Krasilov, 1997. Metaecology-14. General Scheme (Conclusion). Moral

Crises. Biosphere rhythms. Progress. General scheme.

V.A. Krasilov. Metaecology. Moscow: Paleontological Institute of the Russian Academy of Sciences, 1997. 208 pp. Part 14.

General scheme (conclusion). Morality.

Morality (conclusion). Chapter 5. ECOSYSTEM. Tandem.

Intuition tells us that two individuals are insufficient for the stable existence of a population of sexually reproducing organisms. But how many are sufficient? This depends on the degree of risk to which the population is exposed under specific conditions. If conditions are predictable, the risk is lower, and consequently there is no need to build up buffer numbers, which leads to the depletion of trophic resources and destabilises the population if the pressure upon it even temporarily relaxes. The interpretation of redundancy constitutes the principal distinction between the theory presented here and the traditional one. The latter regards population growth as an indicator of high fitness, of a species' flourishing, of its "striving" to fill all available ecological space. Rapidly reproducing species, such as mice and cockroaches, would thus have to be recognised as the most progressive. In contrast to this, ecosystem theory regards the proletarian reproductive strategy (in ancient Rome, proletarians were people considered worthless, capable only of reproducing) as a symptom of population ill-being, of a population "striving" for one thing only — to survive under conditions of permanent stress. Redundancy is precisely the indicator that helps one understand the relationship between stability and diversity. In any ecosystem there exists, determined by the general laws of development, a tendency toward the reduction of redundancy, but its realisation depends on the degree of risk to which the population is exposed during unpredictable environmental changes. If the degree of risk is high, considerable redundancy is maintained, increasing the chances of survival. Accordingly, the growth of diversity is arrested at a relatively low level. On a global scale, sharp declines in biological diversity mark the boundaries of geological eras, which also correspond to episodes of heightened tectonic and volcanic activity, sea-level fluctuations, climatic changes, and frequent geomagnetic reversals. The causal connections between these events are described by the rotational geodynamic model (see above), which also explains their regular periodicity, coinciding with galactic, solar, and orbital cycles. The significance of geological cycles for the evolution of the biosphere consists in their influence, in one way or another, upon all terrestrial and marine ecosystems, disrupting their stability. As a result, a reversal of evolutionary tendencies occurs, the buffer redundancy of populations increases, biological diversity declines, and the accumulation of dead matter relative to the biomass of biological communities increases. The sudden extinctions and the appearance of new life forms, which Darwin explained by gaps in the geological record, are in fact connected with the alternation of tendencies of coherent (in a stable system) and incoherent (in a disrupted system) evolution. The reversal of tendencies is the crisis (literally — turning point). It manifests itself first of all in a sharp, sometimes catastrophic, reduction in biological diversity — the indicator of the structural complexity of ecosystems. The general pattern is that on the eve of a crisis there occurs a certain growth of diversity, owing to the fact that the structure of biological communities becomes more open and admits the incursion of new species. In the next stage, the "tail" of rare species, lacking a buffer in the form of surplus numbers, is cut off. Finally, a full-scale crisis manifests itself as a delay of recovery processes at early or intermediate stages and the elimination of the climax stage. In the geological record, these events are preserved as successions of dominant forms and mass extinctions (although, at first glance, it may seem implausible that a species became extinct not because of low temperature, for example, or competitive struggle, but merely because it occupied a dominant position — the mechanism of abbreviated community development implies precisely this cause). The principal feature of crisis populations is that for them the decisive factor is not so much the efficiency of utilisation of the energy resources of the environment as the capacity for rapid increase in numbers, which permits escape from the "bottleneck" into which the population is driven by catastrophic impacts. The condition for rapid reproduction is most often accelerated development with precocious maturation of the reproductive system. A general tendency arises toward the abbreviation of ontogeny through the loss or merging of successive stages, as well as the elimination of the terminal stage. Such transformations apparently have a saltational character — at any rate, no traces of transitional states in the modified ontogeny are found. Saltations cannot be considered entirely random, since they reflect a certain directionality of the evolutionary process and are manifested in parallel in various evolutionary lineages. Their adaptiveness under unstable conditions consists in the acceleration of ontogeny; however, at the next turn of coherent evolution, they often acquire a new functional significance. At the genetic level, developmental acceleration creates a tendency toward reduction in genome size, principally at the expense of its slowly replicating elements, thereby initiating a shift in organismal formation processes — heterochrony — the most probable source of saltational morphological transformations. In the era of the formation of the synthetic theory of evolution, considerable effort was expended on the refutation of the theory of saltations (or, what amounts to nearly the same thing, macromutations), according to which new organismal forms arise in leaps, as a result of a single mutational event. The chief objection was that macromutations produce a non-viable phenotype, and even in the case of so-called hopeful monsters, the chances of establishment in the population are practically zero. The material for evolution, it was believed, consists of mutations producing an insignificant, sometimes barely perceptible, phenotypic effect. On their basis, genetic polymorphism is built up, partly neutral, partly subject to natural selection. Contemporary views on the genetic system compel a revision of this concept. All that was said about mutations in the synthetic theory of evolution refers primarily to structural genes encoding proteins. It is now known that structural genes, as a rule, constitute only a part of the genome, the system of which includes a large number of repetitive elements performing diverse regulatory functions and, in particular, determining the sequence of activation of structural genes, their expressiveness, and other properties. The regulatory portion of the genome is far more mobile, subject to constant changes — both those occurring in a regular manner during organismal development and those caused by various external influences, in particular the insertion of viral particles which, having established themselves in the genome, themselves act as its regulatory elements. For many classes of repetitive elements, such a symbiotic origin is assumed, and the genome of a higher organism as a whole represents a multicomponent symbiotic system, in which the activity of structural genes is determined by the entire set of complex interactions, including amplification of repetitive elements, their "transposition," insertion into another site, and other processes not yet fully understood. Highly repetitive heterochromatic regions contain few or no structural genes and appear to be ballast, but in reality play an important role in the process of cell division. In a developing organism, even a cytologically visible shedding of heterochromatic blocks may occur. The genome undergoes euchromation, and in old age is again subjected to heterochromation. Such cycles apparently repeat themselves in the history of species as well. In addition, there exists a link between the activity of genes in protein production, which is modulated by external influences, and the rate of their own replication. A reduction in transcriptional activity leads to methylation of genes and their shift into the category of late-replicating, with a subsequent tendency to dropout from the genetic system. This constitutes the feedback mechanism between ecological changes and the genome, determining the adaptive modification of the latter. Virus-like elements possess mobility within the genome and can also be transferred from organism to organism, sometimes carrying along portions of structural genes. This phenomenon has received the name of horizontal (as opposed to "vertical" sexual) gene transfer. In bacteria, the transfer of genes by plasmids passing from one bacterial cell to another serves as a mechanism of gene recombination, as it were substituting for the sexual process in its traditional forms. Thanks to this mechanism, properties useful to the bacterial population — for example, antibiotic resistance — very rapidly become universal assets. It happens that plasmids transfer genes from one bacterial species to another. Higher organisms are more or less protected from the introduction of extraneous genetic material, but in recent times many examples have been obtained of the introduction of genes carried by parasitic microorganisms into the genome of higher plants and animals. Genes carried by viruses and other microorganisms sometimes cause explosive mutagenesis — the very macromutational processes about which the opponents of the synthetic theory wrote. At the same time, the rate of spread of a mutation by the horizontal route is extraordinarily high. Practically all individuals of a population become mutants within a matter of years or even months (see "Horizontal gene transfer": Fallen Leaf Lake Conference, 1996). Thus the principal objection to saltational theory falls away — the impossibility of establishing a single macromutation through its spread by the sexual route. Moreover, horizontal gene transfer allows a better understanding of the relationships between species in biotic communities: they not only compete with one another, but also serve one another as potential donors of genetic material, as it were jointly advancing along the path of evolution. One area of application of this concept is the origin of flowering plants, which, according to palaeontological data (see the author's book "The Origin and Early Evolution of Flowering Plants," 1989), arose as a result of parallel development and the practically simultaneous attainment of the evolutionary level of angiospery by pre-flowering plants (proto-angiosperms) that already existed at the beginning of the Mesozoic Era and belong to various evolutionary lineages. The diverse pre-flowering plants, as it turns out, grew side by side in communities of a special type, which may be called arogenic, since the origin of new classes was a result of their syngenetic development. The decisive role here was played by the acceleration of development under unstable conditions, as well as interaction with insects found in the same fossil communities (pollen of pre-flowering plants has been preserved in their intestines). Phytophagous insects frequently promote the spread of gene-carrying microorganisms among plants. From the standpoint of speciation, during the incoherent stage, protection of the gene pool from the penetration — by sexual or non-sexual means — of extraneous genetic material is not a priority, since the population is more in need of expanding its genetic base for ecological expansion than of preserving its achieved fitness. Species boundaries thus become more transparent to horizontal transfer, and the range of variability considerably exceeds those limits which, under stable conditions, are imposed by stabilising selection. A vivid example of explosive speciation was described by N.N. Vorontsov and E.A. Lyapunova in rodents inhabiting the seismic zones of the Caucasus and Central Asia (N.N. Vorontsov, E.A. Lyapunova: in "Evolutionary Biology of Transient Unstable Populations", Springer, 221-244). In such localities, on a small territory, a fan of forms arises representing all theoretically possible variants of chromosomal rearrangements. Upon entry into the coherent phase, "broad" macro-polymorphic species — linneons, practically exhausting, as N.I. Vavilov showed, the potential spectrum of variability — break down into "narrow" ones — jordanons. These processes are directed by the development of ecosystems and, like the latter, have a cyclical character. The progressive historical tendency, manifesting itself against the background of cyclical changes, is expressed in the increase, from cycle to cycle, of biological diversity and of the stabilising influence of the biota upon the environment. Such progressive groups as flowering plants or mammals are not only themselves more diverse than their predecessors, but also support a higher level of diversity in the organisms consortively connected with them (in particular, insects). Man potentially possesses even greater capacities for environmental stabilisation, including the preservation of its biological diversity. In this connection, the development of reason may be regarded as a biospheric adaptation, a means of increasing stability. Moral In the years when the great Russian moralist writers poured their contempt upon the natural sciences — which, in the words of L.N. Tolstoy, had discovered the struggle for existence but could not discover love — Russian scientists were discovering the nature of lichens — representatives of two kingdoms, plants and fungi, merged into a single organism (A.S. Famintsyn), as well as the origin of cellular organelles, chloroplasts and mitochondria, from bacterial symbionts (K.S. Merezhkovsky, B.M. Kozo-Polyansky). However, Russian spirituality was more associated with the writer D.S. Merezhkovsky than with his biologist brother, and until very recently no one noticed that in the shadow of Tolstoy and Dostoevsky the Gnostic dream of making two into one had been realised: the word "symbiogenesis" was pronounced for the first time. Many years later, the American researcher Lynn Margulis substantiated the symbiogenetic origin of the eukaryotic cell, and another American, Barbara McClintock, discovered mobile genetic particles of viral origin, pointing to the symbiogenetic nature of the genome. The often-repeated thought that scientific discoveries are morally neutral but may be used by society for both good and ill amazes by its naivety and contempt for history. We have already seen how dangerous the game of science's detachment from ethical problems is. Phenomena of the external world are significant for us as elements of the metaecological system in which our spiritual life develops. A scientific discovery complements this system or destroys it. In contemporary pragmatic society, the outburst of passions provoked by Darwinian evolutionary theory may only cause bewilderment. We are readily prepared to acknowledge that orthodox Darwinists erred in many respects, and that their opponents frequently resorted to naive arguments, trusting feeling more than reason. However, one should recall that the European world at that time had not yet advanced so far along the path of pragmatism and that it was precisely the theory of evolution that, in the spiritual plane, most contributed to this advance. It was not Darwin, not the monkey question, and not even the struggle for existence that was the cause of the ineffable anxiety which evolutionary theory instilled in humanists, but rather the not fully conscious choice between the pragmatic and the spiritual which European civilisation was making at that time. All of this no longer seems relevant. In our day, pragmatism defines all spheres of life, including both social relations in the spirit of civilised competition not excluding philanthropy, and the attitude toward nature, which must be both conquered and conserved, for the benefit of man. However, the negative aspects of such an approach are already quite evident, and man, in order to avoid social cataclysms and preserve nature together with himself, will have to revise the basic ethical premises. Systems theory provides us with the sorely needed moral support in the evaluation of evolutionary events and mechanisms. According to the model set forth above, progressive development is accompanied by periodic crises, which signify a reversal of tendencies — movement backwards. We have found a general criterion — the reduction of dead matter production in the ecosystem — which allows us to avoid the cynical identification of progress with success. Victory in the struggle for existence cannot serve as a justification of the means. In war, some win at the price of enormous losses, others through skilful strategy, but only the latter determine the progress of military art. Natural selection and adaptation now appear in a different light as well. The Panglossian attitude toward evolution, in which whatever happens is for the best, compels one to see in natural selection a positive phenomenon. As the principal evolutionary mechanism, selection should improve, become ever more efficient. In reality, all higher organisms (man is no exception) possess means for mitigating selection (mutual aid, redirecting aggression, substituting symbolic confrontation for real confrontation, etc.). Selection — that is, differential survival and reproduction — means the genetic death of the culled individuals and acts as an entropic factor. Progress naturally leads to a reduction in the effectiveness of selection. The weakening of selection in human society is not a counter-natural tendency, as is often thought, but a perfectly natural progressive one. Fitness is traditionally judged by population growth. But rapidly reproducing organisms may increase in number even with very high mortality. Their contribution to entropy production is too great for them to be considered truly fit. An increase in fitness should rather be assessed by a reduction in mortality. Then we would be able, without falling into a vicious logical circle, to determine who is truly the most fit. We would also be able to choose, from among the adaptations that permit the preservation of a population, those that help to accomplish this with fewer losses. These are the progressive adaptations. At the dawn of life, nucleotide particles could penetrate as parasites into protein particles. In time, they formed a memory system permitting the restoration of their habitat — the protein organism. The genetic system continues to feed and reproduce; however, from the standpoint of the protein system, this appears as the servicing functions of transcription (reading the code) and replication (doubling the genetic material in the process of cell division). Since these processes are interconnected, changes in the environment that affect the transcription of certain genes are also reflected in replication and are thereby introduced into genetic memory (without going into the details of this in many respects still hypothetical mechanism, we note that the textbook conception of random mutations — coding errors — as the sole source of heritable changes fails to account for the systemic properties of the genome). The interaction of the nucleotide and protein systems is an example of mutual servicing, showing how in principle the relationship with the environment may be structured. The human genome is the memory of the entire preceding history of life — for millions of extinct species have left their trace in it. It is a record parallel to the geological chronicle. The history of man is a continuation of the history of the biosphere. Awareness of this will transform human society from a parasite on the body of nature into its double and helper (man saw the meaning of life in service to God, but a God ready to serve man has appeared, for servility debases, while mutual service elevates). For the creation of so productive a tandem, it is necessary, on the one hand, to increase the volume of memory (this is the task of science), and on the other, to change ethical premises. For one who has lived his life amid wild and semi-wild nature, it is difficult to agree that natural heritage can be preserved only by fencing it off from man with fines and barbed wire. There are many known cases in which the presence of man promotes an increase in biological diversity. The exclusion of man, as of any other species, impoverishes the biological community and deprives it of developmental prospects. Like any species, man may reduce biological diversity by displacing other species, or may increase it by introducing new forms of interaction, opening new ecological niches. Throughout the entire history of the organic world, progressive development has been connected with this latter possibility.

Morality In those years when great Russian writers-moralists poured their contempt on natural sciences, which, according to L. N. Tolstoy, discovered the struggle for existence but could not discover love, Russian scientists discovered the nature of lichens – representatives of two kingdoms, plants and fungi, merged into a single organism (A. S. Fomintsin), as well as the origin of cell organelles, chloroplasts and mitochondria from bacterial symbionts (K. S. Merezhkovsky, B. M. Kozo-Polyansky). However, Russian spirituality was more associated with the writer D. S. Merezhkovsky than with his biologist brother, and until recently, no one noticed that in the shadow of Tolstoy and Dostoevsky, the dream of the Gnostics to make two into one had been realized: the word "symbiogenesis" was proclaimed for the first time. Many years later, American researcher Lynn Margulis substantiated the symbiogenetic origin of the eukaryotic cell, and another American, Barbara McClintock, discovered mobile genetic elements of viral origin, indicating the symbiogenetic nature of the genome. The oft-repeated idea that scientific discoveries are morally neutral but can be used by society for both good and ill is striking in its naivety and contempt for history. We have already seen how dangerous the game of separating science from ethical problems is. Phenomena of the external world are significant for us as elements of the meta-ecological system in which our spiritual life develops. A scientific discovery either complements this system or destroys it. In modern pragmatic society, the outburst of passions caused by Darwin's theory of evolution can only cause misunderstanding. We readily admit that orthodox Darwinists were wrong in many aspects, and their opponents often resorted to naive arguments, trusting feeling more than reason. However, it should be remembered that the European world at that time had not yet progressed so far along the path of pragmatism, and it was the theory of evolution that contributed most to this progress in a spiritual sense. It was not Darwin, the monkey-question, or even the struggle for existence that caused the hard-to-define anxiety that the theory of evolution imposed on humanists, but rather the not-entirely-conscious choice between the pragmatic and the spiritual that European civilization was making at that time. All this no longer seems relevant. Nowadays, pragmatism defines all spheres of life, including social relations in the spirit of civilized competition, which does not exclude charity, as well as the attitude towards nature, which must be subjugated, and also protected, for the benefit of man. However, the negative aspects of this approach are already quite obvious, and for humanity to avoid social cataclysms, to preserve nature and itself along with it, it will have to revise its fundamental ethical attitudes. Systems theory provides us with such a necessary moral support in evaluating evolutionary events and mechanisms. According to the model presented above, progressive development is accompanied by periodic crises, which signify changes in trends, a movement backward. We have found a general criterion – the reduction of dead matter production in the ecosystem – which allows us to avoid the cynical identification of progress with success. Victory in the struggle for existence cannot justify the means. In war, some will win at the cost of huge losses, others – thanks to skillful strategy, but only the latter define the progress of the art of war. Now, natural selection and adaptation appear in a different light. A Panglossian attitude towards evolution, in which everything that happens is for the best, makes one see natural selection as a positive phenomenon. As the main evolutionary mechanism, selection must be improved, become more and more effective. In reality, all higher organisms (humans are no exception) have means to mitigate selection (mutual aid, redirection of aggression, replacement of real confrontation with symbolic, etc.). Selection, i.e., selective survival and reproduction, means the genetic death of selected individuals and acts as an entropic factor. Progress inevitably leads to a decrease in the effectiveness of selection. The weakening of selection in human society is not unnatural, as is often believed, but a completely natural progressive trend. Adaptability is traditionally judged by population growth. But rapidly reproducing organisms can increase in number even with very high mortality. Their contribution to entropy production is too great to be considered truly adapted. Increased adaptability should be assessed by reduced mortality. Then we would be able to determine, without falling into a vicious logical circle, who is truly best adapted. We would also be able to choose among the adaptations that allow the population to be preserved, those that help to do so with fewer losses. These are progressive adaptations. At the dawn of life, nucleotide particles could embed themselves as parasites into protein particles.