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2nd way - parallelism

Process evolutionary development in a similar direction of two or more divergent groups.

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Saber-toothed tiger

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3rd way - convergence

The process of evolutionary development of two or more unrelated groups in a similar direction.

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Progress and its role in evolution

20s of the 20th century

A.N. Severtsov I.I. Shmalgauzen - created the doctrine of biological progress

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Criteria for biological progress:

1. increase in the number;

2. range expansion

3. increase in the systematic groups of a given taxon

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Ways to achieve biological progress:

• Arogenesis is a way of development of groups of organisms with access to another adaptive zone, under the influence of fundamentally new adaptations acquired by the group (aromorphosis).
• Example: sexual differentiation, pulmonary respiration, 4-chambered heart.

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• Allogenesis is the direction of evolution of a group of organisms, in which closely related species change some particular devices with others, and the general level of organization remains the same (idioadaptation)
• Case Study: Finches in the Galapagos Islands

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• Catagenesis is a special way of development of evolution associated with the penetration of organisms into a simpler habitat and a sharp simplification of the structure and lifestyle (general degeneration)
• Example: bull tapeworm

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Biological regression is

• Decrease in the level of adaptability to living conditions, decrease in the number of the species and the area of ​​the species range.
• Leads the species to extinction
• Example: Ussuri tiger, cheetah, polar bear.

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Macroevolution - site marking

• The study of paleontological finds and modern forms allows us to consider the existence of two main types of evolutionary development of the group (Lamark, 1809, Haeckel, 1866; and others) as fairly firmly established at present: the emergence of a large number of closely related forms that differ in adaptations of the same scale, and development with access to another adaptive zone due to the acquisition a group of some fundamentally different devices that allow you to go beyond the boundaries of the former adaptive zone.
• Adaptive radiation of the same scale is designated differently in modern evolutionary literature (idioadaptation, allomorphosis, allogenesis, cladogenesis, etc.). To use unambiguous terms, it is advisable to dwell on one of these terms; one of the most suitable seems to be the term "allogenesis" (Paramonov, 1966). To describe the development of a group on the way to a different adaptive zone, the acquisition of evolutionary adaptations greater value the terms "aromorphosis" and "anagenesis" were used. ... we, following A. L. Takhtadzhyan (1966), use the term "arogenesis" for such transformations of the group.
• (From Timofeev-Resovsky and others)
• Aramorphosis according to A.N. Severtsov does not have a clear definition. It is usually set graphically.

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• Evolutionary transformations that significantly increase the adaptive capacity of this group (allowing you to expand the old or occupy a new adaptive zone), have a significant impact on the organism as a whole, but do not change the overall level of organization, were called epectomorphoses. …Epectomorphoses persist for a long time during subsequent adaptive radiation, becoming signs of large taxa
• Schmalhausen suggested calling regressive changes catamorphoses, and replacing the term "idioadaptation" with allomorphoses.
• were proposed ... new names for the three main directions of the evolutionary process: morphophysiological progress began to be called arogenesis (or anagenesis), morphophysiological regression - catagenesis, the development of particular adaptations - allogenesis or cladogenesis
• Scheme of the main directions of the evolutionary process:
• A - arogenesis, AL - allogenesis, K - catagenesis, E - epictogenesis
• Planes represent different levels of organization
• From Jordanian

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• A. N. Severtsov called the evolutionary transformations of the organization, leading to morphophysiological progress, aromorphoses. According to A. N. Severtsov, aromorphoses are such changes in the structure and functions of organs that are of general importance for the organism as a whole and raise the energy of its vital activity to a new qualitative level.
• Thus, the undoubted aromorphoses in the evolution of vertebrates were: the development of the mechanism of active gill ventilation (gill pump) in the most ancient vertebrates through movements of the visceral gill skeleton, the acquisition of the jaw apparatus (with restructuring of the anterior gill arches), the intensification of gill ventilation during the development of the gill cover in bone fish, the acquisition by the latter of a swim bladder - a hydrostatic apparatus that allows fish to regulate their buoyancy; the development in the ancestors of higher terrestrial vertebrates - amniotes of embryonic membranes (amnion, serosa, allantois), which provide the possibility of laying eggs on land; the development of a powerful suction (rarefying) respiratory pump of the chest in reptiles; formation aircraft in birds; development of live birth and feeding of young with milk in mammals; improvement of the brain in birds, mammals and humans.
• Aromorphoses have a very high general adaptive value, increasing the body's independence from the external environment.

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• Upper and lower jaws and teeth of various snakes
• 1 - python, 2.7 - smooth-toothed already-shaped, 3, 8 - back-furrowed already-shaped, 4,5,9 - slate, 6, 10 - viper
• Skullviper Rhinoceros
• Modifications of the skull of snakes - an example of epectogenesis
• A free, movable, rotating upper jaw and a free, downwardly shifting lower jaw - an epectomorphosis that allowed snakes
• switch from eating insects (the food spectrum of legless lizards) to hunting vertebrates. The general level of vital activity (energy exchange) remains the same.
• Video

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• Divergence (from medieval Latin divergo - I deviate)
• The wolverine feeds mainly on carrion. Wide paws with webbing between the fingers help her to move on loose snow in the northern taiga and forest tundra. Weight about 15 kg.
• Weasel is the smallest of mustelids. She is able to penetrate mouse holes. It feeds mainly on mice.
• The badger is omnivorous, active in the warm season, hibernating in the northern part of its range in winter. It builds very complex burrows in which it spends most of its time. Weight about 15 kg.
• Divergence within the mustelid family

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• Convergence is not the exact opposite of divergence. Taxa diverge, and convergence is a characteristic of the evolution of organs (similar functions in non-homologous organs) and (or) life forms of distant taxa with the acquisition of external similarity.
• Convergence (from lat. convergo - approaching, converging)
• Mammals adapted to gliding jumps: 1 - woolly wing, 2 - marsupial squirrel, 3 - flying squirrel

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• Parallelism - convergence of homologous organs
• The posture of "standing in a column" and the location of the eyes on the head, characteristic of rodents in open spaces. In the bottom row are related forms in other biotopes.
• 1 - small ground squirrel, 2 - Brandt's vole, 3 - yellow pied, 4 - large gerbil, 5 - common squirrel, 6 - eastern vole, 7 - afternoon gerbil.
• Parallelism in the development of limbs in equines and liptoterns in the Neotropics

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• The term parallelism is also used not to characterize changes in organs, but to indicate the direction of evolution of taxa.
• An example of parallel evolution can be the Quaternary history of whitefish: spreading from a single center (West Siberian sea-lake) throughout the north of the Holarctic, they formed a number of isolated groupings of the subspecies-semispecies-superspecies rank. Within the isolates, independent differentiation occurs into few stamen forms, feeding on benthos, and many stamens, feeding on plankton, as well as a parallel division according to the place of reproduction into lake and river forms.

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• Polyphyly is the union of groups of different origin into one taxon. Poly is a lot, phylum is a branch.
• In the second half of the 20th century, many researchers tried to prove the polyphyletic origin of such well-known groups as flowering, amphibians, and mammals.
• Monophyly is the origin of a taxon from a single ancestral taxon.
• Simpson proposed to consider as monophyletic taxa that are generated by a taxon equal in rank to them (monophyly according to Simpson).
• However, the rank of higher taxa is subjective.
• Ashlock suggested that supraspecific taxa be considered monophyletic, descended from one ancestral species (Ashlock monophyly)
• A rather representative school of cladists accepts a more rigorous criterion of monophyleticity - origin from one species, but how to prove it and how to operate with it? It is pointless to use Ashlock monophyly, which is deprived of the possibility of verification.

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• Neutralist theory of evolution - neutral signs only diverge. Paraphilia and polyphilia are excluded at this level.
• Innovative news: evolution is two-headed, like an eagle on a ruble.
• Head number 1. Adaptive evolution, Darwinian evolution - the evolution of adaptations guided by selection. It can be convergent, divergent, progressive-regressive, etc.
• Head number 2. Neutralist evolution, non-Darwinian evolution - the evolution of traits that are indifferent to selection (the difference in the thickness of the lips between blacks and Indians). If a trait is not subject to stabilizing or driving selection, it always "spreads out".
• Analogy - the divergence of languages ​​in the cultural isolation of their speakers. The divergence of languages ​​is the greater, the longer the isolation: man - man, men - Mensch. They are not tied to either the terrain or the climate.

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Neutral evolution is the key to phylogeny

• Phylogenetic relationships between 8 groups of vertebrates (upper diagram) and differences in the number of amino acid substitutions (Kaa) in the same species (lower graph) for hemoglobin α-chains (black dots) and β-chains (light dots). From Kimura, 1985)
• Additive tree based on differences in transfer RNAs.
• It cannot be done otherwise.
• The phylogeny of such large groups is created only by analyzing the evolution of macromolecules.

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Macroevolution - modern complexities

• To date, the possibilities of morphological, embryological, and paleontological methods for analyzing phylogeny have been practically exhausted. With their help, it was possible to create a relatively complete picture of the evolution of vertebrates and vascular plants, and partially of invertebrates.
• Lower plants and prokaryotes do not lend themselves to traditional analysis.
• Since the 1980s, there has been rapid progress. AT in general terms the contours of phylogenetic relationships of all living things are outlined. The overall design was completely unexpected.
• The new knowledge is based on a new method - the study of the evolution of molecules, first - neutral, then, already in this century, adaptive.
• At the initial stage of evolution, active "horizontal" gene transfer is observed - their exchange between representatives of very distant systematic groups. This is the main factor in the evolution of the Archean and, to a lesser extent, Proterozoic biota.
• In eukaryotes, it is accompanied by cell fusion or the absorption of a cell by a cell with the most varying degrees autonomy of individual components of a common superorganism (symbiogenetic theory).
• The phylogenetic scheme at this level is not a tree, but a network.

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• The evolutionary tree of eukaryotes. Branch points from the main stem above the Euglenozoic are arbitrarily marked.

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Horizontal gene transfer - what it leads to

• Prokaryotes are represented by two kingdoms - archaebacteria and eubacteria
• Domain - a conservative sequence of amino acids present in several (usually many) protein molecules in different organisms. Most domains are characterized by a strictly defined function and are functional blocks of protein molecules.
• In the eukaryotic genome, proteins responsible for operations with the genome (replication, transcription, translation) and proteins that perform operations with membranes are from archaea, proteins, proteins of basic metabolism are from eubacteria.
• there is a hypothesis that the primary prokaryotic non-nuclear organism was formed by the fusion of archaebacteria with eubacteria, and the basic energy metabolism of this organism was of a eubacterial nature (glycolysis, fermentation)
• Quantitative ratio of common and unique protein domains in archaea, bacteria and eukaryotes. The areas of the figures are approximately proportional to the number of domains (from A.V. Markov, A.M. Kulikov, 2004).

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Biological progress: the tension between complexity and success

A.N. Severtsov removes it, separating biological and morpho-physiological progress

Biological progress:

• increase in the number of individuals
• progressive settlement and the capture of new areas
• breakdown of a taxon into subordinate systematic units

Morpho-physiological progress:

• differentiation of the organism
• function intensification
• Severtsov's followers add an improvement in integration, rationalization of the body structure, an increase in the level of homeostasis, etc.

Morpho-physiological progress is one of the ways to achieve biological progress.

As well as morpho-physiological regression.

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The main philosophical problem of macroevolution is the orientation

• Early evolutionists explained the development of nature in terms of final causes, the desire for progress, creative power, and similar agents.
• Darwin removed the predetermination of the course of evolution, but not everyone liked it.
• Periodically, both abroad and in our country, heresies arose, aimed at searching for reasons for evolution other than natural selection.
• Darwinism is a theory that allows you to explain everything “backward”, but leaves no room for predictions – how does it differ from theories of other social and social sciences. Darwinian evolution is random and unpredictable.
• In the USSR, the rejection of Darwinism went under the flag of nomogenesis - an attempt to build evolution on the basis of "laws" (nomos in Greek - law). They ended in nothing, but nomogenetics - L.S. Berg, A.A. Lyubishchev - were such bright and original personalities that nomogenesis became an important page in the history of Russian biology.