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<br />1 <br />j <br />l <br />! <br />\ <br />, <br />I <br />\ <br />l <br />V I <br />; I <br /> \ <br /> i <br /> 1 <br /> ! <br /> i <br /> , <br /> I <br /> I <br /> ! <br /> <br />84 <br /> <br />Nutrient Cycling (Spiraling) <br /> <br />The chemical elements necessary for life tend to ,:irculate from the <br />environment to organisms, and upon death and disintegration of the <br />organisms, are returned to the environment. Thus the term nutrient cycl ing <br />has been coined to describe the cyclic movement of inolrganic and orga<1ic <br />nutrients within ecosystems. Approximately 30 to 40 of the over 100 <br />chemical elements are known to be required by living olrganisms. Both <br />essential and nonessential (mercury, lead, ,!tc.) organic and inorganic <br />nutrients cycle. Elements such as carbon, oxygen, hydrogen, and nitrogen <br />are required by organisms in large quantitiE!s, whereas only small amounts <br />are required of some trace metals such as copper, aluminum, etc. <br />Autotrophs and heterotrophs within stream ecosystems (as in other syste<:1s) <br />can survive only if the cycling process provides a full compliment of <br />essential elements. <br /> <br />The linear configuration of stream ecosystems from headwaters to <br />estuary changes the dynamics of cycling from the conventional "closed" <br />cycles to partially "open" cycles called nutrient spiraling (Webster 1975, <br />Wallace et al. 1977, Newbold et al. 1981, 1982, Elwood et al. 1983, <br />Minshall et al. 1983). The actual recycling of materials and nutrients in <br />stream ecosystems is controlled largely by biological activities related to <br />the biota, temperature, oxygen, and basic nutrient conditions. Spiraling <br />includes both physical and biological processes, but r'~cycling is primarily <br />a biotic process (Figure 6). <br /> <br />For convenience, atoms of nutrients may be design,ated as being <br />components of either a biological pool or a nonbiologi,cal pool. As <br />nutrient atoms are released from the biological pool tl,ey enter the <br />nonbiological pool (or flowing water medium) and may thereby be displaced <br />in a downstream direction. For example, a carbon atom within a dead and <br />disintegrating bacterial cell may be releas.!d to the flowing water. Its <br /> <br />displacement in a downstream direction will depend largely on water flow <br />and, in general, the greater the flow, the greater the distance between its <br />release point and the point where the atom is recycled or incorporated into <br />the biological pool. It is the "spacial" dimension or distance traveled <br />downstream that resembles a partially open cycle and which is described as <br />spiraling. The downstream movement (displacement) is primarily a function <br />of phYSical processes. The greater the wat"r flow, the greater the <br />distance between loops of the spiral. <br /> <br />The rate of nutrient spiraling may be fast or slow. Where biological <br />activities are high, recycling rates may be fast. Where biological <br />activities are low, recycling rates may be slow. The period between <br />nutrient release and its subsequent uptake is termed the turnover time. <br />The distance traveled during:, ~hfl turnover time may be 'lhort or long and is <br />~' referred to as the turnover length. Nutrient turnover distances (length) <br />and time are determined primarily by stream current velocity and the <br />presence and effectiveness of physical instream retention devices (Minshall <br />et al. 1983). A stream's response to dissolved and particulate inputs <br />