THE GREAT LAKES

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knowledge of the entropy flow, the total yalue of the timedependent entropy is unknown, and that means it is impossible to predict favored trajectories and outcomes. Thus, thermodynamics does not provide a general theory of evolution in far from equilibrium situations. A few systems have predictable evolutionary trajectories and outcomes no matter how far from equilibrium they are. These include 1) the isolated system (system exchanges neither energy nor materials with the surroundings); and 2) the closed and open systems (a system exchanging energy but no materials with the surroundinqs, and a system exchanging both energy and mateGials with- the surroundings, respectively) in which all the rate . processes have linear phenomenological (rate) laws. An isolated system has no entropy flow, and evolves to a state of maximum entropy whereupon it ceases to change. Ecological examples are organism death and species extinction in isolated environments. Closed and open systems with only linear rate laws evolve through a suite of predictable steady states to a final steady state of minimum free energy (not always a thermodynamic equilibrium) consistent with any external constraints on energy and material flows. An analysis of force-flux relationships in systems with linear phenomenological laws shows that both entropy flow and entropy production are positive. All final states are stable and withstand perturbations or fluctuations in various parameters. Ecological examples include autotrophic growth in a nutrient-limited environment and diffusional processes in marine plankton leading to patchy and nonpatchy biogeographic distributions. Most ecological systems of interest are open and have some nonlinear dynamics. What happens then? A unique thermodynamic equilibrium still exists. Near to equilibrium, the f orce-f lux relationships are linear, a consequence of truncating the Taylor series expansions of these relationships at the linear terms. Thus, nonlinear systems have linear dynamics and behave like linear systems: they follow the thermodynamic branch. Far from equilibrium, the f orce-f lux equations are nonlinear. Depending on the nonlinearities, the analysis may reveal one or more of the following:
1) multiple evolutionary trajectories, 2) multiple steady states, or 3) a critical point (a bifurcation) at a certain value of the thermodynamic affinity at which the evolutionary pathway can change. For systems of interest, the evolutionary pathway changes at the bifurcation from the thermodynamic branch to a new branch, leading to a new structure. That new structure, called a dissipative structure, explains why systems far from equilibrium and highly disordered can produce new and unexpected ordered structures. This is the new order out of chaos (Prigogine and Stengers 1984) studied intensively by Prigogine and his co-workers. For mathematical models of the biological systems in the required chemical analog format, Glansdorff and Prigogine (1971) provide a method to locate the bifurcation, if one exists. They examined the excess entropy function, a special form of the second-time derivative of entropy for a given system, expressed in terms of fluctuations in the force-flux relationships. Evolutionary trajectories shift at the bifurcation because the original pathway becomes unstable with respect to a perturbation or fluctuation in some factor. The study of dissipative structures is the study of the evolution of system stability, with stability here being what ecologists call resilience--the ability of a system to attenuate a disturbance. To amplify the previous ideas, note that ecosystems are open systems, and several important ecological processes have nonlinear rate laws. A prime example of a nonlinear rate law is the Lntka-Volterra model for predator-prey dynamics (Montroll 1972, May 1973) . A thermodynamic study of a mathematical model of a biological system uses Glansdorf f and Prigogine s method to extract the evolutionary trajectories and their associated steady states. A system having a single trajectory and one steady state requires no further analysis. For systems with multiple trajectories and/or possible steady states, then stability and fluctuation analyses can sometimes be used to assess the relative likelihood that a particular trajectory is favored under assumed external ,and internal conditions.
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ECO TO THE INTEGRITY OF THE GREAT L
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Support of this publication by the
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J. Edwards represented SAB; and Hen
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Theoretical Framework for Developin
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During the Symposium, it was noted
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I can but assert that the essential
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Water quality must be of the highes
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I have only one way to insure no di
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case. . . . From a pragmatic standp
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A program premised upon the establi
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Holding the Line No further degrada
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INTEGRITY AND SURPRISE IN THE GREAT
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exploitation of a scientific concep
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mitigation relates to &d culture an
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mismanagement of sewage works, and
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FOSTERING INTEGRITY IN AN AGE OF SU
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The stress-response approach, as de
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natural associations that are alrea
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Individuals that serve strong econo
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(xi) Degradation of aesthetics; (xi
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Woodwell, G.M. 1977. Biological int
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it is used in the Great Lakes Water
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adapt to perturbation, can both be
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promote this through exercise, diet
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3. For the classic dis~ssion of the
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The GWQA requires that a remedial a
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what is not good about different st
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external influence3 but less adapti
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An extended view of integrity is th
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human systems. Put simply, they awe
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as a human need and, hence, a commo
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measures, if undertaken properly, w
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question of repairing biological pr
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Legally enshrined terms such as int
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See von Bertalanffy, L. 1950. The t
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24. Holling, C.S. Ecosystem design:
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ECOSYSTEM INFEGRITY AND NETWORK THE
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might appear that autocatalysis can
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development, any increase in the pr
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ecosystem elements of interest; e.g
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Ulanowicz , R. E. 1986b. A phenomen
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the members of a society interpret
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coherent knowledge base, yielding a
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Page 89 and 90: een recognized in subatomic physics
Page 91 and 92: culture, human beings become orient
Page 93 and 94: NOTES 1. The ideal type of techniqu
Page 95 and 96: water bcdy resulted in the appearan
Page 97 and 98: 4) Different members of society wil
Page 99 and 100: total costs of ecological damage or
Page 101 and 102: As noted by Everett (1979), this ap
Page 103 and 104: The net result of these problems is
Page 105 and 106: ller, F.G. 1974. Benefit-cost analy
Page 107 and 108: IWPEGRITY AND SURPRISE IN THE GREAT
Page 109 and 110: theory as it is most often applied
Page 111 and 112: a position beyond information towar
Page 113 and 114: them. From the point of view of the
Page 115 and 116: chosen from the group. The standard
Page 117 and 118: est developed and appropriate to Gr
Page 119 and 120: potential of Ulanowiczts technical
Page 121 and 122: Kay, J.J. 1983. Self-oryanization a
Page 123 and 124: hierarchy theory. self-organizing s
Page 125 and 126: configuration. For example, gravita
Page 127 and 128: unpredictable. Middle-number system
Page 129 and 130: Integrity comes through constraint.
Page 131 and 132: Allen, T.F.H., and E.P. Wileyto. 19
Page 133 and 134: greater detail, and only the most g
Page 135 and 136: statistical thermodynamics revealed
Page 137: Force-flux relationships do not alw
Page 141 and 142: 3) reduce the total number of react
Page 143 and 144: input, removal, or accumulation, co
Page 145 and 146: eplacement, the species diversity t
Page 147 and 148: The numerical difference between a
Page 149 and 150: Certain choices of environmental va
Page 151 and 152: On the other hand, thermodynamics a
Page 153 and 154: Morowitz, H.J. 1968. Energy flow in
Page 155 and 156: 2) We must understand basic cause-a
Page 157 and 158: Fontaine and Lesht (1987) used stat
Page 159 and 160: We suggest that exercising control
Page 161 and 162: aggregates, age-class specific stoc
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Page 167 and 168: Bartell, S.M., R.H. Gardner, R.V. O
Page 169 and 170: THEORETICAL FRAMEWORK FOR DEVEIllPI
Page 171: BIOINDICATOR OBTECTIVES IN THE GREA
Page 174 and 175: the Index of Biotic Integrity (IBI)
Page 176 and 177: TABLE 2. integrity. potential metri
Page 178 and 179: of strict funding, public accountab
Page 180 and 181: The managerial advantages in such i
Page 182 and 183: more thorough and competent the exa
Page 184 and 185: organization'' (Kay 1990) . Fig. 2
Page 186 and 187: efficacy of zoobenthos as indicator
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1. Unfortunately, most studies oper
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that is, the index of response is a
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Patrick, R. 1975. Identifying integ
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George Francis Department of Enviro
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FEDERAL FH)ERAC International Joint
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navigation, hydropower generation a
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inational network of groups sharing
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professional experts suggests there
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Ecosystem integrity for the Great L
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Meisner, J.D., L. Goodier, and H.A.
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will be considered to be consistent
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For any real ecosystem, a particula
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Rutledge (1974) showed that a short
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State V&e ~ifru?cation Point State
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An example of this case is the clea
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something big happens in between sa
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When an ecosystem is described as b
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with the idea of an N-dimensional s
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TABLE 1. Environmental factors and
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TABLE 2. A tabular model of ecologi
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Another possibility is that the sta
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State Variable PI PZ P3 State Varia
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1. It is theoretically possible by
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Nicolis, G., and I. Prigogine. 1977
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AQUATIC =NIC COMMUNITIES: SURROGATE
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always retain control of the prey t
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the high levels of abundance of the
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a suite of species exemplified by t
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emergent view, accordingly, is that
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we often observe in natural systems
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ecological structures and functions
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Holling, C.S. 1985. Resilience of e
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Ryder, R.A., and S.R. Kerr. 1989. H
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MYTHS AND BDDEL8 OF SYSTEMIC AND OR
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The first two processes tend to fos
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A complementary process relates to
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that reflect key aspects of integra
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and consumer organisms. Presence of
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~ollingwood, R.G. 1946. The idea of
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Steedman, R.J. 1988. Modification a
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they will be required (by the ecosy
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6) Forbidden zone implies a state i
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3) Lack of a prenrentive .approach.
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2) Can we tell when the processes t
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Government is not wholly consistent
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Awareness of an environmental crisi
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EOOIOGY: ANALYTICAL APmcOACRES TO P
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(Fonnan and Godron 1981, 1986). Suc
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such as photosynthesis and transpir
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8) grid cell analysis: grid cell ov
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Turner 1987). Models at the individ
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Wlchwald, K. 1963. Die Industrieges
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Milne, B.T. 1988. Measuring the fra
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Vesecky, J.F., and R.H. Steward. 19
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THE GREAT LAKES

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