pulses of differing duration per hour and (ii) a constant flux trial that compared periphyton responses at a set hourly P-flux but delivered in pulses of varying concentration and duration.

3. Growth response and alkaline phosphatase activity during the variable flux experiment showed that periphyton responds to the hourly integrated flux of phosphorus, regardless of whether the nutrient is supplied in short concentrated pulses or continuously at much lower concentrations.

4. The constant flux experiment examined the pulse period required to attain maximum biomass for a given phosphorus flux. Periphyton response to 5-min pulses of phosphate per hour approximated the maximum biomass as that attained when the same hourly flux was added continuously. Compared with the control, there was also a substantial increase in biomass with pulses of only 1 min each hour. These results demonstrate that the hourly average phosphate concentration to which periphyton communities are exposed is paramount in determining P-limited growth dynamics.

5. Species composition was not significantly different among treatments in each experiment; however, the design was to evaluate monotonic response with increasing phosphorus flux and species diversity may not respond monotonically. The data are therefore preliminary but suggest the need to determine if species diversity is generally lower when there are brief pulses of phosphate. Unlike pulse experiments that mimic lentic situations, nutrient additions were not used to completion and species success and composition was more dependent on their ability to acquire limiting nutrients rapidly rather than on their ability to take up nutrients at the lowest concentration.