Previous chapter: F. GENETICS, ECOLOGY & EVOLUTION:
Fritjof Capra (1982) describes a paradigm shift as "a profound change in thoughts, perceptions, and values that form a particular vision of reality." Examples of paradigms in Western society, caught up in the scientific revolution, are a "belief in the scientific method as the only valid approach to knowledge; the view of the universe as a mechanical system composed of elementary material building blocks; the view of life in society as a competitive struggle for existence; and the belief in unlimited material progress to be achieved through economic and technological growth."
As did nearly all Caucasian South Africans of my generation, I spent two years in compulsory, military service in South West Africa (Namibia). There was a very particular culture in the army, an attitude that I summarise by a single word - mu. In all interactions in the army, from the generals down through to the majors, lieutenants and other officers that I met, a particular mental balance prevailed. When a person neither agreed nor disagreed with a particular opinion in ordinary conversation, he would say, "Yes-no." (ja-nee) As a paradigm, it was a form of collectively recognised agreement. It meant neither yes nor no, but was a noncommittal or neutral attitude that freed one from comment or opinion and was a form of passive obedience or agreement.
Robert Pirsig (1974) presents the same yes-no concept and calls it "mu" after the Japanese equivalent. Part of our modern paradigm is what he calls the truth trap of yes-no logic. It forms part of conventional dualistic logic and the scientific method. Mu, he says, is a third logical term, not equal to yes and no, and can expand our understanding in an unrecognised direction. Mu means "no thing" and points outside the process of dualistic discrimination. Such an idea serves well in the military and a society such as the Japanese where they expect unquestioning obedience. Mu is also a humble term for it opens one to ideas without the need to form an opinion. Pirsig explains that mu simply says, no class, not one, not zero, not yes, not no.
To me mu is almost the same as a conscious meditative state - the classical "om". It is a form of non-judgmental acceptance and tolerance of a state of affairs. American politicians use a corrupted form of this idea when they refuse to have an opinion such as in comments, "We neither condemn nor condone Turkey's actions." Mu does not belong to our logical scientific paradigm in any conscious way. Without this idea formally included into our vocabulary we do not generally accept thinking of this form. This is what paradigms are all about: what other types of effective thinking and perception have we failed to recognise? How does one change another's subjective reaction to perceive, understand and accept what is being presented? In our encounter with the world, we react subjectively at every moment assigning importance to a small selection from the full vista of experience. How we think and feel forms a paradigm that depends on many factors, including our emotional state and our education.
Such paradigms can become part of a culture or commercialised for others' gain. To get a glimpse of society's paradigms, pretend for a while that you are an invisible observer, and have no intentional influence on those around you. As you begin to realise that you are not causing their subjective reactions to your presence, you will have attained some idea of mu.
In the Bhagavad-Gita we find a description of this state: "A person who neither rejoices upon achieving something pleasant not laments upon obtaining something unpleasant, who is self-intelligent, who is unbewildered, and who knows the science of God, is already situated in transcendence. Such a liberated person is not attracted to material sense pleasure but is always in trance, enjoying the pleasure within. In this way the self-realised person enjoys unlimited happiness, for he concentrates on the Supreme. An intelligent person does not take part in the source of misery, which are due to contact with the material senses. O son of Kunti, such pleasures have a beginning and an end, and so the wise man does not delight in them. Before giving up this present body, if one is able to tolerate the urges of the material senses and check the force of desire and anger, he is well situated and happy in this world. One whose happiness is within, who is active and rejoices within, and whose aim is inward is actually the perfect mystic. He is liberated in the Supreme, and ultimately he attains the Supreme."
I can put several reasons forward why ecologists or others
not discovered the idea that I develop in this book. We may call these
In a discussion of food chains, food webs and ecological pyramids Allaby stated, "In energy terms, for example, the efficiency varies widely from one kind of ecosystem to another, but on average no more than about 0.1% of the solar radiation reaching the ground surface is absorbed and used by green plants. About 0.015% (one thousandth) of the total radiation is used by herbivores, 0.0003% by carnivores, and 0.00004% (four millionths) by top carnivores." He then goes on to emphasise, "This is a very important ecological observation." Disturbance at one level will affect all the levels above it. "This is an example of positive feedback, in which a change in one part of a system has effects in another that feed back to amplify the original change. Positive feedback leads to instability and sooner or later to deterioration of the system itself." This instability only occurs in extreme cases. An example of positive feedback would be an animal getting hungrier as it gorges or an animal producing more offspring as it became more crowded - clearly not reasonable events. Negative feedback is the normal case in ecosystems, without the deterioration described by Allaby (Brewer, 1994). Feedback can lead to uncontrollable escalation or it can produce stability. Using the negative feedback mechanism, a thermostat produces stability through regulating the temperature in a house (Gleick, 1987). Nature often "responds" with this type of stabilising feedback. Often, density-dependent factors cause negative feedback. As animals become more crowded many factors impinge to inhibit further growth, so growth slows. This feedback may be as a lack of food, space, nesting sites, or as increased predation and disease or as excess waste materials. For a plant, as the density of plants increases, available light decreases and so acts as negative feedback mechanism inhibiting or slowing the rate of further growth.
Without a keen awareness of the fine balance resulting from negative feedback mechanisms, it is easier to see nature as an exploitable resource that they can manipulate according to human whim and fancy in accord with the Jewish and Christian tradition of Man having "dominion over the fish of the sea, and over the fowl of the air, and over the cattle, and over all the earth, and over every creeping thing that creepeth upon the earth" (Gen1: 26). With a shallow interpretation of this divine permission and mission there was to no need to concern oneself with the implications of one's activities. The Judaic view of human dominion over nature was reinforced by Platonic and Aristotelian views on the difference between animal and human minds. The human mind was considered a manifestation of the soul, a spiritual entity that possessed rational and creative thought, lacking in animals and plants (Gibson, 1993). This resulted in Christianity and Judaism being "anti-natural", and with humanity severed from nature emotionally and spiritually (Worster, 1994). This does not apply to other religions, beliefs and cultures such as the Taoists that have a deep understanding of our interdependence with nature. Often, pagan-like beliefs, without the use of scientific objectivity, have led to extremely complex superstitions and practises.
In the medieval times, there was little concern with progress. According to biblical teaching, humanity was in a fallen state. All possible secular knowledge was to be found with the ancients and so gained through cultural transmission. It took centuries to change this paradigm into a desire for "progress". Even in the nineteenth century, Darwin feared the social and cultural backlash to his ideas on evolution.
This qualitative uniqueness attributed to humans has dominated Western culture and still does so today. Call someone an animal and you usually insult him or her! My young son, at the age of four happily said that we are humans and animals, while his mother insists that we are only humans. I carefully explain, using physical anatomy, that we are animals and humans and that speech, and intelligence are great and obvious differences. His mother insists that I am confusing him. Her religion does not allow her to admit that we are also animals.
Those who elevate the human status on divine grounds, or the fact of speech and intellect should take heed of the warning from simpler cultures that we have traditionally looked upon as primitive and backward. A Huichol Shaman cautioned, "In ancient times, when balance was lost on the planet, a great flood came to destroy all that which was on the earth so that the world could be reborn. A similar imbalance is occurring in this generation; we have forgotten our life source, the sun, and the sacred sea, the blessed land, the sky, and all things of nature. Unless we remember quickly what our lives are about, unless we celebrate through ceremony and prayer, we will again face destruction, but this time, it will be by fire." (Halifax, 1979).
In a more modern context, Verney (1979) records the history of humanity's destruction of nature and notes: "The story of man on earth is one of competition, often conflict, not only with his own kind, but with the animal kingdom and the resources of the natural world. . . . Too often, . . . man's greed and improvidence - often his ignorance also - have accelerated, and in some cases have begun the process of decline. Man has slaughtered animals for use: he has killed some for sport. He has introduced alien species with excellent short-term intentions, and cataclysmic long-term results. He has annihilated predators to the detriment of the herds he sought to protect. He has allowed, even encouraged, over-grazing and the widespread destruction of forests and natural resources, with no thought for the future." We need to live in harmony with nature, requiring a change in our polluting and destructive lifestyles.
We need to learn from nature the necessary manners to reside upon and be captain of her ship in a civilised and natural manner.
We need to avoid the dire prediction found in the Holy Quraan: "The
of the life of this world is like this: The water We (God) poured from
heavens caused the vegetation to grow upon the earth as food for people
and animals, until the earth became full of decorations and began
look beautiful, and the people upon it thought they were its masters.
command reaches it by day and by night. And We made it like a harvest,
as if it had not flourished only the day before!" (QsXv24).
Ecologists rely very heavily on statistical techniques and the help of computers to represent mathematically many relationships within an ecosystem. Computers allow ecologists to feed measurements and equations into a program to simulate ways in which that ecosystem might respond to change. The model can provide useful guidance as to the behaviour of complex systems. Natural systems are complex systems and so cannot be easily described by mathematics. Getting an understanding of the processes of complex systems (such as ecosystems) from an isolated study of small parts is very difficult. A person studying moths in all their intricacy has a lifetime of work, but may never understand ecosystem processes unless he also studies the whole system. Today, as 150 species per day become extinct, we need to recognise that conservation will fail if we rescue only individual species from extinction. We must preserve whole habitats.
The processes of complex systems, such as nature, are difficult to describe, so that descriptive models as basic as competition in nature, have created much debate and controversy. Natural selection, a descriptive model of the evolutionary process, was not an idea derived from mathematics or the reductionist method, but from a perception of a complex process. Similarly, I did not discover the holistic principle of compatibility through mathematics, yet the process may be demonstrable in such a form. A descriptive approach provides information on the structure and operation of a community of organisms (Putman, 1994).
The niche of a creature is multidimensional and is not easily reduced to a few parameters. A niche is the organism's "profession", while the habitat is its "address" (Odum, 1953). In traditional reductionist approaches, such as mathematics used to develop niche theory, there is a need to measure parameters, so the niche concept is reduced to the relationship of an organism and its various resources (Cooper, 1993) or feeding relationships and competitors (Putman, 1994). Natural selection, however, acts upon the whole animal. If an animal's eyesight is bad and it does not see its predator, it is eliminated from the game of life. An animal uses eyesight to interact within the niche, so sight is a niche parameter. Similarly, an animal may be dependent upon another for its habitat requirements. Many flowers require bees for pollination and others require birds or bats for dispersal. The list of such relationships is long. Maize has evolved alongside humanity changing dramatically over the 5,600 years of association, losing its ability for dispersal without human assistance (Mangelsdorf, 1958). Corn pollen is as old as 80,000 years. Cobs of 5,600 years ago were tiny, only about the size of a single kernel of Peruvian flour corn! Most of the evolutionary changes have gone on through the selection by agriculturists. Modern corn fills a niche, which through close association has led to interdependence with humanity. No mathematician can reflect the niche changes that have occurred to corn.
Also, ecology is not a science with a simple linear structure: everything affects everything else (Begon et al, 1986). In other words, ecology is a holistic science that needs to be analysed using systems theory or the principles of the new science of complexity. Natural systems have a stratified order, forming multilevel structures with integrated, self-organising wholes at lower levels acting as parts of the larger whole. Within this scheme, self-organisation at the larger level influences order at the lower level (according to Capra, 1982). The holistic approach and interpretation are difficult. Putman (1994) noted, "There should perhaps be no a priori presumption of any level of organisation in nature above that of the individual." However, ecosystem patterns and regularities exist. Living creatures have constraints on structure and organisation that a scientist has to define. Putman noted "the sheer difficulty of carrying out studies that attempt to embrace the whole. The tremendous complexity of interrelationships within the community defies any attempt at definition or analysis." He says that detailed descriptions of single systems do not usually lead to the generalised principles of communities. In the reductionist approach, scientists analyse the parameters of a single species or population. With the holistic approach, they study the entire community. The aim of the holistic approach is to find "common denominators of structure or organisation." From this we arrive at sound principles of organisation and a method of rationalisation of complexity.
Over millions of years of natural selection, severe disturbances and dynamic change, as found in the mangrove swamps of estuaries, select for species able to cope with this form of drastic change. Through time such characteristics become common to the surviving species. When a new area becomes available for colonisation, species able to disperse to, persist and survive as separate systems or units, occupy the new habitat. As more species colonise the new site, they form stratified wholes. These provide microhabitats, enabling further colonisation and greater diversity. Such stratified ecological systems can disintegrate and reform again when conditions allow, as each species has its own strategy for perpetuation. Each strategy is adapted to abiotic and biotic environmental factors. Ecosystems have this dynamic quality; the process described in traditional ecology as ecological succession , (Colinvaux, 1973) or today, simply, succession (Begon et al, 1986; Brewer, 1994).
Evolution results in the adaptation of associated species to each other and the rest of the environment, so that the community shows precise adaptation to its "preferred" or natural habitat. As an example, the mangrove habitat has persisted long enough for a community of associated organisms to adapt to it. It is a good example of an ecosystem. Many species cannot live in this environment. Experiments where scientists denuded small mangrove islands of all fauna and then monitored recolonisation, showed a return to almost the same number of species as before the experiment. Species that managed to recolonise the island often differed, but their ecological roles were represented in very similar numbers: the numbers of herbivores, scavengers, detritus feeders, wood-borers, ants, predators and parasites reestablished the original pattern or trophic organisation (Putman, 1994). Some scientists said that this pattern was no different to random colonisation, but only those species that were adapted to the mangrove habitat could return so this argument seems futile. Try to plant roses or maize or introduce giraffes to a mangrove swamp and one will quickly see that colonisation is not so random. There are constraints to the possible colonisers. If it were a random process, one could collect species from nearby, different habitats and introduce them to the mangrove island and they should persist, yet most would not live a single day!
Complex systems are not discernible through the reductionist approach and so the relations pertinent to this level of organisation are not easily identified or discovered. When the first weeds colonise bare soil in a disturbed section of forest, this is the first step in a repair process similar to the healing of a wound. Species that occupy short-lived, ephemeral habitats typically have a high reproductive output, fast growth rates and the ability to disperse widely. This enables them to be the first to colonise a new site acting aspioneer communities. Colonisers have to be able to cope with the extreme or harsh environmental conditions that often occur on such sites (Brewer, 1994). Once dispersed, they may require a long dormant period until the next rains. When germinating, they may need to cope with direct sunlight, little water, few nutrients, wind-blown sand and other rigors.
Eventually, taller, longer-lived shrubs or trees displace the pioneer plants. In whole systems, an organism's role is simply that in which it is found - they have evolved that specific ecological role through their evolution within the whole ecosystem. Their role can only be considered in relation to the whole system for it is within the holistic context that they have adapted and co-evolved. Within the ecosystem, the colonising weed cannot function within any other ecological context. Natural selection of the weed plant in interaction with the other species has "forced" it into this role to survive. It is compatible with the system functioning within this role - its ecological niche within the ecosystem. Destroy the whole forest and the weeds will flourish, but this is no longer the ecosystem to which they were adapted.
Ecological succession as a hypothesis has been very important to the development of ecology. As early as 1742, the French naturalist, Buffon , noted that forest trees might succeed each other in some sort of pattern (Colinvaux, 1973). In European botanical writings, it became fashionable to classify vegetation by successions of relationships from about 1870. Henry S. Cowles (1899) contributed significantly to the development of the ecological succession model. This idea is thus a descriptive model, used to describe a process occurring within a complex ecosystem.
Cowles noted a typical succession process where sand is first colonised by dune grass species. These grasses bound and stabilised the soil, enabling the colonisation of cottonwood (Populus deltoides). Fully grown, the cottonwood is a big tree providing shade and shedding leaves that provide organic matter for the soil and trap moisture. Soil animals that feed on these leaves then become established promoting the enrichment of the soil. Next, pine seedlings blown to this shaded environment, germinate and grow under the cottonwood trees. Eventually the pine trees form woods that displace the cottonwoods. Oaks are the next tree species to follow and replace the pine trees. Over time beech-maple forest establishes itself and as the final stand of forest in this ecological succession, is called the climax community. This successional process takes centuries or millennia within the ecosystem.
This apparent integrity of whole ecosystems led to ideas of nature as a superorganism, an idea strongly promoted by Clements (1916). A common evolutionary history, according to this scheme, resulted in organisms serving the ecosystem, as do the cells and tissues serve the whole body. The Gaia hypothesis of Lovelock is founded on the same observation and Clements’s works helped inspire Smuts , stimulating his ideas Holism (Colinvaux, 1973). Immanuel Kant's ideas on teleology preceded these. All were attempts to describe the processes that govern the whole and complex system instead of employing the reductionist approach. Today they call ecological succession merely succession and the individualistic notion of Gleason (1926) is close to the traditional modern paradigm. In this view, coexistence of species results from similarities in their requirements and tolerance and to some extent chance (Begon et al, 1986). Associations are not obligate, so a single species may be found in two quite different communities. Yet, an organism's "requirements" are what it has been adapting to through natural selection. Even in an apparently random association of organisms, each species "requirements" defines and limits its role within the community. A mangrove plant has an "obligate" niche as it is specialised to unique conditions.
What we are seeing since Kant then, is the continual grappling of the idea of complexity, holism and teleology that is found in nature and the search for a way to describe it. The approach needed for this is far different to the reductionist approach and is still evolving. Science, until today, has been based upon the study and analysis of parts of the system. There is obvious value in the reductionist method of studying units of the whole, but we have to balance this with recognition of the need to identify patterns and processes that characterise the whole system. In a study of the human body for example, doctors interpret kidney function in its relationship to the whole body. A reductionist approach establishes the way kidney tubules function, a combination of structure and chemistry. However, we need a holistic approach to explain how the kidney functions within the context of ("serves") the whole body.
I hope to show through the theory of
perpetuity and compatibility , the evolutionary weave that binds
creating a strong bond within and between communities and between
and their environment. By so doing I provide a new view of ecosystem
We must appreciate the importance of ecosystem complexity. Where
replace whole natural systems by a monoculture crop, hundreds, if not
of species belonging to an intricate system are lost. Only
plants can persist in conditions of high disturbance, so if we maintain
natural areas or habitats, any attempt to re-establish the original
will lead to only pioneer forms establishing as the other successional
will have been lost! In the view of future generations this will have
a criminal act.
Ecologists derive many principles from laboratory studies, but these conditions do not fully duplicate the animal's niche within the ecosystem. They often result in mechanistic cause and effect or stimulus-response results that do not show what really occurs in nature, nor the idea of compatibility. Theodosius Dobzhansky, a Russian-born biologist and geneticist recognised this problem with laboratory studies. In his genetic studies, he performed experiments on fruit flies in the laboratory, but would not draw evolutionary conclusion until he had verified his results in wild populations in their natural environment (Eldredge, 1995). Eldredge (1995) a palaeontologist found patterns in the fossil record that are not evident from small-scale experiments of a few years. To perceive and understand the interdependence and holistic integrity of natural complex systems, we must examine nature in the field (Worster, 1994). To understand the "niche-factor" of living creatures one has to study them in their natural environment; the "classical biology" ideal of eighteenth-century biologists.
Ecologists have sometimes erroneously developed general principles from species specialised to unique roles within the ecosystem. This should not be done, as researchers’ laboratory experiments can neither accommodate an ecosystem's response and the corresponding species' behavioral response, nor the immense period of evolutionary history (of a species within an ecosystem). Commenting on this topic, Colinvaux (1973) said, " . . . we cannot rely very much on the results of simple population-growth experiments for an understanding of the processes leading to the apparent balance in nature." Smith (1974) describes one such experiment:
"Nicholson fed to a culture of blowflies containing both adults and larvae a daily quantity of beef liver, plus an ample supply of dry sugar and water for the adults. The number of adults in the cages varied with violent oscillations. When the population of adults was high, the flies laid such a vast number of eggs that the resulting larvae consumed all the food before they were large enough to pupate. As a result, no adult offspring came from the eggs laid during that period. Through natural mortality, the number of adults progressively declined, and a few eggs were laid. Eventually a point was reached where the intensity of larval competition was so reduced that some of the larvae secured sufficient food to grow to a size large enough to pupate. These larvae in turn gave rise to egg-laying adults - after a developmental period of 2 weeks. Meanwhile the population continued to decline, further reducing the intensity of larval competition and permitting an increasing number of larvae to survive. Eventually the adult population again rose to a very high level and the whole process started over again."
"Competition for limited food held this blowfly population in a state of stability and prevented any indefinite increase or decrease. But the time lag involved between the addition of egg-laying adults by way of larval survival to the declining population resulted in an alternate over- and undershooting of the equilibrium position, causing an oscillating population density."
He called this artificial situation competition. In nature this creature lays its eggs opportunistically upon carrion found through dispersal of adults and their attraction to rotting meat. Once such a food resource is found, they rapidly consume it and complete the larval cycle so that they can produce the next generation of adults. No behavioural adaptation can preserve the resource and carrion feeding represents quite a unique niche, not one from which to derive general ecological principles. The blowfly has no inherent competitive instinct, but a strong drive to perpetuate.
A study of the dungfly, Scatophaga stercoraria , in nature, shows how finely attuned this animal is to its niche. Males wait for females on the cowpats, where the female lays her eggs. Males have the option of waiting for long or short periods for females. Continual natural selection forces only one strategy as a solution. To reduce interactive costs, creating an evolutionary stable strategy (ESS), waiting times fit a random (negative exponential) distribution and each male had a similar mating success whatever waiting time (Krebs and Davies, 1987). This pattern emerges through natural selection reducing interactive costs, for when some strategy results in greater reproductive success, more progeny with this strategy enters the next generation. The proportion of individuals with this strategy increases and interactive intensity increases, leading to decreased reproductive success. Each strategy of waiting time therefore reaches a proportion where its reproductive success equals any other strategy.
Wynne-Edwards (1986) identifies three trophic industries in nature. Green plants and green microbes are primary producers, nourished by photosynthesis and inorganic nutrients. Decomposers consist of microbes and animals that break down dead organic matter and waste. Vivivores are animals and microbes that feed on living tissue and body fluids of other organisms. An important distinction is that there are "food resources which are open to damage from over exploitation and those that are not." A resource that requires conservation is a prey species eaten by a predator. Organic detritus, such as carrion that is completely consumed without detriment to future production cannot be conserved. The detritivore occupies a very specific niche, so ecologists should not develop general principles such as competition from a study of the animal, even in nature.
Far more than half the species in the animal kingdom are dependent upon food resources of the over exploitable kind (Wynne-Edwards, 1986). Creatures devouring a rotting carcass do not fall into this category. We can describe the niche and ecology of these creatures, but to say that these creatures are competing is a subjective attribute and anthropomorphic. However, to study how this species is compatible with the ecosystem in which it is found can create a new field of understanding. From a holistic perspective the key question is how this creature contributes to ecosystem processes through the role that it performs. This would require a teleological explanation of the type described by Emmanuel Kant when asking what purpose a part (such as the heart) serves in relation to the whole (the body).
Recently, researchers have run experiments dealing holistically, with fascinating results. In London, England, is the Ecotron , an experiment that attempts to match reality. It is a controlled community system. The 'biodiversity factor' is added to experimentation, in that many species are involved. Out in the American tall-grass prairie, David Tillman and associates from the university of Minnesota have been manipulating natural plots for the last 12 years. Already, increasing diversity has been found to achieve the following:
 Organisms can harness energy from sunlight more efficiently;
 They consume more carbon dioxide and they convert this more efficiently into plant tissue;
 Increased productivity (from  above);
 Prairie studies showed that complex systems were more stable in
they were less affected by extreme conditions such as drought and
from such effects more rapidly (New Scientist, No.1937, Aug. 1994)
With the domination of the concepts of "survival of the fittest" and competition, to describe natural processes, there is little chance of perceiving the process in terms of the principle of compatibility. Often where scientists note competition in nature, I can propose a different interpretation. An animal's behaviour, physiology and physical form may reflect adaptations to other life forms within the ecosystem. Coadaptation occurs when two species adapt to each other. Adaptation occurs in response to biotic and abiotic components of the environment. I will show that, following the principle of compatibility , the animal adapts to enhance ecosystem stability through improved efficiency. Organisms achieve this through the evolution of interactive efficiency so that long associations lead to less energy expended in interactions. "Fitness" achieved through natural selection can be defined as the holistic process of perpetuity and compatibility instead of competition. It is an invisible "fitness" in that it involves the energy required to maintain the association. In all interactions between life forms there is a " window of opportunity" that involves reduced energetic costs.
A question that arises, which blinds most people to the possibility of compatibility, is how two or more interacting species switches over from competitive to compatible interactions. A species in decline due to the "competitive" (interactive) effect of associated species can do better through three mechanisms:
i] Either it increases its reproductive rate (r-factor),
ii] or it increases its competitiveness (c-factor) or interactive effect upon the associated organism causing the decline,
iii] or it reduces the interactive effect (i-factor) incurred upon it by the other species through any possible mechanism. This last mechanism is the "window of opportunity" that those ecologists with the competitive paradigm miss.
I will show these three mechanisms in the simple model below. The third mechanism leads to speciation (species formation). Methods employed are numerous. Brewer (1994) describes this third mechanism as the ability to "disaffiliate - escape the interaction . . . " Disaffiliation involves the evolution of traits that enable an organism to reduce the harmful effects of interactions such as parasitism and predation. Compatibility includes this, but goes further, as its expression in nature is more diverse.
Edward O. Wilson (1992) recognises that "it is possible to proceed down the catalogue of evolved procedures known to biologists by which species avoid hybridisation and seldom see one repeated in exact detail." These all fall under the category of evolved mechanisms that reduce intraspecific interactive intensities (classical "competition").
[a] Silkworm moths (family Saturniidae) reduce the interactive effects of chemical scents released by the females, through having different times of release for these scents: 1600 to 1800, the Promethean moth, Callosamia promethea; 2200 to 0400, the polyphemus moths, Antheraea polyphemus ; 0300 to 0400, the cecropia moths, Hyalophora cecropia.
[b] Bird species of the same genus, such as the flycatcher Empidonax , occupy different habitats and are reproductively isolated through isolating mechanisms. E. minimus occupy open woods and farmland, E. alnorum occupies yalder swamps and wet thickets and E. flaviventris are found in coniferous woods and cold bogs (Wilson, 1992).
[c] Various species of the falcon, genus Falco, are similarly distributed. In the Nearctic region, F. rusticolus are found in cold ice and tundra terrain, F. peregrinus frequent taiga and woodland with many trees and F. mexicanus are found in desert and prairie. In the Palearctic region, three species of Falco subdivide and occupy a habitat similar to F. mexicanus. These are F. cherrug, F. jugger and F. biarmicus (de la Fuente, 1971).
[d] Species need not be closely related for this separation to occur. Studies in the Spanish Sahara have shown that five species of birds, which are all insect hunters living on the fringes of an oasis, do not compete with each other. They do this by having separate hunting ranges. Sand martins (Riparia riparia) search for flies and mosquitoes over water; barn swallows ( Hirundo rustica) utilise the water edges and fly high above streams and rivers; the swift (Apus apus) flies high above all; the crag martin (Hirundo obsoleta) occupies the craggy cliffs and the house martin (Delichon urbica) a niche closely associated with human dwellings. These animals live side by side in relative harmony by specialising upon a specific food resource (de la Fuente, 1971).
[e] At some critical junction in human evolution our chromosome number was reduced from the 24 pairs found in the great apes (chimpanzees & gorillas) to the 23 pairs found in humans. This was achieved through the fusion of two different chromosomes into a single no. 2 human chromosome (Strickberger, 1985). Such a change would result in instantaneous reproductive incompatibility between the mutant and the rest of the population. Hypothetically, such a mutation would be beneficial in enforcing differentiation in a promiscuous species. It could be beneficial if the environment were exerting two conflicting selective forces on a diverse population that readily interbred. If hybrids of the two races had a lower survival potential, then the change in the chromosome number would make such hybrids unviable. Differentiation of the two populations would then occur more rapidly, as there could be no gene transfer. This change in chromosome number is therefore a very significant marker in our evolution. Establishing when this occurred is important. It need not have been at the chimpanzee-bipedal ape juncture, but may have occurred at the Australopithecus-Homo habilis juncture where so much morphological variation is found.
People devoted to competition miss the third mechanism. If they constrain reproductive output and competitiveness, what can the animal do to survive in the face of competition? Darwin saw the answer. The third option is to differentiate in some way to reduce the interactive effect that is so detrimental. Geneticists such as Richard Dawkins (1983) note quite correctly, "We must not forget that natural selection is all about relative success." Instead of the confusing term "competition" one should speak of the relative intensity of interactions or of interactive intensity.
It is the intensity of interactions that determines the intensity of the selective pressure and thus the rate of evolution. If species (or some factor) "C" is driving species "A" to extinction due to the intensity of interaction, a survival strategy open to species A is to reduce the effect (relative intensity of interaction or "i-factor") of species C on A. In the physical world, competition will force this option on Species A, if it is to survive.
People viewing nature with the anthropomorphic competitive paradigm of the survival of the fittest do not see this option, yet it is fundamental to ecosystem evolution and structure! The best solution that they can arrive at is the "Tit for Tat" solution of the "Prisoner's Dilemma". Instead of opting for the tit for tat solution, which is competitive, the animal may adopt the "once bitten, twice shy" solution of the Taoist . Over time (generations, computer iterations), Species A evolves to reduce the effect of C on A. The actual means employed is as diverse as nature. Many examples can be drawn from nature! Organisms may become active at a different time of day as do bats or in a different part of the habitat, as do various antelope. It may become physically adapted to a different food type such as a larger seed, as did some of Darwin's finches. A small change that improves reproductive success rapidly increases the proportion of that type within the population.
We need to explore dominating ideas of our culture to better understand what I have just said. The process leading to compatibility is very different to the competitive Prisoner's Dilemmas of game theory, such as Tit for Tat. There are many ways to perceive and interpret a situation. A slight change of emphasis can change a whole body of thought. Instead of the reduction of interactive "costs" being selected as a survival strategy through the process of natural selection, modern ecologists have a different view. Within an ecosystem any physical, behavioural or physiological attribute or feature that reduces the relative intensity of interaction (i-factor) of the individual with its environment will evolve through natural selection. Through this reduction of interactive costs, the individual becomes more efficient and therefore more effective. Its relative fitness increases.
Researchers working in a different context recognise the significance of reducing interactive and therefore energetic costs as a fundamental part of the process of natural selection. Cohen and Stewart observed that physical systems tended to minimise their energy. This is a principle of physics. Of complexity, the same authors state, "Genetic variation arises through random mutations, and natural selection then weeds out those mutations that don't improve the organism's chances of survival, allowing mutations that do improve survival odds to replicate. This process leads to increased complexity and sophistication of organisation of living creatures; complexity is "downhill" to evolution - it's the path of least resistance." A Taoist would say "Whoever forces it spoils it. Whoever grasps it loses it" (Watts, 1975).
Richard Dawkins also discovered this principle. In economics is a term called a utility function, which means, "that which is maximised." Economic planners and social engineers have to optimise something. By reverse engineering Gould tried to find the utility function of cheetahs and other animals by studying their present form and asking why it is so. While disproving God and showing that "the quantity that is diligently optimised in every cranny of the living world is, in every case, the survival of DNA responsible for the feature you are trying to explain," he makes another significant observation. Natural selection leads to a reduction in costs through an optimisation process. "Natural selection favours a levelling out of quality in both downward and upward directions until a proper balance is struck over all parts of the body."
Wilson (1992), in describing species formation, says, "The archives of natural history are filled with other cases of species formation as a response to ecological opportunity." He reinforces this perception a few paragraphs later: "What selection force drives the herbs to larger dimensions and assembles the island forests? Evidence from many sources suggests that it is ecological opportunity afforded by the absence of conventional trees." This is an anthropically based, teleological statement for the animal must respond to some "opportunity". I could reword this as: "Most cases of species formation are the result of the animal's response to biotic selective pressures by evolving to reduce the interactive cost between it and associated animals or the environment." An animal occupies different habitats (where possible) as a response to the effects of interaction with its own species and other organisms. A species will start to diverge as it adapts to two or more environments within its range. One may say that the organism competes by finding a way to not compete or it survives by evolving ways to reduce the intensity of the interaction. Specialised animals become more specialised. Similar, associated species differentiate to occupy different parts of their habitat.
If we take herbs or weeds as an example, they are usually well specialised for wide dispersal. On arrival on a new island, this plant weed will begin to propagate and perpetuate there. As the island is likely to be heterogeneous in character, the weed will be more suited to certain terrain than others. It may prefer the shaded, moist fissures between rocks. Inevitably, the plant begins to interact with its own species and perhaps other species, as seeds germinate in the suitable habitat and attain maturity. If there is no suitable habitat, the dispersed weed perishes.
With sexual reproduction comes genetic variation, a simple biological fact. An organism in a new environment experiences less intense biotic interactive pressure, so more of the variants may survive in this economically less demanding environment and the genotypes and phenotypes diversify. Simultaneously intense abiotic forces will rapidly eliminate varieties that do not conform to the physical (abiotic) demand. Forms that do not expend energy on this adaptation will swamp those energy-costing adaptations that hark back to the organism's previous niche. (This is why most cave dwelling creatures are albinos.)
Eventually a genetic variation arises, perhaps giving the plant tougher and thinner leaves, enabling it to occupy more exposed and wind swept areas more successfully than its parent. Immediately, it can counter new environmental stresses and disperse to an unoccupied niche where it can live and perpetuate. Offspring from this variety occupy a similar habitat successfully. Natural selection maintains this fibrous, thin-leafed form, for the ancestral form is not able to occupy the new habitat of the new variety. Natural selection will then be acting upon the species, selecting for two distinct niches in the physical world and two distinct genotypes within the population. Two incompatible alleles are being selected for through natural selection.
Dobzhansky a Russian naturalist noted the importance of the principle ofreproductive isolation evolving to reduce variation and allowing the animal to meet the specific adaptive requirements of its environment (Eldredge, 1995). The corollary of this is that diversification reduces interactive costs. As I do, he saw it as a part of the evolutionary process. The windswept niche requires a different genetic makeup to the moist, shaded-fissure niche. Different alleles on the same locus are being selected for. Reproduction of the survivors in each habitat generates more variety. Hybridisation of the two forms may not be as successful as either of the varieties in their preferred habitats. Mechanisms that evolve to prevent such hybridisation will be a selective advantage in that they increase the probability of successful propagation of each form. With each generation, the forces of natural selection, wind, sun, cold, heat, and so forth, promote the differentiation of the plant types. Natural selection leads to separate, non-breeding populations and new species.
One can interpret this event as a response to ecological "opportunity", but it is a case lacking interacting species. In the more normal case, many species are interacting. Usually the animal or plant has to cope with the abiotic factors of temperature, moisture and the like and many biotic factors, such as other animals using the same food resources, and nesting sites. In such a situation of biotic interaction, the animal has three "choices" as mentioned above. It either interacts more intensely (competes aggressively), or increases its reproductive output or changes its behaviour or niche to reduce the energetic cost of interaction. The third option leads to the coexistence of associated species as with the birds of the oasis in the Spanish Sahara. As it is the least costly option, it is the most likely where form, behaviour and physiology are plastic and subject to natural selection.
On a broader scale animal associations extend across a range of habitats. Their form and behaviour reflect adaptation to the abiotic conditions of their niche and the sustained selective forces resulting from biotic interactions. Traditional ecologists look at these interactions as competitive and essentially this is true. However human competitive events have a winner and a loser, while when species interact over many generations, the winning solution is one that increases the survival potential of the individual. In these instances, a decrease in the "cost" of interactions can be an advantage to both interactors, so both win.
I have placed a mirror within a territorial bird's range and seen it spend hours in conflict with its image, smashing its beak against the mirror, ultimately to its own detriment, as it has less time for foraging and other survival activities. The intensity of interaction with the mirror is obviously a waste of energy and time and may even cause physical damage. Two responsive animals, subject to natural selection for generations will evolve interactive behaviour that is economical and recognised by both sides. The ecological solution to intense interactions is thus mechanisms that reduce the competitive effect or "cost".
Examples of coexistence are numerous. Within the family Sciuridae is the squirrel and many other species distributed all over the world (except Madagascar and Australia). In the Mojave Desert of California two species of ground squirrel coexist, the antelope squirrel (Citellus leucurus) and the Mojave squirrel (Citellus mojavensis). Scientists initially assumed that the smaller, more active antelope squirrel excluded the larger and less active relative from its territories through competition, and a higher reproductive output, but apparently they live together in their natural surroundings without interactive detriment or competition (de al. Feunte, 1971). The solution resulting in this compatibility is to be found in the nature of the animal's niches.
Both animals are diurnal omnivores that do not need much water. They have adapted to cope with the high temperatures and dry conditions typical of the desert. Antelope squirrels and the Mojave squirrel possess different physiological and behavioural adaptations to heat and drought. If the physiological mechanisms of the antelope squirrel begin to fail under extreme conditions, it resorts to a behavioural solution to stay cool. If it cannot control it's over heating by finding a refuge, it begins to produce saliva, and wiping this over its head area until it is completely wet. This allows it a period of grace to find a better refuge, at the cost of some moisture.
Mojave squirrels copes with drought by spending seven months of the year (August to March) in a state of intermittent sleep called aestivation. It sleeps for between three and five days per week during this period. Between sleeping it goes about its normal activities. Each sleep phase lasts about six hours, its temperature drops to below the surroundings, oxygen consumption decreases, breathing and heart rate slow and very little energy is used. Over 172 days, it needs only 2 oz of energy in this state and normally carries at least 4 oz of fat.
Antelope squirrel numbers are limited by the months of food shortage during the same months when the Mojave squirrel goes into its state of aestivation. From April through to July, food is plentiful and the two species coexist. During the period of insufficient food the two species cannot coexist. Aestivation is partly a response to abiotic factors but biotic interactions must have also been a selective "force" leading to the solution displayed. The Mojave squirrel has managed to reduce the "cost" of interactions with the Antelope squirrel, by reducing competitive interactions for food and appearing during times of plenty. I call this a reduction of the i-factor (interactive factor). What a solution to competition - compete better by going to sleep and not interacting, for to interact with a more efficient animal is a waste of energy! This is why "competition" is such an unsuitable term for interactions in nature. Which of these two species is the better competitor?
Where there is the possibility to reduce the cost of interactions, solutions may be found, even within a single species. The now extinct huia bird ( Heteralocha acutorostris) of New Zealand had a unique adaptation to its habitat. Huia males had a straight, stout bill, used to chisel open dead wood and green saplings in search of any insect food. Females had long slender curve bills used to probe deeper into crevices to pry out other insects. The two different adaptations allow each sex to use a different feeding niche (Wilson, 1992).
In northern Mexico is found the fish, Cichlasoma minckleyi, living in streams, ponds and canals. It coexists with several other fish species and occurs in two different forms or morphs. Papilliform morphs have slender jaws and slender teeth. Molariform morphs have heavier, stronger jaws and pebble-shaped teeth. They are not separate species as they interbreed freely and feed together on the same prey of insects, crustaceans and worms. During periods of food scarcity, the two forms reduce competition (reduced i-factor) through the molariform morph switching to snails that it can crush with its teeth. Papilliform cichlasomes cannot do this (Wilson, 1992) (p103). Because of natural selection upon interacting biotic forms, animals evolve mechanisms to reduce the energetic cost of the association by reducing the intensity of the interaction.
The implications of natural selection favouring
reduction in the relative intensity
of interactions within ecosystems are important. We can interpret the
skin colours of humanity as due to natural selection reducing the
of interaction of sunlight with the body. Geneticists often explain
as the result of shared genes and are therefore able to reduce the unit
natural selection to the gene. With the "modified energetic
Lotka-Volterra model" (MELV
show that natural selection can lead to coadaptation between
interactors due to the selection of economic factors that operate
the ecosystem at the individual level. A decrease in the
cost through natural selection is a relative economic factor. The
of coadaptations is the result of the evolution of improved economic
Just as an animal that evolves a better camouflage has an improved
potential, so the animal that evolves a more (energetically) efficient
mechanism is better adapted to its environment. Two different species
adapt to each other in this way, evolving an association that is
and reflects interdependence.
Ecosystems possess a dimension that theoreticians do not consider when pondering game theory. To lead into the effect of game theory upon modern thought we need to briefly delve into its origins. I will then explain how this mode of thought so differs from holistic thought that game theories could not evolve into the resolution of conflicts through the perpetuity-compatibility idea. This should lead us to an understanding of the impact dominant ideas have had upon the route society has taken.
A simple game theory type conflict of interests is illustrated by asking two children to share a piece of cake. The piece of cake can never be split perfectly in half, so in the sharing, both children will want to optimise their share by getting the biggest piece. They usually end protesting that the other got the bigger piece. In this dilemma, the "solution" is to let the one child cut the cake and let the second child choose his piece first. It is then in the first child's own interest to cut the cake in half as precisely as he can. In this interaction, the child anticipates what the other will do (Poundstone, 1992). Game theory provides solutions or rational outcomes to such conflict situations.
Some great mathematicians have devoted much effort to game theories. John von Neuman (1902-1957) was possibly the most brilliant of these. As a pioneer in the development of the electronic digital computer, and a mathematician working on the Manhattan Project, leading to the manufacture of the first atomic bomb by the USA, his influence was significant. He had a photographic memory, able to recite a book, such as The Tale of Two Cities, which he had read years previously! As a professor at the Princeton Institute for Advanced Study from 1933, he had an office near Albert Einstein's. A member of the institute compared the two in a strangely distinctive way: "Einstein's mind was slow and contemplative. He would think about something for years. Johnny's mind was just the opposite. It was lightning quick - stunningly fast." This memory and computational ability enabled him to create and revise computer programs as long as fifty lines of assembly language code in his head!
Von Neuman and Oskar Morgenstern studied the mathematical structure of games and published "Theory of games and economic behaviour" in 1944. Applications of this study extend into economics, politics, foreign policy, and other spheres of life (Poundstone, 1992). They hailed this book as "one of the major scientific achievements of the first half of the twentieth century" (American Mathematical Society Bulletin, in Poundstone, 1992). Economists, social scientists and military strategists recognised the important implications of game theory to their fields. Biologists use it as a tool to explain or predict animal behaviour (Maynard Smith, 1978) (Sigmund, 1993). Game theory uses precise mathematical analysis to study the logic of conflicts among humans or animals. In this context, a game is a conflict situation where they must make decisions and choices while an associate or interactor is responding and choosing as well. Usually game theory involves situations where the two players' interests are completely opposed. Von Neuman showed mathematically that in these conflict situations there is always a rational or optimal course of action. He called it the "minimax theorem". In some applications of the minimax theorem, more than two players could be involved and there could be a partial overlap of interests.
Missing from these calculations is the impact of the interaction upon the whole system, and relative fitness of the interactors (achieved through natural selection). Interactions within ecosystems are more complex than the applications of game theory because of the effects of the interactors on the system upon which they depend. Game theory seeks to figure out the optimum strategy to pursue in conflict situations, but as seen in the flight response of so many animals, there is the option of avoiding conflict. Natural selection at the level of the individual may favour strategies of decreased interaction. This is the basis of the theory of compatibility that undercuts the game theory hawk-dove and hawk-dove-bourgeois games so loved by some biologists (Maynard Smith, 1978). We need to take one of Maynard Smith's Hawk-Dove games to understand how the compatibility theory is not subject to Game theory rules.
"Game theory is about perfectly logical players interested only in winning" (Poundstone, 1992). In ecological terms, it is about survival through competition and in holistic terms, it is about perpetuity. It is essentially competitive. Interacting opponents in game theory want to win. In game theory, a strategy used is as complete as possible description of the behaviour employed to win the game, no matter what the other player does or how long the game lasts. If figuring out every outcome of a game (such as chess) was possible, each strategic move or possible pairing of strategies could be tabulated. The early moves in the game would lead to very specific outcomes, but opponents are pursuing their own strategies, constantly optimising their situation. They cannot derive solutions to such problems from probability theory of statistics.
In the children's dilemma of how to split a piece of cake, the cake-cutter can, at extremes cut it unevenly or cut it evenly. The chooser can take a large or smaller piece. The cutter knows that the chooser will take the bigger piece, not the smaller piece, so to maximise his own portion; he cuts as evenly as possible. They arrive at equilibrium, the best strategy enforced by both players' self interest. Mathematicians call such games zero-sum games. One player's loss is the other's gain. When two rational beings find that their interests oppose the other's, they can settle on a rational course of action in confidence that the other will do the same - an equilibrium enforced by self-interest and mistrust (Poundstone, 1993). Not all equilibrium states require a zero-sum condition. There are also non-zero-sum games, where the rewards of strategies are dependent upon the strategy of the interactor.
"Prisoners Dilemma" evolved from game theory in 1950 and is now a universal concept. Theorists applied it to biology, psychology, sociology, economics and law. It is fundamental to all conflicts of interest, even where the conflict is between unthinking beings. We can even explain human and animal social organisations by the prisoner's dilemma idea. Poundstone (1992) called the Prisoner's Dilemma a major philosophical and scientific issue, tied to our survival. Here we have a paradigm woven into our society without most of us having ever heard of the term "Prisoner's Dilemma".
How does the perpetuity-compatability (p-c idea) concept's resolution of conflict situations differ from game theory? The answer is that the p-c idea considers the whole environment. In this context, to avoid energy-consuming conflict, the rational (human) player evaluates the environment, and decides that, as the two interactors have to remain together as friends, leaving the whole cake to his brother is better. He should go and find another piece or forgo the pleasure of cake altogether (flee). In evolutionary (non-rational) terms, the animal's form and behaviour are "plastic". Natural selection moulds form and behaviour. Through competition it may be forced to occupy a suboptimal environment, but natural selection results in morphological, physiological or behavioural changes, so adapting this animal to its new niche. To improve survival potential a reduction of the i-factor may be necessary. This is not a solution to be found in game theory. Game theory seeks the optimal resolution of a conflict, while the p-c idea leads to the avoidance of conflict behaviour as far as possible. Also if coevolving interactors destroy their environment, they become extinct, while this is not a consideration in game theory as evident by the nuclear stockpiles around the world.
In Tit for Tat, if an animal bites, the bitten bites back, while in the "flight" strategy, the bitten become "twice shy" and adapt to reduce the interactive effect. This is not a conscious endeavour for evolution drives it, nor is it the Christian turning of the other cheek. The motto for this strategy would be "flee until free". Flight is the most common interactive strategy in nature, yet because fleeing it is cowardly, anthropomorphism is evident in an absence of any reference to this behaviour in the index of two books on animal behaviour (Apps, 1992; Krebs & Davies, 1987), and four books on ecology (Begon et al, 1986; Brewer, 1994; Colinvaux, 1973; Wells et al, 1930). Only Smith, 1974 devotes half a page to the discussion of this response. By his 1990 edition it no longer features. Such a universal response should provide a clue to an evolutionary trend. An animal does not 'measure' the interactive effects of other species, rather, the evolution of mechanisms that reduce the interactive "cost", improves its survival potential. It is a mechanism based on economics.
A study of the history of human events confirms the futility of competition in life. When the Serbians of Yugoslavia adopted the ultimate competitive strategy of war and ethnic cleansing (see earlier in this book), they were sure their nationalistic goals were right. After years of war and destruction a Serbian (1995) observes, "This whole war was for nothing" (Time, 147(4)). Peaceful means could have achieved more.
In Maynard Smith's hawk-dove game, possible interactions are hawk-hawk, dove-dove and dove hawk with the following payoffs:
|HAWK (H)||DOVE (D)|
Serious injury = -20
Victory = +10
Long contest = -3
Hawks follow no convention, but to fight with escalating intensity until they either win or perish. Doves fight conventionally, but if fighting escalates, they run away before they are injured. A population of hawk interactors is less efficient (E(H,H)=-5) than a hawk-dove interaction (E(D,H)=0). Hawk interactions of pure competition are therefore totally destructive, while a dove-hawk interaction results in no return for the dove. As Maynard Smith (1978) notes: "Hence, dove mutants would reproduce more often than hawks."
In nature intraspecific interactive fitness is a component of total fitness. Other aspects of fitness that could be important are digestive efficiency (e.g.. cattle), eyesight (e.g. eagles), stamina (e.g. pronghorn antelope) and so forth. Many animals take advantage of this fitness, proven through natural selection, and then become territorial, dominating an area of habitat. Territoriality is a selective advantage, but not in combination with the hawk strategy. Territory holders mostly display ritualised behaviour that reduces the cost of interactions. An animal therefore uses its superior fitness in a hawk fashion, by occupying a territory. Natural selection then favours the selection of dove behaviour between these territory holders (interactors). As the dove strategy is more efficient than the hawk strategy (see above), a mutant that ignores the territorial convention and acts as a hawk loses. As an added bonus, territory holders conserve their own habitats as described below!
Geneticists often bring up the issue of "cheaters". In any conventional co-operative system, a cheater benefits by not following the behavioural convention. Territoriality serves to prevent the evolution of "cheater" behaviour.
Go to next chapter: 5. ANTHROPOMORPHISM AND GAIAN