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Structure and Productivity of Aquatic Systems Functional Lake Zones Pelagial Living Things in Lakes Distribution & abundance of living things in lake controlled by physical and chemical conditions in different zones Organic Matter in Lakes Living things make up only small portion of organic matter in lakes Most is in form of non-living detritus Both particulate and dissolved Organic Matter in Lakes In most lakes, dissolved organic matter is 10 X more abundant than particulate Living things make up small portion of particulate Detritus is habitat & energy resource for living things Organic Matter in Lakes Much of the organic production of photosynthesis within a system is not consumed, but becomes part of detritus reserve Primary Producers in Lakes 3 major categories of primary producers: Phytoplankton Photic zone throughout lake Generally small, unicellular or colonial organisms Primary Producers in Lakes Emergent macrophytes Shallow portions of littoral zone Roots and lower portions in water, tops above water surface Primary Producers in Lakes Submersed macrophytes Deeper portions of littoral zone Completely underwater Productivity Hierarchy Emergents most productive (Carbon fixed/area/year) More productive than terrestrial grassland, forest Submersed much less productive Phytoplankton least productive Phytoplankton Cyanobacteria or blue-green algae Important nitrogen fixers High densities in late-summer Odor (and taste) problems Phytoplankton Desmids Green algae Tremendous diversity Planktonic, but can be attached, benthic (often filamentous) Phytoplankton Golden-brown algae Low diversity, but can be important segment of phytoplankton Dinobryon important under low P conditions Phytoplankton Diatoms Very important group Planktonic and attached forms Cell walls with silica -- maximum abundance in spring when silica is most abundant Phytoplankton Cryptomonads Extremely small May reach high densities during cold periods with low light intensities (winter under ice) Phytoplankton Dinoflagellates Unicellular, flagellated, with spines Strict requirements for Ca, pH, temperature, dissolved organics Phytoplankton Some exhibit cyclomorphosis seasonal change in size & form Ceratium - more spines, longer spines, more divergent spines as water temperature increases Reduce sinking rate out of photic zone in less viscous water Phytoplankton Euglenoids Unicellular Most abundant in areas with high ammonia, dissolved organics Shallow farm ponds in cow pastures Paradox of the Plankton Lakes usually have a few dominant species and many rarer species Theoretically should have only single dominant species (niche overlap leads to competitive exclusion) Paradox of the Plankton Multispecies equilibrium in open waters 4 possible explanations: Paradox of the Plankton Environmental change too rapid for competitive exclusion to occur Symbiotic relations among species (commensalism) Selective grazing on competitive dominants by zooplankton (size-based) Some species alternating between plankton and benthos Not truly competing with pure planktonic forms Phytoplankton and Water Quality Assemblage indicates level of nutrient enrichment Desmids and certain diatoms in nutrientpoor systems Different diatoms, greens, and bluegreens dominate as enrichment increases Phytoplankton and Environmental Factors Temperature and light control type, abundance of plankton Diatoms have lower temperature optimum, blue-greens higher optimum Phytoplankton and Environmental Factors Many can adapt to changing light intensities Chlorella changes pigments per cell to maintain same rate of photosynthesis Blue-greens regulate gas pressure in vacuoles to position themselves at depth with optimum light intensities Phytoplankton and Environmental Factors Some phytoplankton experience photoinhibition High light intensities near lake surface may temporarily destroy enzymes and decrease photosynthesis Sunny days - less photosynthesis near surface than at greater depths Phytoplankton - Seasonal Succession Changes in light, nutrients, temperature drive a shift in phytoplankton during the year Phytoplankton - Seasonal Succession Low growth in winter Diatoms and cryptophytes dominate in spring Greens take over in summer, joined or replaced by bluegreens as N runs low in productive lakes Less productive lakes few greens, blue-greens, only peaks of diatoms spring and fall (silica) Phytoplankton - Seasonal Succession Seasonal abundance varies much more in temperate (1000 X) than in tropical (5 X) lakes, but total populations are much greater in tropical lakes Selective grazing by zooplankton can influence succession Eating some, providing nutrients for others Phytoplankton - Nutrient Enrichment Enrichment can greatly increase productivity (per volume) up to a point Eventually self-shading develops and thickness of photic zone reduced Inhibits further increases Productivity/m2 of surface remains virtually unchanged Photosynthetic efficiency low (<1% of incident light) Phytoplankton - Variation in Production More production in littoral zones than pelagial areas Peak production during midday (except at surface - earlier in day) Seasonal production peaks in summer Macrophytes Restricted to the littoral zones In small, shallow lakes with no profundal zone, macrophytes may occur basin-wide Emergent Macrophytes Rooted in water or saturated soil with aerial leaves/stems Upper littoral - out to 1.5 m depth Typha - cattail Emergent Macrophytes Special category occupying midlittoral region - 0.53.0 m Floating-leaved plants Water lily Submersed Macrophytes All depths within photic zone down to ~10 m for vascular plants Macroalgae - may occur slightly deeper Coontail, curlyleaf pondweed, Elodea Free-floating Macrophytes Not rooted May have welldeveloped submersed roots, or no roots Lemna - duckweed Aquatic vs. Terrestrial Aquatics mostly similar to terrestrial macrophytes One major difference - rooting tissues grow in anaerobic substratum Aquatic vs. Terrestrial Roots need O2 to respire Only can get it by transporting it from tissues in other parts of plant Extensive system of intercellular gas lacunae for gas transport, exchange Aquatic vs. Terrestrial Emergent macrophytes have leaf structure similar to terrestrial plants Linear, thick leaves no problem obtaining light, CO2 High transpiration lose lots of water Aquatic vs. Terrestrial Submersed macrophytes often look much different than terrestrials >70% of volume is intercellular lacunae Leaves very thin, divided and broadened to increase surface area to volume ratio Better absorb sunlight, CO2 Aquatic vs. Terrestrial Some submersed forms also capable of assimilating bicarbonate for use in photosynthesis Based on relative scarcity of free CO2 in most environments Nutrient Needs Most nutrients required by macrophytes come from sediments Free floaters get it from water Nutrient Needs Interstitial waters generally contain much higher concentration of nutrients than waters above sediments (anoxic conditions) Most macrophytes can assimilate nutrients from water if concentrations rise (just like phytoplankton) Leaky Macrophytes Submersed macrophytes are very leaky Lose nutrients to surrounding water during active growth Developed on land and not adapted to water? Compromise improved light, CO2 uptake at cost of losing some nutrients? Light Limitations Emergent macrophytes are seldom lightlimited - tremendous capacity for production Submersed macrophytes are light-limited Depth distribution regulated by light, in part Depth Limitations Even in systems with light penetrating to great depths (unproductive systems), macrophytes only occur down to ~10 m Results from hydrostatic pressure - doubles atmospheric pressure by 10 m Inhibits movement of gas through lacunae Macrophytes vs. Phytoplankton Phytoplankton productivity may be very low in littoral areas with many macrophytes - 3 reasons: 1) Competition for nutrients 2) Shading 3) Release of inhibitory organic chemicals by macrophytes Macrophytes vs. Algae Productivity of some types of algae may be very high in close proximity to macrophytes Grow attached to macrophytes and live off materials leaking out