Full text in pdf format
Conflicting genomes, the demictheory & biodiversity*
H.F. Hoenigsberg
Fundación Instituto de Genética-Ecológica & Biodiversidad del Tropico Americano, Cra. 4 No. 71-69, ap. 203, Santa Fé de Bogotá D.C., Colombia. E-mail: hoenigsb@colciencias.gov.co
ABSTRACT
Organismic-centered Darwinism, in order to use direct phenotypes to measure natural selection's effect, necessitates genome harmony and uniform coherence plus large population sizes. However, modern gene-centered Darwinism has found new interpretations to data that speak of genomic incoherence and disharmony. As a result of these two conflicting positions a conceptual crisis in Biology has arisen. My position is that the presence of small, even pocket-size, demes is instrumental in generating divergence and phenotypic crisis. Moreover, the presence of parasitic genomes as in acanthocephalan worms, which even manipulate suicidal behavior in their hosts; segregation distorters that change meiosis and Mendelian ratios; selfish genes and selfish whole chromosomes, such as the case of B-chromosomes in grasshoppers; P-elements in Drosophila; driving Y-chromosomes that manipulate sex ratios making males more frequent, as in Hamilton's X-linked drive; male strategists and outlaw genes, are eloquent examples of the presence of real conflicting genomes and of a non-uniform phenotypic coherence and genome harmony. Thus, we are proposing that overall incoherence and disharmony generate disorder but also more biodiversity and creativeness. Finally, if genes can manipulate natural selection, they can multiply mutations or undesirable characteristics and even lethal or detrimental ones, hence the accumulation of genetic loads. Outlaw genes can change what is adaptively convenient even in the direction of the trait that is away from the optimum. The optimum can be "negotiated" among the variants, not only because pleiotropic effects demand it, but also, in some cases, because selfish, outlaw, P-elements or extended phenotypic manipulation require it. With organismic Darwinism the genome in the population and in the individual was thought to act harmoniously without conflicts, and genotypes were thought to march towards greater adaptability. Modern Darwinism has a gene-centered vision in which genes, as natural selection's objects can move in dissonance in the direction which benefits their multiplication. Thus, we have greater opportunities for genomes in permanent conflict.
Keywords: genomes; demictheory; biodiversity.
REFERENCES
Alexander, R.D. and Borgia, G. (1978). Group selection altruism, and the levels of organization of life. Ann. Rev. Ecol. Syst. 9: 449-474.
Bartz, S.H. (1979). Evolution of sociality in termites. Proc. Natl. Acad. Sci. USA 76: 5764-5768.
Bartz, S.H. (1980). Correlation to evolution of eusociality in termites. Proc. Natl. Acad. Sci. USA 77: 3070.
Beatty, R.A. and Gluecksohn-Waelsh, S. (1972). The Genetics of the Spermatozoon. Department of Genetics of the University, Edinburgh.
Bethel, W.M. and Holmes, J.C. (1973). Altered evasive behavior and responses to light in amphipods harboring Acanthocephalan cystacanths. J. Parasitol. 59: 945-956.
Bethel, W.M. and Holmes, J.C. (1974). Correlation of development of altered evasive behavior in Gamarus lacustris (Amphipoda) harboring cystacanths of Polymorphos paradoxus (Acanthocephala) with the infectivity to the definitive host. J. Parasitol. 60: 272-274.
Bethel, W.M. and Holmes, J.C. (1977). Increased vulnerability of amphipods to predation owing to altered behaviour induced by larval acanthocephalans. Can. J. Zool. 55: 110-115.
Brahmachary, R.L. (1995). Altruism of selfish DNA and evolution. Curr. Sci. 68: 1081.
Campos, H.A. and Hoenigsberg, H.F. (1995). Fluctuating sexual characters in Drosophila pseudoobscura from Colombia. Curr. Sci. 68: 1084.
Clark, J.B. and Kidwell, M. (1995). Molecular biology and evolution of P-transposable elements in Drosophila. Evol. Biol. 8/9: 1-36.
Clemente, M., Remis, Ma. I., Vilardi, J.C. and Alberti, A. Influence of supernumerary heterochromatin on chiasma condition and abnormal sperm formation: Intra and interpopulation analysis (in press).
Cronin, H. (1994). The Ant and the Peacock. 3rd edn. Cambridge University Press, Cambridge, pp. 490.
Crow, J.F. (1979). Genes that violate Mendel's rules. Sci. Am. 240: 104-113.
Darwin, C. (1845). Journal of Research into the Natural History and Geology of the Countries Visited During the Voyage of H.M.S. Beagle Round the World, under the command of Capt. Fitz Roy, R.N. John Murray, London; new edition with a biographical introduction by G.T. Bettany, Ward, Lock. London, 1891.
Darwin, C. (1859). On the Origin of Species by Means of Natural Selection or the Preservation of Favoured Races in the Struggle for Life. John Murray, London; facsimile reproduction with an introduction by Ernst Mayr, Atheneum, New York, 1967.
Darwin, C. (1862a). On the Various Contrivances by which British and Foreign Orchids are Fertilised by Insects. John Murray, London. 2nd edn., 1877.
Darwin, C. (1862b). On the two forms, or dimorphic condition, in the species of Primula and on their remarkable sexual relations. J. Proc. Linnean Soc. (Botany) 6: 77-96. Reprinted in Barrett, 1977, pp. ii, 45-63.
Darwin, C. (1865). On the sexual relations of the three forms of Lythrum salicaria. J. Proc. Linnean Soc. (Botany) 8: 169-196; reprinted in Barrett, 1977, pp. ii, 106-131.
Darwin, C. (1868). The Variation of Animals and Plants Under Domestication. John Murray, London. 2nd edn., 1875.
Darwin, C. (1871). The Descent of Man and Selection in Relation to Sex. John Murray, London. 2nd edn., 1874; facsimile reproduction of first edition with an introduction by John Tyler Bonner and Robert M. May, Princeton University Press, Princeton, 1981.
Darwin, C. (1872). The Expression of the Emotions in Man and Animals. John Murray, London; facsimile reproduction with an introduction by Konrad Lorenz, University of Chicago Press, Chicago, 1965.
Darwin, C. (1876a). The Effects of Cross and Self Fertilization in the Vegetable Kingdom. John Murray, London, 2nd edn., 1878.
Darwin, C. (1876b). Sexual selection in relation to monkeys.Nature 15: 18-19; reprinted in Barrett, 1977, pp. ii, 207-211.
Darwin, C. (1877). The Different Forms of Flowers and Plants of the Same Species. John Murray, London, 2nd edn., 1892.
Darwin, C. and Wallace, A.R.W. (1858). On the tendency of species to form varieties, and on the perpetuation of varieties and species by natural means of selection. J. Linnean Soc. Lond. (Zoology) 3: 45-62. Reprinted by the Linnean Society (1908), pp. 87-107.
Dawkins, R. (1976). The Selfish Gene. Granada Publishing, 1979, London, pp. 224.
Dawkins, R. (1978). Reply to Fix and Greene. Contemp. Sociol. 7: 709-712.
Dawkins, R. (1979). Twelve misunderstandings of kin selection. Z. Tierpsychol. 51: 184-200.
Dawkins, R. (1980). Good strategy or evolutionarily stable strategy? In: Sociobiology: Beyond Nature/Nurture? (Barlow, G.W. and Silverberg, J., eds.). pp. 331-367. AAAS Selected Symposium 35. Westview Press, Inc. Boulder, Colorado. pp. 627.
Dawkins, R. (1981). In defense of selfish genes. Philosophy 56: 556-573.
Dawkins, R. (1982a). The Extended Phenotype: The Gene as the Unit of Selection. W.H. Freeman, Oxford, pp. 307.
Dawkins, R. (1982b). The necessity of Darwinism. N. Sci. 94: 130-132.
Dawkins, R. (1989). The evolution of evolvability. In: Langton, 1989, pp. 201-220.
Dawkins, R. (1990). Parasites, desideratum lists and the paradox of the organism. In: Keymer and Read, 1990, pp. 263-273.
Desmond, A. (1994). Huxley: The Devil's Disciple. Michael Josep, London, pp. 475.
Dobzhansky, Th. (1937). Genetics and the Origin of Species. 3rd edn. Columbia University Press, New York, 1951, pp. 364.
Dobzhansky, Th. (1940). Speciation as a stage in evolutionary divergence. Am. Nat. 74: 312-321.
Dobzhansky, Th. (1956). What is an adaptive trait? Am. Nat. 90: 337-347.
Dobzhansky, Th. (1970). Genetics of the Evolutionary Process. Columbia University Press, New York and London, pp. 503.
Dobzhansky, Th. (1975). Analysis of incipient reproductive isolation within a species of Drosophila. Proc. Natl. Acad. Sci. USA 72: 3638-3641.
Dupuis, C. (1979). La Systématique Phylogénétiqué de W. Hennig (Historique, discussion, choix de références). Cah. Nat. 34: 1-69.
Fisher, R.A. (1930). The Genetical Theory of Natural Selection. Clarendon Press, Oxford, revised edition, Dover, New York, 1958, pp. 360.
Gadagkar, M. (1995). Observational study of animal behaviour. Curr. Sci. 68: 185.
Gould, S.J. and Lewontin, R.C. (1979). The spandrels of San Marco and the Panglossian paradigm a critique of the adaptationist programme. Proc. R. Soc. Lond., Series B 205: 581-598.
Haldane, J.B.S. (1932). The Causes of Evolution. Longmans, Green, London; reprinted Cornell University Press, Ithaca, 1966, pp. 300.
Hamilton, W.D. (1963). The evolution of altruistic behavior. Am. Nat. 97: 354-356.
Hamilton, W.D. (1964). The genetical evolution of social behaviour. J. Theor. Biol. 7: 1-32.
Hamilton, W.D. (1967). Extraordinary sex ratios. Science 156: 477-488.
Hamilton, W.D. (1970). Selfish and spiteful behaviour in an evolutionary model. Nature 228: 1218-1220.
Hamilton, W.D. (1972). Altruism and related phenomena, mainly in social insects. Ann. Rev. Ecol. Syst. 3: 193-232.
Hennig, W. (1966). Phylogenetic Systematics. 2nd edn. University of Illinois Press, Urbana, Illinois.
Henry, S.M. (1967). Symbiosis. Vol. II. Academic Press, New York.
Hoenigsberg, H.F. (1968). Temperature induction of phenodeviants in Drosophila. J. Genet. 60: 1-17.
Hoenigsberg, H.F. (1989). Traditional neo-Darwinism is not enough! A hierarchical demic theory of evolution. Evol. Biol. 3: 383-391.
Hoenigsberg, H.F. (1990). Replicators & pleiotropic interactors, species, demes, size & life history traits in r & k selection, some problems in biology. Evol. Biol. 4: 269-306.
Hoenigsberg, H.F. and Dobzhansky, Th. (1987). Population Genetics in the American Tropics XXVI. Colonizing attempts of Drosophila pseudoobscura in Colombia. Evol. Biol. 1: 273-295.
Hoenigsberg, H.F. and Rojas, N. (1995). Population Genetics in the American Tropics XLV. The courtship behavior of Drosophila pseudoobscura from Aguas Calientes, Colombia violates the fequency dependent principle of population genetics. Evol. Biol. 8/9: 237-273.
Hoenigsberg, H.F., Ordoñez, M., Montano, D.A. and Bustos, E. (1988). Population Genetics in the American Tropics XXXV. The pattern of genetic variation in marginal and isolated Drosophila pseudoobscura. From Colombia. Evol. Biol. 2:7-34.
Holmes, J.C. and Bethel, W.M. (1972). Modification of intermediate host behaviour by parasites. In: Canning, and Wright, 1972, pp. 123-147 (quoted by H. Cronin, 1994, pp. 62).
Huxley, J.S. (1942). Evolution: The Modern Synthesis. George Allen and Unwin., London, pp. 652.
Jones, R.N. (1985). Are B-chromosomes selfish? In: The Evolution of Genome Size (Cavallier-Smith, T., ed.). John Wiley & Sons, Ltd.
Kidwell, M., Kidwell, J.F. and Sued, J.A. (1977). Hybrid dysgenesis in Drosophila melanogaster: A syndrome of aberrant traits including mutation sterility and male recombination. Genetics 36: 813-833.
Lewontin, T.C. (1978). Adaptation. Sci. Am. 239: 156-169.
Loray, M.A., Remis, M.I. and Vilardi, J. (1991). Parallel polymorphisms for supernumerary heterochromatin in Dichrophlus elongatus (Orthoptera): Effects on recombination and fertility. Genetica 84: 155-163.
Lyttle, T.W. (1977). Experimental population genetics of meiotic drive systems. I. Pseudo-Y chromosomal drive as a means of eliminating cage populations of Drosophila melanogaster. Genetics 86: 413-445.
Mackay, T.F.C. (1986). Transposable element-induced fitness mutations in Drosophila melanogaster. Genet. Res. 48:77-87.
Mayo, O. (1990). R.A. Fisher's contribution to evolutionary theory. Evol. Biol. 4: 1-21.
Mayr, E. (1942). Systematics and the Origin of Species from the Viewpoint of a Zoologist. Columbia University Press, New York, pp. 334.
Mayr, E. (1963). Animal Species and Evolution. Harvard University Press, Cambridge, MA, pp. 797.
Mayr, E. (1974). Cladistic analysis or cladistic classification? Z. Zool. Syst. Evolutionsforsch. 12: 94-128.
Moore, J. (1984). Parasites that change the behavior of their host. Sci. Am. 250: 82-89.
Myles, T.G. and Nutting, W.L. (1988). Termite eusocial evolution: a reexamination of Bartz's hypothesis and assumptions. Q. Rev. Biol. 63: 1-23.
Nelson, G.J. and Platnick, N.J. (1980). A vicariance approach to historical biogeography. Bioscience 30: 339-343.
Ostergreen, G. (1945). Parasitic nature of extra fragment chromosomes. Bot. Notiser 2: 157-163.
Patterson, E. (1982). Cladistics. Evolution Now. In: Nature (Maynard Smith, ed.). MacMillan Press, pp.239.
Platnick, N.I. (1980). Philosophy and the transformation of cladistics. Syst. Zool. 28: 537-546.
Remis, M.I. and Vilardi, J. (1986). Meiotic behavior and dosage effect of B-chromosomes on recombination in Dichroplus elongatus (Opthoptera: Acrididae). Caryologia 39: 287-301.
Remis, M.I., Remis, M.I., Confalonieri, V.A. and Sequeira, A.S. (1993). Clinal variation of B-chromosomes in the South American grasshopper Dichroplus elongatus (Acrididae). 1st B-Chromosome Conference Miraflores de la Sierra. Madrid.
Sequeira, A.S., Confalonieri, V.A., Remis, M.I. and Remis, M.I. (1995). B-chromosome and enzyme polymorphisms in the grasshopper Dichroplus elongatus: geographical gradients that are not explained by historical factors. Evol. Biol. 8/9: 283-299.
Simpson, G.G. (1975). Recent advances in methods of phylogenetic inference. In: Phylogeny of the Primates (Luckett, W.P. and Szalay, F., eds.). Plenum, New York, pp. 3-19.
Van Valen, L. (1978). Why not to be a cladist. Evol. Theory 3: 285-299.
Vrba, E.S. and Gould, S.J. (1986). The hierarchical expansion of sorting and selection: sorting and selection cannot be equated. Paleobiology 12: 217-228.
Wallace, A.R. (1905). My Life: A Record of Events and Opinions. Chapman and Hall, London.
White, M.J.D. (1973). Animal Cytology and Evolution. 2nd edn. Cambridge University Press, Cambridge, pp. 511.
Williams, G.C. (1966). Adaptation and Natural Selection: A Critique of Some Current Evolutionary Thoughts. Princeton University Press, Princeton, New Jersey.
Wright, S. (1932). The roles of mutation, inbreeding, crossbreeding and selection in evolution. Proceedings of the Sixth Inal. Congress of Genetics, pp. 356-366.
Wright, S. (1945). Tempo and mode in evolution: A critical review. Ecology 26: 415-419.
Wright, S. (1951). Fisher and Ford on the Sewall Wright effect. Am. Sci. 39: 452-458.
*Conference presented at 41st National Congress of Genetics, September 6-9, 1995, Caxambu, MG, Brasil.