Natural Systems Theory

by Hugh M. Lewis

http://www.lewismicropublishing.com/

Chapter Fourteen

Multi-cellular Organismic-Population Systems

            The multi-cellular organization of eukaryotic cells resulted in the evolution of an entirely new level of systems organization of living systems that can be referred to as organismic-population systems. One of the key attributes of these systems is the process of the specialization of function of cells through progressive mitosis, and the developmental differentiation of organismic organization of these cells. This remains one of the unsolved mysteries of the contemporary biological sciences. It is unknown clearly how DNA structures work to control the growth and development and maturation of the individual multi-cellular organism. The process in the first few hours seems remarkably identical for all plants and animals. But the outcomes are remarkable different for all species of organism that ever existed by the final stages of gestational development. This level of articulation of living systems is thought to have emerged sometimes between 1 and .5 billion years ago, as an immedite precursor of the Cambrian Explosion of organismic life forms on earth. It represented a revolutionary saltational jump in the organization of life and in evolutionary development of living systems.

            What is quite evident is this process of the organiismic differentiation in the development of multicellular organisms that confer properties and functions on the organism as a whole individual, highly integrated, is part of a kind of living system that is little understood in terms of its control functions and pathways of articulation.

Consideration of multicellular organisms as systems brings us to the individual and, indirectly, the species-population level of the organization of living systems. We can refer to the individual level of the multi-cellular organism as the organiismic level of living systems, which are distinguished by the functional and organismic differentiation and specialization of cells to certain organ systems the result of which integration results in one or more emergent properties that are associated with the organism as an entire integrated entity, or living individual.

These emergent properties that are the consequence of systems integration of entire organ systems, with distinctive specialized tissue cells, confer adaptive advantages to the individual as well as reproductive advantages to the specific population. It comes as a profound paradox that natural selection of traits for multi-cellular organisms is on the basis of phenotypic expression of these emergent organiismic properties, probably controlled pleiotropically and polygenically, such that point mutation becomes either unproductive or counterproductive to such expression. Genetic reshuffling of haploid sex cells and sexual reproduction confer a level of genetic variability for multicellular organisms that would otherwise be missing.

Most natural selection and evolutionary changes in fact has occurred with multicellular organisms that reproduce sexually, and that evolve rapidly in terms of the phenotypic expression of the organiismic properties associated with the behavior and response patterning of individual organisms.

We can refer to species specific starter cells, which grow and colonize in patterns of developmental differentiation to create a specific individual organism, that is defined by emergent properties from organ system integration derived from the differential functioning of these cells. There is a unity of all multi-cellular organisms in that the original starter cells have a common identity. It is the ontological developmental outcomes that the distinct characteristics of species specific organisms emerges.

Sexual reproduction provided another revolutionary boost in the development of multi-cellular life forms, for this speicalized form of reproduction greatly amplified gene-reshuffling that enhanced the individual variability of complex multi-cellular organisms, conferring in general an adaptive and long term evolutionary advantage upon such popoulations that were sexually reproduced from both male and female progenitors.

The entire evolutionary direction in the rise of multicellular organisms was the movement away from rapid rates of reproduction towards increasingly differentiated and specialed modalities of organismic reproduction that conferred greater K or equilibrium to the resulting individuals and their populations of which they were a part.

Multicellular organisms are composed exclusively of eukaryote cells, with true nucleus and cellular organelles that make them larger and more complex systems than their bacteria cousins. If cells are the prototypical system in many basic respects, then multi-cellular organisms represent surely the epitome of the emergent properties possible with systems integration. Multicellular organisms are in an analytical sense nothing but bags of miscellaneous kinds of cells. These collections of cells have amazing properties--locomotion, sophisticated perception of the environment, some level of intelligence which enables the processing and response to the environment as a whole. All of these properties by which multicellular organisms are identified form the basis by which we define and identity these organisms as whole and complete. Another way of looking at this is to see that if we remove any part of such an organism, then there is surely some critical loss of functional properties.

            The cells that compose these organisms become highly specialized and differentiated in function, and perform functions that go beyond that of bacteria or single celled organisms. There has been an evolutionary trade-off in independence of function for the enhanced survivability of the functional properties that are conferred upon the organism as a whole. The internal environment of the organism provides a "metasystem" context that appears suitable for the survival and functioning of these highly specialized eukaryotic cells. The cells are adapting to this internal environment, and evolving within it, which indirectly relates to the functional properties externally exhibited by the organism as a single whole.

            It is evident that when these multicellular organisms evolve, they are not evolving so much on a point-by-point basis with standard mutations, but there is a streamlining of property traits occurring that appears to encompass simultaneously numerous genetic points of mutation and change.

The eukaryote cells that are part of an organism all differentiate from a single set of fused sex cells, and early embryionic development is remarkably similar in all multi-cellular organisms. These cells are capable of differentiating through cellular mitosis into a myriad of highly specialed types of cells.

It is the relative sophistication of a typical eukaryotic cell compared to a prototypical bacterium or prokaryote, that serves to explain the possibility of greater differentiation of functional specialization and form of eukaryotic cells. The presence of a true nucleus, of cellular organelles, of mitochondria, golgii bodies, endoplasmic reticulum, and of a lipid bilayer cell membrane with specialized protein transport and communication structures, all serve to enable to larger eukaryote to achieve greater differentiation of form. The presence of a full complement of DNA, in multiple chromosomes, which is reproduced completely with each subsequent cellular generation through mitosis, entails that the cell has the informational capacity for differentiation along a species blue-print for the entire organism.

The cells have yielded a degree of independence found and exhibited by bacteria or single-celled organisms, and trade off this independence for an increased functioning capacity in adaptive survival and reproductive success.This tradeoff confers adaptive survival for the organism as a whole, and, indirectly, for the species or specific type of organism. Most multicellular cells in fact yield their own reproductive function to specialized sex cells, and grow only for replacement purposes.

Organiismic multicellular integration of living systems above that of the single-cell represented a revolutionary step in the development of living systems upon earth, and was a significant step towards the development of complex evolutionary systems. It was a step not achieved overnight in the natural history of life on earth, and appears to have required from 1 to 1.5 billion years for the development of multicellular life forms, in increasing varieties and complexities, to take off during the Cambrian Explosion. The success of this level of living systems development can be witnessed in the fossil record that yields myriad life forms and types, and the emergence of millions of new species of multi-cellular organism. For all that is discovered in a relatively sparse fossile record, there have been that many more that probably once existed and became eventually extinct without a trace.

In all multi-cellular organiismic systems, we witness the integration of highly differentiated and specialized cells into a single organiismic whole, or individual organism, that is conferred with a set of living properties that are unique both to that individual and to its type of species. Survival of the individual cell becomes dependent entirely upon the adaptive success and reproductive success of the individual organism as a whole, and, again indirectly, upon the species or that particular type of organism.

No multi-cellular organisms occur only as a lone individual, but always occur as part of a larger population of related individuals, and it is a key design strategy of multicellular life forms of all types that there is strength in numbers, that though a few individuals may perchance perish unsuccessfully, there will remain enough of viable population balance to continue the line successfully.

Adaptive functioning for multicellular organisms therefore occurs upon two levels simultaneously, and these two levels interact with one another. We can speak of the adaptive function and biological equilibrium of the individual organism that primarily, in the context of sexual reprodution, involves the first of the two imperatives of biological systems, and then we can, for sexually reproducing species, refer to the second level of social adaptation and equilibrium, which leads to successful reproduction of both the individual and the population as a whole. These two levels of adaptation for sexually reproducing taxa clearly interact in a systems manner, providing evolutionary advantage to species specific communities. The trade-off of K-adapted species, of course, is both the problem of over-specialization and narrowing of niche profile, which is related to the second problem of the tendency towards environment circumscription due primarily to niche exploitation and relative over-population, resulting in taxon cycles of growth and collapse of entire populations and the degradative stress of the ecosystems of which these populations are a component.

Therefore, in multicellular organisms, especially those involved in sexual reproduction, evolutionary development occurs from the feedback between two levels, the organiismic level of the adaptive functioning and health of the individual organism, which is the host body of the cells it harbors, and the larger specific population of which that individual is a potential reproductive member. Sexual reproduction favors social behavior of the group, while individual health favors adaptive success of the individual, meeting both requirements in successful evolutionary development, adaptive and reproductive success.

Adaptive and reproductive feedback between the level of the individual organism and the larger population of which that organism is a part is a critical system underlying natural selection and speciation of organisms, and this has been a relatively continuous process resulting in long term cellular differentiation and the emergence of new organiismic systems and properties.

Each individual multicellular organism that is born and developed to maturity is a unique individual, unless the primary process of reproduction was by means of cloning. High levels of individual variation along almost every trait profile assures a high degree of genetic and ultimately adaptive variability for a population, which assures that some members of the population will be capable of successfully adapting to widely fluctuating environmental circumstances.

Playing the Game of Life

The Almost Closed World Hypothesis, Fitness-Selection Matrices & the Systematics of Living and Dying

I have reached the point in the development of a model of evolution that we must venture into a more complete mathematical description of the processes involved in natural selection. In general, I build a basic set of descriptive explanations that attempt to account for and describe these processes. From this, I propose to apply models derived from game theory and various mathematical theories to the understanding of natural selection in organic evolution.

At least on one level, nature, being blind to the outcomes, plays a game of chance in producing genetic variability even at the cost of adaptive fitness. This can be described as a min-max optimization strategy in which the outcomes of different selection pathways cannot be known beforehand. Of course, theoretically Evolution is a non-constant sum game, and therefore it is impossible to model in finite and general terms, but a simple matrix model is useful in understanding the fundamental process in simplified form.

Life plays a game that is always with itself, so to speak. This is a part of an almost closed world hypothesis. Selection, fitness, and evolution are all blind processes. Each time an organism reshuffles the deck and each time a population redistributes its genetic pattern, it does so largely without knowing the consequences and without really being able to consider them. The question implicitly being asked in the reshuffling act, which is the act of reproduction, is what will be the net outcome of the next round, especially in terms of its reiteration. Not being able to know outcomes, or plan strategies, life has adopted a purely chance or random min-max strategy, which, in the structure of the long run, tends always to be the strategy of optimization of gains and losses.

This game can be characterized as one that is played by an organism with itself, a "game of n-tuple evolutionary double solitaire." The analogy of a card game is a fitting one in understanding natural selection, genetics and evolution. Each kind of organism has its own way of laying out the cards, its own rules for playing the deck. Each card would represent both a set of traits (trait-fitness value) that have some kind of adaptive-reproductive bearing on the individual, and the possible outcomes of this value (trait-selection value) depending on prevailing conditions.

The attribution of intentionality structure to evolutionary process, when we speak of selection strategies, expectations of death, or even intelligence itself, is an unfortunate consequence of our own limited language. The attribution of "intentionality" is even smuggled implicitly into the term "natural selection" itself, as if nature is making some choice and selecting in some deliberate or preplanned way. This term was derived from Darwin's observation of "selection" in the process of the cultural breeding of plants and animals. We could as well call it natural election but it is more like a grand lottery system. Intentionality structure that connotes deliberate planning can only be attributed to fairly large brained animals that exhibit some degree of planning in their adaptive survival, and this is almost always in very limited and bound contexts.

The almost closed world hypothesis arises from the notion that if possible, all life forms would seek closure upon their world. Closure in this sense would be a form of static un-changeability--a form of equilibrium that is permanent. This is a natural consequence of their own world closure. Each organism, biologically, exists bound within a world of its own making. It is the world that it has inherited, and that in a fundamental sense it cannot change except through accident. For most creatures, competition with other forms of life or even other organisms, is merely a fact of life that occurs much like a reflexive response. There is little equivocation. If anxiety arises in an animal, it is a consequence of hormonal adaptations to stress and fear. A closed world leads to equilibrium and stasis within eco-systemic frameworks.

But we must emphasize an almost closed world. Though all life (except perhaps human beings) tend to seek its own closure with its world, and many more or less achieve this, if life were completely existent in closed bubbles then there would have been no evolution in the first place.

If the world were closed, each time the genetic deck is reshuffled, the outcomes would be the same--what is reflected and theoretically expected in the normal model of Hardy-Weinberg equilibrium. But there is a large element of random chance introduced into this process, and this chance reflects noise and uncertainty in the transmission process. This chance occurs on two levels, not just in chance mutations of alleles, but in the relative fitness-selection that afterwards visits upon those alleles.

It is like death, disguised as a statistician, knocking randomly upon houses in a neighborhood, on perchance to see if anybody is home. K-A fit dominant homozygous individuals stay home every third day. Heterozygous individuals can be found at home ever other day, and homozygous recessive individuals everyday.

Thus, individual organisms get to play two games of chance. The first, the game of adaptive survival, is to decide whether it should remain at home on any given day, or go out to look for food. It cannot know if death will come knocking at the doorstep, or is in the field counting heads and counting coup in the old food chain.

This game is played every day in the same way. The odds are usually not completely random, as evolution always is built upon successful foundations. Creatures, trying to close their worlds just like their ancestors tried to do, inherit patterns that are largely automatic or habitual. They are nocturnal or crepuscular, they restrict activities to certain zones and avoid others; they eat only fruit or grains or leaves from trees, so on and so forth.

In this game, death will eventually visit, whether an organism remains at home to breed or goes out to feed. The question is when, and how soon. Less fit individuals will be chosen earlier than more fit individuals, on average. This means that unfit embryos with weak hearts do not even get out the door. It means that those old individuals that survive countless days represent the peak of their generation's outdated fitness curve. We might say that the relative equilibrium and adaptational success of a population can be measured by the number and age of post-reproductive organisms it can sustain. If death comes counting before the individual organism is able to move to the second round of the game of life, then the final verdict of "non-fitness" is rung in.

Thus selection regimes that directly affect reproductive rates through removal of premature individuals that do not have "non-deleterious" traits are disruptive regimes that leads to disequilibrium. This form of selection I have called non-differential selection.

If an organism makes it to the second round, the reproductive round, the individual actually gets to reshuffle the cards its been playing with all the time up until that point. It does so at this stage in a sexual manner, mostly. Thus it must mate with some sexual counterpart.

I do not want to get sidetracked on the issue of mate choice and patterns of sexual selection, which itself, as choice or selection implies some kind of intentionality structure. Again, life has a supreme and often sublime way of working out all these minor details.

The point I wish to emphasize here is that of the possibility of sexual reproduction itself, as a gradational capacity of evolutionary development. Sexual reproduction accomplishes an assorting and genetic accounting that is far more flexible and lends its self to much greater chance variability of pattern than asexual forms of reproduction.

Sexually reproductive creatures get to mix the deck with one another, so to speak, whereas asexually reproductive forms of life are stuck with the same deck they are born with. This basic gradational evolutionary trait complex lends great credibility to the notion that life tends to maximize its variability of genetic potential in each round of the game.

Why is one allele dominant and another recessive? This can only be explained by a genetic relationship. Selection in some previous population favored the expression of the dominant allele and the repression of the recessive. Dominance-recessiveness for any given trait complex or set of traits is therefore genetically controlled. We can talk hypothetically of a set of genetic controlling traits that act as pattern determination devices that govern the phenotypic expression of the total trait-configuration of an individual.

Thus, all traits may not be genetically or functionally equal in either the ontogenetic or the evolutionary development of an individual. Certain controlling genes may have decisive influence in the outcomes of an individual. It stands to reason therefore, that mutational patterns that are alleged to affect all genes in a random and more-or-less equality of opportunity manner, may have uneven and unequal consequences depending on the genes that they do affect.

An outcome of this would be that while most mutations might be relatively neutral and non-deleterious, most such mutations may also be rather unproductive and non-positive as well. These forms of genetic changes would occur on average most of the time, and lead to a storing of both a great fitness load and a great adaptational hoard of hidden variability within a reproductively successful population.

At the same time, it can be expected that the chance mutation of key-genes might most often result in deleterious trait configurations, pushing these kinds of variations out to the tails of the normal population curve. But it is also remotely possible that selection and mutation favoring certain variant key or controlling genes might have a revolutionary adaptive significance for its successful organism. A consequence of this might be a switching of dominance-recessiveness of pattern between alleles, among other kinds of trait-outcomes.

A principle rule of the evolutionary structure of the long run is something like Murphy's law: If something can happen, it eventually will happen.

Thus, in the second round, the organism reshuffles the deck it plays with, most often always exchanging half its cards with other members. In the second round, it gets to reshuffle the cards as many times as is allowed for that species. Each time the cards are reshuffled, a new deck is created and assigned to a new organism. Once the parent organism shuffles as many times as possible, its game is essentially over, and it returns to playing its own cards on a daily basis.

The challenge of the second round is not only to produce as many new decks as possible but to produce decks that are as substantial and as robust as possible without being able to know beforehand what these may be. In an absolute sense, we can say that an r-deck will tend to be smaller and more limited in choices than a K-deck. Of course, there are trade-offs. Producing large robust decks means making fewer overall decks, and producing many small decks precludes the possibility of making hefty decks. If one or two new cards are added at each second round of the game, then decks gradually grow in size with successive generations. This sense of allocation leads to consideration of an important and probably long neglected facet of life. The total number of cards, or instances of cards, allowed to any one organism to play with in the second round are probably more or less the same for all organisms, whether big or small, simple or complex. There is an inherent, or approximate, conservation of cards being played with per organism, per generation, on average. But the relational values of the cards must change considerably.

Of course, we need to reiterate in this analogy the almost closed world of the organism. Already, the sexual exchange of cards represents a kind of openness not otherwise available to organisms. It creates a world more open than otherwise, but still not unbounded in an absolute sense.

There is a paradox in this such that creatures can be seen to behave in ways that lead to the opening of their worlds, through maximizing and increasing trait variability. At the same time, they also function normally in ways that lead to the closing of their worlds, through trait fitness and adaptive equilibrium, or maximizing trait-predictability and hence reduction of variability. I believe this is the min-max aspect of the game of life.

Creatures create a world that is as open as possible for their offspring, and then the offspring attempt to close this open world as much as possible back upon themselves until their turn to play the second round comes up.

In what ways is the world open for a creature? In terms of the trait-card game, we may say that the relative openness of the game is an uncertainty value that is based on the fact that other creatures make choices that influence the outcome of the organism's game. Thus it is an imperfect game of solitaire, polluted by the choices that other organisms may make in the course of playing their own games that limit the choices an organism may make in any turn of the game. In each turn, competitive interactions with other creatures, only if indirect, set constraints on the number and kinds of cards an organism may get to play. The cards at this stage are not exchanged in interaction. The choices made between and by organisms are mostly random and blind.

At this stage, the game becomes a kind of poker or gambling game in which each player's deal determines some finite outcome for all players, even if the game is played in a solitary way. Each player has chips that are controlled by the cards. The chips are the expression of fitness and survival, the outcome values. These chips are exchanged in the course of the play, and there may be a central pool of chips that are taken from at each turn by each player. If a player loses all its chips before it gets to the second round, it is out of the game. It is negatively selected.

Attributions of structural pattern or of even implicit strategic intentionality to indirect selection processes again bespeaks a kind of fallacy of misplaced intentionality. Indirect selection patterns are to be understood as natural responses to a range of circumstances that are themselves largely undetermined products of chance and probability. Selection in a direct form is a completely random process. If it occurs as the result of indirect patterning, we can say that the process becomes probabilistically or stochastically nonrandom of occurrence. A non-random pattern of indirect selection therefore is a semi-determined system.

We might say, as Fisher said in relation to an ideal form of optimal adaptation that always tracks environmental change, that on average, over the long run, any selection outcome and any adaptational change at a particular point is expected to result in a 50/50 chance of outcome.

Furthermore, I believe that any systematic model of natural selection must explain it as a feedback process operating on two levels simultaneously. A model based only on normal population genetics describes only one level of this patterning. And it describes even that level inadequately as it by definition factors out the exceptions by favoring the normal curve.

In this framework, we can see that the cycle can be interrupted at almost any point, the significance of which is the death of the organism. This cycle can be interpreted in two ways, either from the standpoint of an individual organism and its environmental and social interaction with other organisms. It may also be construed from the standpoint of a population as a collection of organisms in interaction within the group and between other groups. Therefore, in terms of its representation of group dynamics, we can see that interruption of the process at any point is the equivalent of extinction.

We can say that the optimal strategy in the long run is one that seeks to minimize losses and maximize gains at every throw of the dice--losses being construed as population in terms of loss of life, and gains in terms of reproductive increase. We may construe this in terms of a simple matrix contrasting reproduction and adaptation, fitness and selection:

If we put this simple four-square table into the form of a graph along the axis of selection/fitness and reproduction/adaptation, then we get the following considerations:

            We can infer from this kind of model that there is an oscillatory cycle that defines stable min-max and central points in a pendulum pathway, as well as intermediary transitional saddle points that are inherently unstable in this state trajectory. According the first diagram of the model, inputs and outputs from without the system would tend to occur at stable nodes rather than at saddle nodes, making such a system internally resilient and coherent in relation to outside social interactions. Two other sets of inputs help to govern this process, that is environmental fluctuation, which is inherently unpredictable and therefore uncontrollable, and genetic mutation, which is also uncontrollable. These kinds of forces tend to occur at saddle transition points in the overall cycle, hence tend to be inherently disequilibriating, except that they can serve indirectly in favor of the min-max strategy by further minimizing variability by adaptational selection and maximizing reproductive variability by reproductive selection.

Each cycle can be thought of as a generational round. We can see that this generational round sets the relative evolutionary clock for each species, and the absolute rate of this clock for all species is a size-dependent relationship, such that it spins faster for smaller, shorter-lived, r-selected species, and slower for larger, longer-lived, K-selected species. This relationship is a relatively linear regression, and is related directly to the absolute size of the organism and also is based upon the presumption of an absolute constant rate of expected mutation, in the large.

We can see in this model an important intrinsic difference between r- and K-selected populations and individuals. K-selected groups will have populations that are by definition more age-heterogeneous than r selected groups, which will tend to be more age-homogeneous. The standard deviation and variance of age of a population profile of a K-group is expected to be much greater than the standard deviation and variance of age of a population profile of an r-group. This is in part related to the idea that in K-selected groups, larger body size on average is linearly related to longer life span and longer rates of growth. This is also related to longer lag periods between changing rates of death and birth, and also delayed reproduction and the retention of post-reproductive individuals in a population. This kind of age heterogeneity may have a consequence of minimizing the effects of non-differentiating selection upon the population.

In other words, non-differentiating negative selection that is tied to relative density independence is less likely to affect a highly age-heterogeneous group than an age-homogeneous group. Environmental fluctuations that affect rates of birth and differential rates of death in any given population in an ecosystem are less likely to adversely affect the entire K-population, rendering such effects relatively density-dependent, than an entire r-population. K-populations, by slowing down their clocks, are more evenly distributed through time, hence widening the limits of tolerance to environmental fluctuations by the population to very broad bands. If the net reproductive potential of an entire population is conserved over the long run, then fluctuating seasonal conditions would impact differentially upon the net reproductive capacity of the population as a whole. K-type organisms engaged in reproduction are probably inherently more susceptible and vulnerable to fluctuating uncertainty values in the environment than otherwise. A pregnant female is probably less able to evade predators and more susceptible to fluctuations of food supply. Such a population could not afford to invest all its reproductive potential in a single seasonal bloom, but must rely on distributive spacing over time to minimize the null effects of adverse selection.

Another way of looking at this is to see that within any K-population, individuals are more evenly distributed around the entire pathway of the cycle, hence the cycle itself is inherently, internally more coherent and balanced. If selective factors impinge at any one point in the cycle, which can always be sent as episodic and temporary, the likelihood is that relatively fewer individuals will be involved. Thus, K-populations can be said to be intrinsically more stable than r-populations.

On the other extreme of the continuum, r-selected populations must invest their total reproductive potential in a single seasonal bloom, a climactic mass reproductive episode. While a K-population is bent on a pathway that attempts to minimize the effects of adverse selection, at the cost of minimizing reproductive rates, the r-population is bent on an opposite course of maximizing the benefits of reproductive success, at the cost of maximizing selective adversity. The result is that for any given r-population, not only is the cycle more rapid, but the population is more likely to be unevenly distributed around the curve, such that the entire system is inherently fluctuating and imbalanced and occurs in periodic or episodic, rather than in a continuous, manner. Negative selective factors impinging on the curve at any particular point, are therefore inherently more likely to affect the entire group, or more members of the group, proportionately.

This brings up the question and possibility of age-selection as a form of natural selection. It is known for instance that some diseases occurring in the wild affect the reproductiveness of the female and the survivorship of the offspring. These types of diseases appear to "target" the reproductive and pre-reproductive.

But if r-populations are bent on a crash and dash, boom, bloom and doom cycle than K-populations, they are in a sense also inherently more crash-resilient than K-populations. If K-populations do crash, it spells disruption of the entire cycle, rather than just at one point or phase of the cycle, such that they will crash in proportionately more devastating ways. K-populations cannot rebound as quickly nor as completely as r-populations from a crash. Leftover members of a K-population crash are probably inherently more susceptible to negative selection factors during the post-crash period. This is true just on a statistical basis alone, because they can only reproduce fewer numbers over longer periods of time. Thus any random point-selection is liable to have greater resonance upon the remaining group. It suggests that once a species has climbed up the r-K gradient to the apex of the pyramid it occupies over a long evolutionary curve, it has no where else to go but back down to the bottom. Phyletic size increase in the fossil record might be an indicator and precursor of a pathway that is headed eventually towards extinction.

If each cycle of this model is a generational round, it also represents the reproductive lifecycle of an individual. If this round is a single year, or a single season, then all individuals of a population within this cycle can be expected to arrive at the terminus of the cycle at approximately the same time. The only way of avoiding this would be to make reproduction a continuous and random process in the population, as might be the case with some single celled organisms. If, for instance with canines, the reproductive life cycle can be placed over a decade, rather than just in a single season, then multiple breeding seasons would tend to distribute a population out over a larger spectrum of the cycle. Such intermediate cycles would still be subject to perturbations and susceptible to fluctuations, such that in any given season there may be a bloom or a bust in net reproductive success.

In reconstructing our model, two further considerations need to be taken into account. This model encompasses a single cycle and thus entails a reiterative or recursive process of oscillation. It is still defined in terms of only a single normal population, rather than in terms of an entire ecosystem encompassing multiple possible populations and it refers us back to the original problem in natural selection. We can identify selection fairly easily, but at what point does selection result into speciation.

In a simplified sense, we may state that a speciation event would be represented by significant changes in initial values of the cyclical system we have previously described, such that the profiles resulting from the articulation of the system would be minimally different as to characterize two different population matrices. This is sort of beating the bush around the issue of reproductive isolation, which alone appears insufficient as an explanation for con-specific differences.

We can understand speciation in a simple way if we assign some mostly arbitrary composite values to each of the four corners of the system, plus input and output values that occur stochastically, and we develop transformation equations regulating the transition between states. At each round, we would record the changes in initial values that have occurred. We would do this for each subsequent round. We would develop a matrix to represent the entire population, and we would perform the same sets of operations for each individual. Over time, we would compare the restart values with the initial start values.

If the differences in the primary variables are substantially altered, then there is an overall profile of change in the two decks of cards. There must be some cut-off point at which we can say that the subsequent population is substantially separate and different from the initial population, such that we can presume phyletic speciation has occurred. This can be judged perhaps by comparing the decks of cards, such that if we reshuffled two dissimilar decks together, it would yield a fresh set of decks that would be reproductively dysfunctional, minimally speaking. The resulting deck could no longer be used.

This issue relates indirectly in simple form to the second question of inter-specific interaction within ecosystems. We can see that inter-specific interaction can occur at several levels in the system, and influence the variables that are attached to the transition equations in the model, as well as the outcomes of the individual and population parameters. The transition equations have the challenge of taking these different sets of interactive variables with other species, especially, I believe, with con-specific species, into faithful account.

Species appear, over the short term at least, relatively stable and robust. Oscillatory cycles of selection can reverse the fortunes of any breeding population countless times about some normal standard of deviation. One round might push the curve in one direction, another in the opposite direction, but always to rebound again about some intermediate norm defined by the initial population parameters.

We must construe this oscillatory mechanism as something that in its state-trajectory may lead to a number of different patterned outcomes, depending on its initial starting conditions and intermediate variations. It is the alternative state-trajectories that such oscillatory mechanisms may take in the large that determines speciation pathways and evolutionary outcomes.

In one sense, the cycle above describes the pathway of a pendulum that oscillates about a "center of gravity." We can depict this as below:

This model emphasizes the periodicity of fluctuation, where there are limit lines about reproductive rate and death rates on either end about a centerline of equilibrium. This model would characterize r-selected populations more than a K-selected population, because in a more K-selected population in equilibrium, individuals would occupy all three stable nodes at the same time, and would be distributed in a balanced way in the entire system. A hypothetical r-selected population would more likely occupy only or mostly one node at any one moment.

Another way of describing the difference, I believe, is to say that an r-type system is in general uni-directional at any one time, hence inherently imbalanced, whereas a K-type system tends to be bi-directional at any one time, hence inherently balanced.

In a simple pendulum system representing extreme r-type species, mechanisms like drift, shifting balance, bottleneck and founder-effect, geographic dispersion and isolation are probably sufficient to account for speciation over the long run. But even for these r-selected populations, this model may in fact be over-simplistic, and hence the traditional mechanisms of speciation described for it may be insufficient.

For a K-type system at least, the correct state trajectory is defined by a second-order non-linear control mechanism, in which there are possible four alternative state trajectory patterns depending upon initial starting conditions. Speciation directions or pathways are therefore to be considered the result of these alternative state-trajectories.

I believe these models provide the appropriate conceptual framework for understanding evolutionary process in terms of natural selection and speciation. From these models, I propose the hypothetical construction of an analog of a form of natural intelligence that functions upon two levels simultaneously and that sufficiently describes the patterning of evolutionary process as a form of "blind strategy."

Faithful mathematical representation of longer-term speciation processes takes us to nonlinear control and nonlinear programming theory, underlying optimization theory as well as stochastic programming of minimum and maximum problems in which some of the variables are random and therefore uncontrollable. In general, it can be said that the values used in the original equations are themselves complex and composite variables that are solved by the use of differential equations and linear functions. In other words, a rendering of speciation processes in nature might be approached by means of computer programming involving functions inscribing differential equations, matrices, and vectors. To approach the problem of speciation evolution in less complex terms is to risk over-simplification. Such full model is beyond the scope of this book.

The presupposition of a min-max optimization theory suggests the influence of a relay control system that operates upon the system at minimum and maximum values, which can be interpreted as birth rates and death rates respectively, or alternatively as trait-fitness and selection. It can be expected that the cycle described above will oscillate in a normal manner about its central axis, within range of its maximum and minimum limits, but will become unstable if these limits are exceeded. At this point, disruptive selection occurs, the consequences of which would be the system adopting an unstable equilibrium pattern, resulting in a long-term state trajectory leading to outcomes reflecting either phylogenic or cladogenic speciation, or extinction.

Programming challenges suggest it is better to begin the construction of such a model from the ground up, rather than theoretically from the top down, and to build one program at a time, tying these together to create integrated larger programs that would represent ecosystems. Co-evolutionary structures of ecosystems can be depicted as interdependent coalitional and competitive network structures derivable from the multiplication of such systems in nature.

This kind of programming model is beyond the scope of this current work. Here suffice it to say that the basic variables representing fitness and selection are composite and complex, representing Eigenvalues of vector equations. Thus, at the level of trait description, we may distinguish between separate values of positive and negative fitness, as well as positive and negative selection, among other values that are part of that function. We are led as well into complex combinations of matrix, differential and integral equations. These combinations occur at multiple levels of the individual, the population and the species as a whole, which can be considered to be a continuous series of recurring and multiple populations over a definite period of time.

The conclusion to be derived from this is that there is no one equation, nor one single set of equations, which can describe all processes occurring in evolution in a sufficient manner. Equations like Hardy-Weinberg Equilibrium describe normal distributions of populations but it becomes increasingly difficult to represent change mechanisms representing differential selection and speciation, especially in a systematic way, in terms of simple linear equilibrium equations.

I have reached the conclusion that under normal circumstances most trait variability is probably neutral about a modal line and therefore non-deleterious. If any trait-change has a 50/50 chance of being deleterious or non-deleterious, then in the long run trait-changes should balance out and be largely self-canceling. Trait variability and change by itself does not necessarily lead to a reduction of fitness, and in general may be good for a population in the long run of wide-spectrum adaptation. Therefore, especially large populations might normally accommodate a heavy load of variation without suffering substantial penalization for carrying such a load. The question of load is more than offset by the savings account or reservoir of genetic flexibility that maintaining genetic variation entails. Thus as much as there may be genetic load associated with maintenance of less fit individuals in a population, there might also be a hidden hoard in doing so that is only expressed during periods of eco-systemic destabilization and disequilibrium.

Most populations probably regularly emit members of their groups, which emission is a form of selection. I will call this diffusion selection or dispersing selection and I construe it to be a complementary aspect of stabilizing or disrupting and diversifying selection. In such a manner, groups regularly explore the boundaries and beyond the boundaries of their world, in several senses. Diffusion selection is the better alternative than direct selection, as it does not represent the waste and absolute loss entailed in the latter form.

To the extent that there can be said to be an expectation of direct selection, it seems reasonable to conclude that migration would be the better alternative. To what extent this is a deliberate strategy, rather than an automatic consequence of conditions, is probably a moot point. Creatures normally explore the limits of this system by whatever means they have available to them. They do this naturally and as a matter of course, without considering the consequences of their actions.

The chances are that the more marginal or peripheral types of individuals are those that are the candidates for emission from the system. They can be said to be more r-type in their patterning. But this is not by any means a golden rule to be obeyed by all members. A great deal of this shedding process is probably purely happenstance, and may be about as random and stochastic as mutation itself.

Such processes offer several possibilities. It offers the possibility for adaptive radiation and gene flow over broader areas. It offers the possibility for reduction of load and of stabilization of a saturated system about equilibrium. In strong forms, it offers a bottlenecking process that can lead to rapid evolutionary development. It can also lead to a founder effect in a new colony. Colonization of a new area, or even just admission in a new system, can alter the balance of that system. Even if the survival rate of immigrating individuals is very low, the impact of this process of immigration can have a counter-adaptational effect within a neighboring ecosystem. It could stimulate counter-migration in the reverse direction of other displaced life forms.

This perspective offers two points of view. First, all selection is by definition in population genetics direct selection meaning direct removal of non-fit members by death. Therefore, all other forms of selection are essentially indirect and derivative patterns, of life attempting to avoid or minimize the expectation of death and simultaneously trying to achieve or maximize the expectation of reproductive survivorship. In this sense, direct fitness is the success of reproduction, and is the complement of direct selection. Therefore, varying forms of fitness represent indirect patterns of fitness adaptation that life adopts to hedge its bets for success.

From an eco-systemic point of view, varying forms of indirect fitness and selection are classifiable in one of two ways:

1. Whether the system is rising (succeeding) through successful reproduction;

2. Whether the system is falling (failing) through preclusive negative selection that affects premature individuals before reproduction.

Therefore a selection pattern has a positive form if it results in a net gain for the system, and a negative symmetrical form if it results in a net loss for the system. In this manner, diffusion selection in an extreme form that has negative consequences becomes a kind of directional, disruptive or peripheralizing selection.

Defining functional systems in terms of their consequences, and not their causes, are of course tautological, but this is admissible tautology if we are describing feedback systems that are essentially on some level partly closed, as the consequences indirectly become the causes.

Therefore, all forms of life tend to increase trait-variability as much as possible, and the most direct way of achieving this is through maintaining high rates of reproduction, even at the expense of carrying capacity, equilibrium and increased negative selection. Populations will thus become more K-selected in a focused way, while retaining a fundamental r-selection strategy that directs towards the periphery of its realm.

Excess population is normally thrown off, or cast off from a system, at regular rates defined by relative measures of differential fitness and saturation of the system. Thus, the costs of increased reproductive rates are avoided and eco-systemic stability can be maintained for the indefinite long run.

Normalizing and diversifying diffusive selection is an optimization pattern that represents an excellent means for a population to control itself, as it accomplishes two goals at the same time--that of maximizing its reproductive gain, while at the same time minimizing its world-openness. Casting off individuals achieves a state of parity within the boundaries of the normal population.

The expected rate of migration will be tied within an island model not only to a rate of extinction, but to the equilibrium of death and birth of a population. Old species do not necessarily have to go extinct to accommodate the entrance and colonization of new species in the same ecosystem.

The consequences are a net reduction of K of each species as a result of increasing species diversity, in order to maintain the net-K or saturated equilibrium of the overall system. Hence, if an area is by definition bounded, then species diversity must be inversely proportional to relative average K achieved by any one species within the system. In this sense, K is maintained not only by internal factors of equilibrium of death and birth rates, except that these are influenced by patterns of diffusion displacement or by invasion.

If relative K is somehow inversely proportional to species diversity and area, then the higher the diversity, the greater the net resistance of such a system to intrusive colonization by non-member populations. At the same time, the more critical or delicate the balance of the system at a supersaturated state, such that if and when invasive colonization does occur, it can lead to disruptive selection occurring with other competitive populations within the system.

Birth rates are always maximized under normalizing conditions, leading to maximization of post-reproductive death rates. This should be an inherently disequilibriating pattern, but it tends to be countered by adaptational patterns of an organism and other organisms in relative K states, by attempts to maintain niche-closure over the world represented by the eco-system. This is accomplished either by regular diffusion or periodic "infusion." Diffusion will serve to maintain equilibrium or increase equilibrium to a higher level, while "infusion" will serve to lower the level of equilibrium by crowding or restriction of space.

Thus, added gain in maintaining a pattern of regular diffusion is the possibility of increasing the niche territory that a breeding population can occupy, especially if back flow can occur between colonized areas. Diffusion itself, and its rate, must be a consequence of a system that is locally at carrying capacity. It is a high-pressure system that increases the likelihood of either random or selective diffusion occurring. Saturated systems approximating equilibrium can afford, indeed need, to emit fit and non-fit organisms alike if it is either to maintain equilibrium at high levels or increase these levels of equilibrium.

If diffusive selection is indeed selective, it means that nonrandom patterns will occur such that some sub-segment of the population curve are expected to become displaced out of the ecosystem to a disproportionate degree. This does not necessarily have to be the least adapted or less fit organisms, though on average it probably usually is. Maintaining random diffusive selection would be a means for a population to insure equality of death opportunity, which is a min-max strategy for reproductive optimization.

A high pressure system tends to be a healthy system as well, at least in the short term, in that it can resist the infusion of competing organisms into the area, on average. It thus maintains a positive balance or positive bank account with the outside world. It can afford to trade off part of its total value, to maintain the stability of the internal reservoir. If a population system slips into a negative net balance, it means that it becomes depressurized and susceptible to invasive infusion. It takes in outside value that tends to be chaotic if it crosses the reproductive boundary. It falls into a "debit" cycle.

Thus complex relative K states of population groupings exist in dynamic equilibrium to one another based on exchange rates between competing systems. We thus have an econometric model of evolution at the level of the ecosystem, as well. The absolute K of any population grouping would be measured by the area it effectively encompasses, or its total niche, and therefore the resources that are in its behavioral purview to control.

Relative K arises as the result of the shifting balance of control and area that results from continuous and periodic processes of diffusion and infusion between contiguous areas. Successful colonization of an invading species or group cannot but help lead to eco-systemic restriction of absolute K for all other species of the system, either directly or indirectly.

If these other alternate species do not restrict their behavioral range, their system becomes supersaturated or over-pressurized. In a fundamental sense, it becomes relatively depressurized in the sense that it occupies an area too great for its population to control, yielding adaptational advantage to other populations for invasion or expansion of control by default of restriction. It accomplishes this restriction through directional selection or extreme normalizing selection patterns. It must either increase its rate of diffusion or its rate of negative selection will follow.

It is to be expected, as a consequence of this pattern, therefore, that the opposite disequilibriating pattern of selection represents a failure to resolve the inherent paradox of an organism's min-max game. It cannot achieve maximization of reproductive gain, and it can no longer maintain relative closure upon its world. The net result is a failure to maintain relative K, and general disequilibriation of the organism's game.

In such a condition, a population can be considered to be open to invasion and displacement by an invading or colonizing out-group. Back migration of like or similar species may by reverse gene flow restabilize such a population. This is a side-benefit of continuous diffusing selection. Whether such a population gets "invaded" and displaced or replaced, remains a matter of chance.

High rates of population reproduction is a sign of healthy adaptations and positive blanket fitness, and continues in face of an implicit expectation that there will be high rates of death randomly distributed in a population due to negative selection.

Counter-adaptations by other competitive species are an expected outcome of systems in which relative K is defined by the maximization of reproduction and internal density, and by gradients of diffusion-infusion between contiguous systems. Ecosystems under stable conditions can tolerate a wide latitude of trait and behavioral variability. They can tolerate both high rates of reproduction and high rates of death without fundamental disturbance. Thus they can run in high gear at rapid rates, without fundamental disturbance. Ecosystems start off in low gear and gradually accelerate in time. In healthy systems, there is a net gain in population, and increased diversity of life.

In positive selection regimes, trait variability is amplified with increasing density and high rates of population growth. In such regimes, death is essentially non-selective and equal. Deleterious trait characteristics will lead to selection before reproduction. Great stores of trait variation, describable as alternate alleles that govern the same trait complexes accumulate in a species over time as the long-term consequence of mutation, drift, population increase, wide-spread adaptation, differential selection and gene flow, and can become hyper developed in such contexts. There is a hidden load paid for such development, as the average fitness of any single trait complex is thereby reduced for any given set of circumstances, even though the overall range of fitness is considerably broadened. At any time when circumstances prevail leading to uneven trait selection, death becomes selective and no longer equally distributed.

The development of a broad-spectrum alternate allele trait complex in a large population near equilibrium describes a form of diversifying selection that is uniformly positive (or potential equality) for all non-deleterious alleles.

Contrasted to this is a form of uniformly negative but unequal trait selection that occurs under unusual but not infrequent conditions. This form of selection I call disrupting. In general, there is high death rate and low birth rate. In particular, individuals are removed prematurely from the population, either antecedent to or consequent to partition. Death rate falls abnormally heavy or disproportionately upon individuals before they have a chance to successfully reproduce. In an equilibrium model, it can be expected that counterbalancing would remove all post-productive individuals of the population, but selection in this case, tending to be negative, is also blind.

Even if birth rates increased, selection would remove normal individuals with almost the same force as it removed those characterized by deleterious alleles. This entails that there is no direct connection between birth and death rates as implied in equilibrium theory. It also entails that selection and fitness, though interdependent and complementary processes are also fundamentally independent of one another. Selection can take out fit as well as non-fit individuals on a random basis. A sign of maladaptation of a population to an ecosystem, and a symptom of distress in ecosystems, are high rates of relatively nonrandom death coupled with decreased rates of reproduction. There is a net loss of population and population instability.

In general, conditions causing disrupting selection are any combination of environmental degradation, environmental shifts, competitive exclusion or displacement, and the loss of a foundational linkage within the food chain at a lower trophic level or within a co-evolutionary network. A condition is describable where a species is adapted to several different co-evolutionary frameworks at the same time, and as such serves as an intermediary linkage between both frameworks. Removal of such a linkage, or its disruption from interdependencies within one or another framework, can result in reverberations in other frameworks as well. In such a way, a regional or global system that has achieved super-saturation of all or most of its ecosystems is primed for mass extinction following gross systemic disequilibrium. It is a chain reaction that sets in throughout the biotic realm.

Other forms of trait-selection are describable within this framework. Balancing and stabilizing selection are differential forms of diversifying selection that favor either the center or the margins. Social selection, adaptation and counter-adaptive selection processes are "specialized" forms favoring varieties of certain allele complexes, but not necessarily at the expense of other alleles in other conditions. In general, directional selection can be seen as an incipient form of disruptive selection that tends to favor only one or a limited number of genotypic forms and that can easily lead to situations that degrade into disruptive selection. The extreme forms of disruption selection are marginalizing or peripheralizing selection, which pushes trait variability to a minimum on the tail of a normal curve, or else mass selection, which leads to extinction.

In conclusion, it is to be seen clearly that in an almost (but not entirely) closed systems, states of equilibrium, hence normal population distribution patterns of genetics, is not entirely sufficient for the explanation of why speciation occurs, and appears to have occurred as frequently as it has. "Normal" models stress a conservative view of evolutionary development that naturally assigns a slow even and gradualist tempo to evolutionary changes. Populations do little more than drift along, with an ocassional shift in balance occurring.

More "marginal" and less conventional models would view evolutionary process as occurring at the peripheries of the population, at the edge, so to speak, and generally credits much higher rates of development and speciation. Such models tend therefore to be more dynamic and less conservative concerning the forces of selection and change operating on populations.

A good compromise, born out to some extent in the fossil record, is that of punctuated equilibrium in which a normally stable population, typified perhaps by a basic and well represented type specimen that is assumed to be prevalent. Indeed, in terms of dominant-recessive alleles this sense of a dominant normal type may be embedded in the very character of genetic redistribution and expression patterns.

The pattern over the long run intermittently steps into a phase of rapid peripheral speciation, either leading to extinction of the line or the radical emergence of different types. These new types are fundamentally unlike the previous stem progenitor, or else phylogenically they lead to a new group that replaces the old group with a new level of stability and represented by a fundamentally different, but related, type specimen.

I am not sure that punctuated equilibrium is the best compromise solution that we can come up with to explain patterns in the natural history record. I think it is one possible outcome, that is the product of one kind of pathway that evolution can take. The difficulty with the fossil record is that it is inherently sketchy, made up of a few specimens, which, however they are grouped, become the type-specimens for the lines they almost exclusively represent, and many wide gulfs of missing linkages that would explain smoother transitions between types.

Diffusive selection would under most circumstances appear to be a very important if somewhat neglected mechanism underlying evolutionary process, both normal and punctuated. It suggests a kind of model in which diffusion resulting in successful colonization and adaptive radiation, can lead to saturated systems that result in reinvasion of closely related but reproductively isolated species, congenitorial or sibling or cousin species, that would be the most trophic-taxonomically similar and therefore highly competitive. This reinvasion of a newer and probably more fit species into the territories or niches of its cousins would result either in displacement and restriction, or else total replacement. If no reproductive boundary existed, the two groups would remerge and the population characteristics would become more homogenized and stabilized. But if speciation was carried to its natural conclusion, then niche invasion or even just indirect selection circumscription, can shift the balance of equilibrium in favor of the new group. This is an on-going process. If it is intermittent, it is so only in the structure of the short-run. Gaps develop in the fossil record because a lot of species and groups get lost along the way, either displaced or replaced completely.

Thus, my model of evolution construes speciation as more steady and regular than strongly punctuated models suggest, and yet as intrinsically more dynamic and diversifying than strongly normalized models imply with slow, slow rates of speciation.

Over the structure of the long run, odds that any species or group of species will become extinct or evolve may be just 50/50. This is just the same as the odds of the adaptive survival and reproductive success of any individual organism, in the structure of the long run, is liable to be just 50/50, all other things being equal. In this way, evolution is just about as blind as anything can be. In such a framework, the best strategy is one that optimizes one's random chances.

In closing this overwrought dialog, I will go back to some primes. I will state the following:

1. Maximum evolutionary rates of speciation are a function of the rates of reproduction and the relative size of the species, such that, if plotted, there is a near linear regression line in which the slope of size to reproductive rate is nearly constant.

2. Actual or achieved evolutionary rates of speciation fluctuate about an optimal line as the result of random and nonrandom selection processes that determine the net outcome. Rates can speed up or slow down within some range of standard deviation about this line.

3. The line of speciation optimization tracks the curve of speciation maximization.

4. Species act in such a way as to attempt to slow down speciation by achieving some state of homeostatic equilibrium. They attempt to achieve world closure of their system that is by definition almost closed but always partially open.

5. Species are led therefore to act in such a way, by maximizing rates of reproduction while maximizing adaptive fitness, upon which such equilibrium rests, that leads indirectly to the acceleration of their speciation, since partially open systems are, by definition of the long run, inherently unstable systems.

6. There is always an adaptive trade-off in trait-modification the net consequence of which is to induce greater adaptive fitness but at the cost of greater reproductive fitness. If a population acts to increase its reproductive rate at all costs (r), it must eventually sacrifice adaptive fitness. If a population acts to improve its adaptive fitness (K) it must eventually sacrifice its reproductive capacity.

7. This point beyond which trade-offs must be made is defined as the relative equilibrium line of the group in relation to other groups within its ecosystem. This is equal to the line of optimal reproduction.

8. Only by means of diffusion selection or random direct selection can a group increase its carrying capacity or saturation without incurring great trade--offs. The inherent biological imperative of all organisms, which comes before and is independent of the evolutionary reproductive imperative of all life, precludes for the most part the second option. Diffusion selection is the primary means a population has to maintain control over a partially open system.

9. Diffusion indirectly destabilizes systems by destabilizing neighboring systems to which they are partly attached. Ecosystems coexist and co-evolve in dynamic equilibrium with one another. To introduce changes into one system is to induce reverberating changes in other systems.

10. The very mechanisms that induce stability within the system established and controlled by a population, in the long run lead to instability of that system as the indirect result of that system's interaction with other systems. The differential functions that create instability in the system established by a population, will, under another prevailing set of conditions, cause that population to speciate more rapidly. The results of rapid speciation can be the reestablishment of some new niche equilibrium in the resulting population.

The object of this chapter has been to explain the model of natural selection in as systematic terms as possible, short of the delineation of a complex mathematical programming model, which would be a minimum construct necessary to the problem of description. Even such a sophisticated model though, could neither recount the steps of evolution that life has already taken, nor could it accurately predict the steps that life will take in the future. At best it would be an applied heuristic model to help us to understand and better explain in theoretical terms the processes that do appear to regularly occur in evolution.

It has been done with the idea of developing a more complex and sophisticated account of natural selection processes underlying evolutionary development. In general, this account leads to an understanding of natural selection in evolution as a form of natural self-organized "intelligence" that is essentially problem-solving in a blind, distributed, and stochastic sense, but problem-solving nevertheless.

I wish to complete this final chapter of the second part to emphasize the connection between biological systems and human systems, which are a subset of biological systems. We live in an evolutionary epoch that is distinguished by one single set of characteristics. These are the distinctive characteristics of the human system that has come to dominate the biosphere in many basic ways. I further address these issues in the first chapter of the third part treating human systems.

Life has been a grand biological synthesis. Its complexity and order is more than sublime when we consider that it was founded essentially on stochastic mechanisms alone. The regularity exhibited in the informational patterning of life is anything but chance, and can only be faithfully understood in terms of systems science. This is essentially no different than the grand stochastic pattern underlying the structure of the total universe itself. I will undertake to show that the same stochastic pattern of self-organizing systems can be employed in the accounting for human systems and intelligent systems in general.