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Livro Evolution, Coevolution And Biodiversity

Livro Evolution, Coevolution And Biodiversity

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Published by: Jitendra Mishra on Dec 21, 2011
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A
NSELM
K
RATOCHWIL
Fachgebiet Ökologie, FB Biologie/ChemieUniversität OsnabrückD-49069 Osnabrück, Germany
A
NGELIKA
S
CHWABE
 Institut für Botanik/Abt. GeobotanikTechnische Universität Darmstadt D-64287 Darmstadt, Germany
Evolution,Coevolution, andBiodiversity
Thetheoryofbiologicalevolutionisofoneofthemostimportant,centraltheoriesofbiology.Manyevolution-theoretical theses can be proven, owing to theenormous scientific progress recently made in many biological fields (e.g., paleontology, biogeography, and molecular biology). The most important evolutionary factors are mutation, recombination, and selection.Of particular relevance is speciation, which is usually due to the geographic separation of populations and  genetic isolation, in combination with an adaptationprocess induced by selection pressures, for example,of a special environment (allopatric speciation).However, speciation may also occur within an areaand population, for example, by polyploidization(sympatric speciation).Coevolution is defined as a process in the courseof which two partners or entire partner systems(animals, plants, bacteria, or fungi) depend on oneanother in their evolution. Both acquire specific adaptations as a consequence of mutual selectionpressure. Examples for a gene–gene coevolution,for a close coevolution between species (specific coevolution), as well as for a coevolution betweenspecies groups (diffuse or guild coevolution) will bepresented.Theseexamplesconcernflower–pollinatosystems, the dispersal of plant species by animals,protectionofplantsbyanimalspecies,andcoevolutionbetween plant species and phytophagous insects.Evolutive processes bring about biological diversity (biodiversity).Theconceptbiodiversitywillbedefined,and the hierarchical levels of various biological organization forms will be differentiated. Thedifferent elements of diversity (taxonomic diversity,diversity of life-forms, diversity of spatial patterns,etc.), the diversity of organismic interactions,evolution-biological and ecological factors causing diversity, and functional processes of diversity will bediscussed. Special attention will be given to intra-and interbiocoenotic diversity, the many and diverse phenomena occurring within and betweenbiocoenoses. It will be shown that the preservationof biodiversity is of paramount importance; it is anessential prerequisite for the survival of man onEarth and has thus been incorporated into theconcept of sustainable development.
PART TWO / DISCOVERY AND SPOLIATION OF THE BIOSPHERE
395
SECTION I /
THE BIOSPHERE
................................................
Basic Principles
Thetheoryofbiologicalevolutioninfluencedman’sconcep-tion of the world—also on humanistic, social, and religiouslevels—in a radical and far-reaching way. Man’s anthropo-centric conception was first shaken in the sixteenth centuryby Nicolaus Copernicus, whose Copernican system movedtheearth—andthusman—outofthecenteroftheuniverse.The selection theory of the English naturalist Charles Dar-win (1809–1882) upset it even further: The species
Homo sapiens
is related to all other organisms.Biological evolution, coevolution between species, andthe phenomenon of biodiversity (which is the result of evo-lutiveprocesses)arecausallycloselyconnected.Thefactors
 ◗    
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ANSELM KRATOCHWIL, ANGELIKA SCHWABE
396
VOLUME IV / THE LIVING WORLD
discussed in the following sections are essential prerequi-sites for evolutive processes.
Energy
Without the sun as an extraterrestrial energy source, lifewould not be possible. The development of higher molecu-larchemicalcompoundsduetophotosynthesis,theprimaryproductionofautotrophicorganisms(producers)—withoutwhich the consumers and decomposers (animals, bacteria,and fungi) could not have evolved—and temperatures fa-vorableforthemetabolismprocessesoflivingorganismsarean essential prerequisite for organismic life. However, theordered variety of different life-forms (biodiversity) withinbiocoenoses and ecosystems, reflected by numerous plantand animal species, is also based on a constant energy sup-ply. The thermodynamic laws (basic principles of thermo-dynamics)applybothtoinorganicmatterandtoorganis-mic life. Order (neg entropy) within a system (in this case,a biosystem) is only possible when energy is available (sup-ply offree enthalpy). Without energy supply, the compo-nents of a system (compartments) cannot ensure any trans-fer of matter or information. Such a transfer, however, isnecessary for the functioning of the entire system, which ismaintained by system-inherent characteristics—that is, theinteractions of its compartments. Energy failure, on theother hand, leads to entropy (disorder) and breakdown inthe system.
Organisms and Their Genetic Information
The genetic code (DNA and RNA) of organisms con-tains information regarding the preservation of the species-and organism-specific life processes and the organizationof the constructive and the energy metabolism. Moreover,it ensures that the organism is able to respond to environ-mentalinfluencesandthatthegeneticinformationispassedon to the next generation.
Organisms and Their Structuresand Functions
Organisms have structures: To survive in a specific abi-otic and biotic environment, they have acquired morpho-logical,physiological,andbiochemicaladaptations,andani-mals have also acquired ethological adaptations. The manyand diverse environments (habitats as well as habitat com-plexes) bring about a diversity of structures. As a rule,structures of organisms (adaptations) are linked to certainfunctions and vice versa.
Mutation
Mutations are the basis of evolution. A mutation is achange in the genotype, either spontaneous and natural orinduced by certain mutagenes (substances provoking mu-tations, high temperatures, and UV rays). The variation of genetic information within populations and between them(genetic variability) is due to mutations.
Selection
Mutation provides a wide variety of possible forms andphenomena; selection then “chooses” from the existing or-ganisms those reaction types which are best suited, for ex-ample, to the prevailing environmental conditions. Thesetypesreproducebest(“highfitness”)andthuspassonmanytheir genes and alleles to the next generation.
Time
Thehistoricityofthetaxonomicsystemsisoneofthefun-damentalcharacteristicsofbiology.Biodiversityistheprod-uct of evolution and thus the result of phylogenesis effec-tive over millions of years. Dollo’s law on the irreversibilityof evolution-historical processes implies that a species canonly originate once. Intricate structures lost in the courseof evolution can never be regained in their original formsince combinations of random mutations and directionalselection,asarule,arenotreproducibleandevolutiontakesa linear course. However, there are examples showing, forinstance, that some metabolic pathways which occur in dif-ferentrelatedgroupshaveobviouslybeenacquiredbymoresimple mutation and selection steps (e.g., the Crassulaceaeacid metabolism in plants).In the following section, some basic principles of thebiological evolution are explained. The evolution processmay be considerably accelerated and the degree of coor-dinated adaptations may be especially pronounced whenspecies or species groups exercise a significant selectionpressure on each other (coevolution). Some particularlyspectacular phenomena are due to coevolution. The formsof diversity will be described on the different hierarchy lev-els of biological systems (species, biocoenosis, ecosystem,and landscape).................................................
Evolution
The Evolution Theory
The theory of the phylogenetic development of organ-isms, first outlined in Charles Darwin’s epochal work
TheOriginofSpeciesbyMeansofNaturalSelection
(1859),ises-sentiallystillvalidtodayandhasbeensubstantiatedbydatafrom nearly all fields of biology.The theory is based on two central principles: (i) Allorganisms are phylogenetically related to each other, and(ii)evolution is due to undirectional changes in the geneticmaterial (mutation), leading to transformations of shape,function, and mode of life of organisms (species) and to di-rectional selection by the abiotic and biotic environment.Darwin’s selection theory (with regard to phylogenesis
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PART TWO / DISCOVERY AND SPOLIATION OF THE BIOSPHERE
397
EVOLUTION, COEVOLUTION, AND BIODIVERSITY
designated as
theory of evolution
or
theory of descent 
) con-tradicts the thesis of the French zoologist Jean-Baptiste deLamarck (1744–1829). According to Lamarck’s thesis (de-veloped in 1809), there is no genealogical relationship be-tweentheorganisms.Evolutioniscausedbyachangeinthegenotype, provoked by altered environmental conditions.Thenewlyacquiredcharacteristicsarepassedontothenextgenerations (Lamarckism) due to an organism-inherentimpulse of perfection (psycho-Lamarckism). The selectiontheory, however, implies that only the phenotype, and notthe genotype, can be changed, via modifications, by envi-ronmental influences, but environmental influences deter-mine the selection processes and favor specific genotypes.At the same time, but independent of Charles Darwin,the biogeographer Alfred Russel Wallace (1823–1913) as-certained the same main mechanisms of evolution. He didnot work out a detailed theory, however.Modern genetic (especially population-genetic) datahave contributed to a better understanding of mutationprocesses and genetic variation and have thus confirmedand further substantiated the evolution theory. Disciplinessuch as ecology, ethology, developmental biology, system-atics, special botany and special zoology, and paleontologyhave made considerable progress, and differentiated math-ematical procedures have been developed. Consequently,evolutionary biology could be established as synthetic sci-ence (synthetic evolution theory; Huxley, 1974).The course of phylogenesis and the factors which in-duced it are a topic of current research.
Evidence for Evolution
Important tasks of evolution research are the analysisofthe change in species (anagenesis), reconstruction of thephylogenetic development (“phylogenetic tree” research),and demonstration of evolution factors (causal evolutionresearch).The following evidence for evolution has been gathered.
Data of Homology Research.
Organisms can be dis-tinguishedfromeachotherbycertainorgansandstructures.Organs and structures of organisms whose similarities aredue to the same hereditary information are homologous.As homologous organs change in the course of evolution,they have to be investigated with regard to certain criteria.The following homology criteria apply (Osche, 1966):Criterion of position: Organs and structures are homol-ogous when they occupy the same position within an or-ganism (
homotopic organs
); for example, the position of la-bium, labrum, mandible, and maxilla of the mouthparts of insects (Fig.1).Criterion of continuity or steadiness: Organs and struc-tures which are neither similar to each other nor occupy thesame position within an organism (
heteromorphous
or
het-erotopic organs
) can be recognized as homologous by tran-sitions and intermediate forms. Examples can be given(i) from the embryonic developmentconversion of the serially and segmentally arranged, homologous ap-pendices within the insect embryo into mouthparts andextremities (see Fig.1);
FIGURE 1
Homology of the mouthparts in different orders of insects. Dark blue, mandible; red, maxilla; lightblue, labium. (a) Ventral view of the embryo of an insect, with the segmental rudiments of the extremity budswhich in the head develop into mouthparts and in the thorax into the extremities. (b) Cockroach (genus
Blatta
);the mouthparts (shown separately) have the typical original chewing form. (c) Honeybee (genus
 Apis
); maxillaand labium parts have been converted into a sucker and the mandibles are fully developed. (d) Butterfly (orderLepidoptera); parts of the galea are stretched into a proboscis, whereas the mandibles are reduced (adaptedfrom Czihak
etal.,
1976).
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