Plant Nutrients - Nitrogen and Phosphorous
Aside from O2 and CO2, there are a variety of other chemicals needed by living things. Animals, almost by definition, obtain these other chemicals along with the carbohydrates and proteins they ingest when they consume other animals or plants, therefore, animals are usually content as long as the water has enough O2 and a decent salinity. Plants, on the other hand, are more self-sufficient, and they can synthesize a wide variety of complex molecules from simple inorganic precursors. The ecological community that will develop in a body of water is thus often dependent on the suitability of the habitat for the growth of photosynthetic organisms. Exceptions to this include the deep ocean (which is dependent, however, on the growth of phytoplankton above), or headwater streams (which depend on adjacent trees for most of their organic input through leaffall), or cave streams (bat guano) and so on.
Along with sunlight and CO2, the major needs of a plant include macronutrients nitrogen and phosphorous (used for proteins, DNA, RNA, ATP, etc.) and micronutrients such as sulfur (protein), magnesium (chlorophyll), and iron (cytochromes) (this list is not comprehensive). The micronutrients may be found in very small concentrations; plants (I will use the term plants to refer to all photosynthetic organisms) are good at obtaining them even if they are in low concentration; and they are rarely a limiting factor (be sure to review your ecology to be sure that you understand the concept of limiting factor). Some micronutrients, such as sulfur (as SO4-2), magnesium (as Mg+2), calcium ( as Ca+2) and potassium (as K+) are important constituents of both seawater and freshwater, as we have seen earlier.
Phosphorous (P) and nitrogen (N) are critical to plant growth, and they (usually P, but sometimes N) are often limiting factors to plant growth. Before you object, recalling that dissolved nitrogen is common in water, remember that it is dissolved nitrogen gas, N2, which is inert and cannot be used by most plants. The exception here are the cyanobacteria, which can fix N2 in the heterocysts, which provide a local anoxic environment for the nitrogen-fixing enzymes (and bacteria in anoxic root nodules of legumes and other anoxic places in the soil). Bodies of water with a low N/P ratio are thus prone to blooms of cyanobacteria. For most plants, N must be in the form of nitrate (NO3-) or ammonia (NH3, NH4+ in water). Ammonia, of course, is the nitrogenous waste of choice for many aquatic organisms, and even more is released by bacteria breaking down dead plants and animals or other nitrogenous animal wastes such as urea. Nitrate is a product of the nitrogen cycle; the nitrogen cycle in water differs slightly from the nitrogen cycle that takes place on land (which you are probably familiar with).
On land, N2 is fixed by bacteria in the soil such as Rhizobium, Clostridium, and Azotobacter; in the water (both freshwater and marine) N2 is fixed by such cyanobacteria as Anabaena, Plectonema, and Nostoc. The reaction requires energy and proceeds as follows:
Nitrogen Fixation: 2N2 + 6H2O ---> 4 NH3 + 3O2
Ammonia, whether generated by nitrogen fixation or by the breakdown of amino acids by animals or decomposers, is toxic. As the pH of water increases, more of the ammonia exists in the water as NH4+. NH4+ is even more toxic than NH3, and the fact that it is more prevalent at higher pH leads to one of the significant differences between keeping a marine and a freshwater aquarium. A marine aquarium typically has a pH of 8.0 to 8.5; a freshwater aquarium will usually have a pH of about 7. At pH 8.0, there is far more NH4+ present, and, if too many animals are producing too much NH3, then NH4+ levels will soon become toxic. Therefore, marine tanks must be "aged", that is, stocked slowly, to allow populations of bacteria to develop to remove the ammonia. The role of these latter bacteria will be explored below.
Once produced, ammonia (NH3) is used by a variety of plants and bacteria as the source of the amino group for amino acid synthesis (another reason that freshwater tanks are more tolerant than marine tanks in regards to ammonia is the ready availability of freshwater plants which help reduce ammonia levels). The amino acid synthesis reaction also requires energy and looks like this:
Amino acid synthesis: 2NH3 + 2H2O + 4CO2 ---> 2 CH2NH2COOH + 3O2
Note that both this reaction and the preceding one release O2 into the atmosphere; photosynthesis is not the only source of O2 in the atmosphere! While amino acid synthesis does remove some ammonia from the water, much more is usually present. Another reaction, nitrification, takes ammonia and converts it to nitrite (NO2-); this reaction releases energy to the organism which carries it out:
Nitrification I: 2NH4+ + 3O2 ---> 2NO2- + 4H+ + 2H2O
In water, this reaction is carried out mostly by bacteria of the genus Nitrosomonas. Nitrite is less toxic than ammonia, but is still toxic; high levels of nitrite can kill many aquatic organisms. Fortunately, a further nitrification reaction can occur (also with a release of energy):
Nitrification II: 2NO2- + O2 ---> 2NO3-
The end product here, nitrate (NO3-), is even less toxic than nitrite, and can be used by many plants as a nitrogen source. In aquatic systems and terrestrial systems as well, this reaction is carried out by bacteria of the genus Nitrobacter. In a typical marine aquarium, nitrate may approach toxic levels, but this process takes months. In addition, a number of denitrification reactions take place and reduce nitrate levels, as does uptake by plants.
To retrace the nitrogen cycle, let us consider the marine aquarium again (Fig. 7). Ammonia levels build as animals excrete nitrogenous wastes; as they die and decompose; as food (with protein) is added; and as N2 from the atmosphere is fixed by cyanobacteria. Because of the high pH (8.0), most of the ammonia will exist as toxic NH4+. Nitrosomonas bacteria will convert the ammonia to nitrite, and Nitrobacter bacteria will convert the nitrite to nitrate, which can be utilized by plants. Denitrification will remove some of the nitrate from the water. In a freshly established marine tank with a few fish, it is not uncommon for the ammonia levels to peak, then drop as the Nitrosomonas bacteria take hold and begin to convert ammonia to nitrite. As nitrite levels build and peak, Nitrobacter populations will thrive and convert the nitrite to nitrate, reducing nitrite concentrations to near zero. Typically, it takes about one month for the bacterial populations to become established, and it is usually wise to monitor the process by daily tests of ammonia and nitrite levels. The number of organisms that can be maintained in a marine tank is usually proportional to the amount of surface area on the gravel of the aquarium available for the Nitrosomonas and Nitrobacter bacteria to attach to.
Figure 7. Diagram of the Nitrogen Cycle in water. N2 in the air is fixed by cyanobacteria and put in the form of protein, which is eaten by fish. Fish release NH3, which is taken up by heterotrophic bacteria, plants, cyanobacteria, and the bacterium Nitrosomonas. The first three reincorporate the NH3 into protein, Nitrosomonas converts it to NO2, which is taken up in turn by Nitrobacter, which gives off NO3. The NO3 is a plant nutrient and is also utilized by anaerobic bacteria, which can produce N2, NO2, or NH3.
The story with phosphorous is much simpler. Phosphorus is typically available to plants as organophosphate (PO4-3); this compound is a common weathering product of igneous rock. Other sources of phosphate are decaying animal bodies, animal wastes, bones of vertebrates, and bird guano. The last is mentioned specifically because it is such a rich source of phosphorous; in fact, huge quantities of bird guano that accumulate near nesting areas of shorebirds are often mined for fertilizer. Phosphorous trapped in the bodies of dead organisms which sink to the bottom may accumulate in bottom sediments; this phosphorous will become available again to living plants (animals do not take up phosphorous directly from the environment) when currents sweep the bottom and bring it to the surface where there is sufficient light for photosynthesis (remember the LCP?). In nature, two such currents are noteworthy, the overturn of lakes (which occurs when the water is all of the same temperature and density, more on this later), and upwellings, places where cold ocean currents meet continents and rise up from the bottom. The most famous example of an upwelling is off the coast of Peru; as the cold water reaches the surface (replacing warm surface waters blown west by prevailing winds), it brings up phosphorous, which encourages the growth of abundant algae, which in turn are appetizers for anchovies, which are then fished in great numbers by humans (for pizza topping) and birds (where do you think the guano comes from?).