PRIMARY PRODUCTIVITY •Productivity is the rate of biomass formation •Primary productivity (photosynthesis) of phytoplankton can be measured directly by O2 production •Photosynthesis can be summarized as 6CO2 + 12 H20 ----> C6H12O6 + 6O2 + 6H2O •Respiration is the reverse reaction The most common techniques for PP measurement are: •the light-dark bottles technique for O2 production •14C uptake measurements of CO2 assimilation •harvest method •Whole water column O2 or CO2 flux Primary Producers in Aquatic Ecosystems Cyanobacteria, Protista (microalgae, macroalgae), Aquatic plants These groups are all found in both standing and flowing water In standing water the two main primary producer communities are: a.) the phytoplankton (suspended cells or colonies of microalgae or cyanobacteria) b.) the littoral community (benthic micro- or macroalgae or cyanobacteria, epiphytic microalgae, and macophytes (macroalgae or plants)-emergent/submerged In running water the two main primary producer communities are: a.) benthic algae attached to rocks (generally in fast current) b.) submerged macrophytes & associated epiphytes—in areas of slow current Nightime mg/L O2 Afternoon mg/L O2 2 4 6 5 8 10 2 4 6 8 5 Depth m Depth m 10 10 Average [O2] Epilimnion = 9.8 mg/L Average [O2] Epilimnion = 6.9 mg/L 10 Van Dorn sampler obtains samples of water from a desired depth A Yellow Springs Instrument DO meter being calibrated Changes in Dissolved Oxygen •Water samples from various depths are enclosed in light (transparent) and dark (completely opaque) bottles, •For each depth initial readings of dissolved O2 are taken (IB) and the light (LB) and dark (DB) samples are incubated for a period long enough to produce measurable changes in O2. •During the incubation, we expect that the initial DO concentration (IB) at a given depth will decrease to a lower concentration in the dark bottles (DB) due to respiration of phytoplankton. •Conversely, we expect light bottle (LB) should increase from their initial values (IB). IB DB respiratio n rate/unit volume t LB IB net photosynth etic rate/unit volume t LB DB gross photosynth etic rate/unit volume t Productivity measurements are usually expressed in terms of carbon not oxygen Therefore we need to convert the changes in oxygen concentration to corresponding changes in carbon. For this we use the photosynthetic quotient (PQ) and the respiratory quotient (RQ), which are ratios describing the relative amounts of oxygen and carbon involved in photosynthesis and respiration. : PQ = (molecules of oxygen liberated during photosynthesis) / (molecules of CO2 assimilated)= usually around 1.2 RQ = (molecules of CO2 liberated during respiration) / (molecules of oxygen consumed) = usually very close to 1.0 Calculations •Gross photosynthesis = [(LB - DB) * 1000 * 0.375] / (PQ * t) •Net photosynthesis = [(LB - IB) * 1000 * 0.375] / (PQ * t) •Respiration = [(IB - DB) * RQ * 1000 * 0.375] / t •Gross and net photosynthesis and respiration are expressed as mg C/m3/t •LB, DB, and IB are dissolved oxygen concentrations in mg/L • t is the incubation period eg. hours •1000 converts L to m3 (1 L = 1000 cm3) •0.375 converts mass of oxygen to mass of carbon and is a ratio of moles of carbon to moles of oxygen (12 mg C/32 mg O2 = 0.375) Because we are using bottles incubated at various depths in the photic zone to measure primary productivity in situ, and the measurements vary with depth, we need to obtain a profile of primary productivity with depth. % of surface irradiance Primary productivity (mgC/m3/d) Depth (z) Let us say that we wish to incubate samples at the 75% , 50%, 25%, 10% and 1% light levels. How do we decide at what depths to incubate samples at? The Secchi disk—a simple way to estimate light extinction 1.7 k Secchi disk trans parency (m) Light extinction --Light enters from above and its intensity (I) is sharply attenuated with depth (z)—absorption by water or solute molecules or scattered by particles Section 10.6 Iz z 50% z 10% At what depth z 1% while the light intensity be 1% of I 0 Iz e kz so 0.01 e kz and ln 0.01 kz... Photic I 0 zone z 4.6 , k for the depth at which light intensity is 10% of I 0 2.3 , k for the 50% light level z z 1% z z Page 144 in text 0.69 k For a lake with Secchi disk transparency of 3m k=0.57 (1.7/3) %light z=-ln (%light/100)/k (k=0.57) ln(%light/100) 0.75 -0.29 0.50 m 0.5 -0.69 1.22 m 0.25 -1.39 2.43 m 0.1 -2.30 4.04 m 0.01 -4.61 8.08 m If we place dark and light bottles at each of these depths, and calculate the Gross Primary productivity at each GPPz gC/m3/d 1.5 0.5 1.0 z 75% z 50% x x x z 10% x m z 1% z x Photic zone Find the area under the GPP vertical profile Its units will be gC/m2/d, Why? If we place dark and light bottles at each of these depths, and calculate the Gross Primary productivity at each Find the area under the GPP vertical profile Its units will be gC/m2/d, Why? GPPz gC/m3/d 1.5 0.5 1.0 0.45 sq in x 1.0 B x x 3.0 Photic zone A Area A represents [4.10/0.45] *1.0 gC/m2/d=9.1 gC/m2/d Average productivity per unit volume 9.1gC/m2/d / 9.1 m = 1.0 gC/m3/d x 5.0 7.0 x zm Area B represents 0.5 gC/m3/d * 2 m = 1.0 gC/m2/d 4.10 sq in If you average all the GPP estimates 1.2,1.65,1.70, 1.1,0.22, =1.174 gC/m3/d x 9.1 m = 10.7 gC/m2/d Averaging the readings leads to a nearly 20% overestimate, Why? Primary producers differ in their photosynthetic responses to light intensity Comparing the Photosynthethis/irradiance curves among species A GPP/t per unit of biomass B C 10% 25% 50% Light intensity Assuming that A, B and C are similar in all respects other than their P/I curves, which of these species would you expect to perform best in the well mixed water column of a deep lake (25% light level at 10 m, max depth 100 m) a) A b) B c) C d) A and B would do equally well e) A and C would do equally well A GPP/t per unit of biomass B C 10% 25% 50% Light intensity Assuming that A and B are equal in all respects other than their P/I curves, under what conditions would you expect B to outperform A? a) in the epilimnion of a clear stratified lake, (Assume 25% light level at the thermocline) b) in the well-mixed water column of a deep lake (Assume 25% light level at 10m, max depth 100 m) c) in the hypolimnion of a clear stratified lake (Assume 25% light level at the thermocline) d) growing on the substrate near shore in the littoral zone of a clear lake (assume 25 % light level at the outer boundary of the littoral) e) both a and d f) both b and c A GPP/t per unit of biomass B Could C ever be expected to dominate? C 10% 25% 50% Light intensity The seasonal dyanamics of the phytoplankton in lakes •Temperature adaptations of different algal groups •Thermal stratification, sinking rates and nutrient dynamics •Food-web interaction—effects of grazing zooplankton mid-summer low biomass community shifts to inedible forms Early spring—diatoms dominate--under cold temperatures and low light conditions plenty of nutrients in the well mixed water column Summer—lake warms up, thermocline forms diatoms fall out of the mixing layer—low viscosity and low mixing depth Asterionella the only diatom that can still hang in. Mid-summer—nutrients lost from mixed layer (sedimentation of algae), warm temperatures favour green algae, and zooplankton herbivory is high favouring fast growing small species eg Chlorella Late summer—herbivores eliminate edible species, large colonial cyanobacteria dominate eg. Microcystis Fall—water cooling, thermocline breaks up, mixing depth increases, nutrients increase, diatoms dominate Winter—low light and cold temp low biomass Cyclotella:centric diatom (around 20 microns) and Stephanodiscus (around 50 microns) Usually abundant in spring plankton Asterionella colonial diatom—elongate cells joined at the base to form stellate colonies Fairly large for planktonic diatoms, each cell 50-70 microns Commonly found in dense blooms during May, prior to the onset of thermal stratification Chlorella: a unicellular non-flagellated green algae Lake Zooplankton Cladocera Rotifers Copepod Larvae--Nauplius Copepods Anabaena: cells in large colonies (filaments) with no gelatinous matrix, coiled or straight, heterocysts and akinetes usually present, cells 3-5 microns Microcystis: cells in large colonies irregularly arranged within a gelatinous matrix Colony of Microcystis A common bloom forming cynabacterium that sometimes can be highly toxic Colonial growth pattern—cells embedded in a gelatinous matrix (3b) Adaptations of benthic diatoms to life in streams They can live either as solitary cells, usually raphed—capable of slow gliding movement on rocks or in chains joined at the valvular surface, attached to substrate either by mucilage pad or by a stalk http://craticula.ncl.ac.uk/EADiatomKey/html/Achnanthes.html Achnanthes longipes: alternation between a motile, solitary phase and a stalked sessile phase When inoculated into fresh media, the cells are at first motile (I), then become sessile and produce a stalk that anchors them to the substratum (II). This stalk continues to be synthesized as the cell is pushed away from the substratum (III), and eventually mitosis occurs producing a row of cells stacked one upon the other (IV). These cells eventually detach from one another (I) and begin the cycle again. Why is this life cycle be advantageous for life in streams? http://www.bio.mtu.edu/the_wall/integrated_microscopy/a_longipes_general.html Daily fluctuations in O2 provides and integrated measure of community metabolism over a reach. Daily cycle in dissolved oxygen (flow weighted average) at Pavan study site Pavan 12.50 12.00 11.50 11.00 Dissolved O2 mg / l 10.50 100% saturation 10.00 9.50 9.00 8.50 8.00 0:00 8AM 2:24 4:48 2PM7:12 9:36 8PM 12:00 Tim e 14:24 2AM19:12 16:48 21:36 8AM 0:00 A B C O2 mg/L Day Night Which one of these curves is from a treated sewage effluent and which is from an untreated sewage effluent? You find that C outperforms both A and B during the summer months but not in the early spring. Assuming that all three species have similar temperature optima and nutrient uptake affinities, which of the following explanations is most plausible? a) C is the least palatable species to herbivorous zooplankton b) B does best at low light intensity, and A does best at high light intensity, but C does best under fluctuating light intensities c) C has the most eccentric shape d) a and b are both plausible e) a and c are both plausible A GPP/t per unit of biomass B C 10% 25% 50% Light intensity