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[Evolution as resistance to entropy. I. Mechanisms of species homeostasis].

The idea is discussed that the common output of any evolution is creation of the entities that are increasingly resistant to further evolution. The moving force of evolution is entropy, the tendency to disorder. This general aspiration for chaos is a cause of the mortality of organisms and species, however, being prerequisite for any movement, it creates (by chance) novelties, which may occur (by chance) more resistant to further decay and thus survive. The surviving of those who survive is the most general principle of evolution discovered by Darwin for particular case of biological evolution. The second law of thermodynamics states that our Universe is perishing but its ontology is such that it creates resistance to destruction. The evolution is a history of this resistance. Not only those who die do not survive but also those who evolve. The entities that change (evolve) rapidly disappear rapidly and by this reason they are not observed among both the fossils and now-living organisms. We know only about long-living species. All the existing organisms are endowed with an ability to resist other changing. The following main achievements of the species homeostasis are discussed: high fidelity of DNA replication and effective mechanisms of DNA repair; diploidy; normalizing selection; truncated selection; heterozygote superiority; ability to change phenotype adaptively without changing genotype; parental care and the K-strategy of reproduction; behavior that provides independence of the environment. The global resistance of the living systems to entropy is provided the state that all the essential in biology is determined not by physical-chemical interactions but could semantic rules. A conception of "potential zygotic information" that determines the rules of ontogenesis is proposed. A zygote does not contain this information in explicit form. It is created de novo step by step during ontogenesis and it could not be decoded beforehand. The experimental data on the adaptive mutagenesis and the relevant hypothesis are discussed. It is concluded that the special mechanisms for speeding-up of evolution as created by evolution are impossible conceptually.

Adaptation, Biological↗

The evolution of the neocortex in mammals: intrinsic and extrinsic contributions to the cortical phenotype.

The neocortex is that portion of the brain that is involved in volitional motor control, perception, cognition and a number of other complex behaviours exhibited by mammals, including humans. Indeed, the increase in the size of the cortical sheet and cortical field number is one of the hallmarks of human brain evolution. Fossil records and comparative studies of the neocortex indicate that early mammalian neocortices were composed of only a few parts or cortical fields, and that in some lineages such as primates, the neocortex expanded dramatically. More significantly, the number of cortical fields increased and the connectivity between cortical fields became more complex. While we do not know the exact transformation between this type of increase in cortical field number and connectivity; and the emergence of complex behaviours like those mentioned above, we know that species that have large neocorticies with multiple parts generally have more complex behaviours, both overt and covert. Although a number of inroads have been made into understanding how neurons in the neocortex respond to a variety of stimuli, the micro and macro circuitry of particular neocortical fields, and the molecular developmental events that construct current organization, very little is known about how more cortical fields are added in evolution. In particular, we do not know the rules of change, nor the constraints imposed on evolving nervous systems that dictate the particular phenotype that will ultimately emerge. One reason why these issues are unresolved is that the brain is a compromise between existing genetic constraints and the need to adapt. Thus, the functions that the brain generates are absolutely imperfect, although functionally optimized. This makes it very difficult to determine the rules of construction, to generate viable computational models of brain evolution, and to predict the direction of changes that may occur over time. Despite these obstacles, it is still possible to study the evolution of the neocortex. One way is to study the products of the evolutionary process--extant mammal brains-and to make inferences about the process. The second way to study brain evolution is to examine the developmental mechanisms that give rise to complex brains. We have begun to test our theories regarding cortical evolution, generated from comparative studies, by 'tweaking' in a developing nervous system what we believe is naturally being modified in evolution. Our goals are to identify the constraints imposed on the evolving neocortex, to disentangle the genetic and activity dependent mechanisms that give rise to complex brains, and ultimately to produce a cortical phenotype that is consistent with what would naturally occur in evolution.

Animals↗

Comparative Genomics Reveals Convergent Evolution Between Avivorous Bats (Ia io and Nyctalus aviator).

Investigating the genetic basis of dietary specialization can provide insights into the evolution of niche breadth. In this study, we employed comparative genomics to investigate the adaptive mechanisms enabling two bat species (Nyctalus aviator and Ia io) to shift from insectivory to seasonal bird consumption (avivorous bats). Our findings revealed adaptation related to immune response and lipid metabolism in avivorous bat species. Avivorous bats exhibit strong positive selection and convergent evolution in immune-related genes, which are under heightened selective pressure compared to those of non-avivorous bats. These species also display significantly fewer endogenous retroviral elements. These findings emphasized the significance of immune-driven adaptive evolution in avivory. Additionally, our results showed that the dietary evolution of avivorous bats is accompanied by convergent evolution associated with the lipid metabolism. Notably, CEPT1, the upstream gene required for the activation of the PPAR pathway, underwent positive selection and convergence, which may have affected lipid metabolism. These adaptations may enable avivorous bat species to face the challenge of immune response and nutrition during dietary niche expansion. These findings not only provide comprehensive insights into the adaptive evolution driving the unique diet of avivorous bats but also offered novel perspectives on the molecular mechanisms underlying ecological niche evolution in a dietary context.

Animals↗

Clinical and cytogenetic significance of myelodysplastic syndromes with disease evolution.

We performed a retrospective study of 83 patients with myelodysplastic syndrome (MDS) to clarify the clinical and cytogenetic implications of disease evolution. Twenty-three patients showed disease progression; six of the 11 patients whose disease evolved within 100 days showed complex cytogenetic aberrations and most of them died within 300 days. Of the patients who survived more than 300 days, those with high bone marrow (BM) blast percentages experienced significant disease progression, but we noted no cytogenetic indicators for disease evolution at the later phase. Sixty percent of patients showing karyotypic evolution without disease evolution had deletion-type chromosome changes. The most frequent anomaly in patients with disease evolution who survived more than 300 days was an additional numerical change, whereas patients with disease evolution who survived less than 300 days showed karyotypic instability. It was difficult to predict disease progression for patients whose disease evolved more than 300 days after diagnosis, but in some patients the presence of additional numerical changes was related to disease progression. The cutoff level of early disease evolution was 100 days after diagnosis, and most patients with complex abnormalities survived less than 300 days with or without disease evolution.

Adult↗

Chromosomal alterations associated with evolution from myelodysplastic syndrome to acute myeloid leukemia.

Several studies have demonstrated the prognostic value of cytogenetic analysis in MDS both for survival and progression to AML. However it is unknown which are the numerical or structural abnormalities required for leukemic transformation. In this report we studied clinically and cytogenetically 127 patients: 125 with primary MDS and two with AML with a previous history of MDS. Thirty-one patients (24%) showed evolution of the disease during the follow-up study. Chromosomal abnormalities found at diagnosis in patients that progressed toward AML included: del(5)(q15), +6, del(6)(q21), t(5;8)(q32;q22),-7, del(7)(q22), der(7)t(1;7)(q10;p10), t(7;11)(p15;p15), +8, del(11)(q23), del(12p), del(3)(q21), del(20)(q12) and complex karyotypes. Eight of these patients were studied cytogenetically during transformation and showed acquisition of chromosomal alterations involving dup(1q), +8, del(11)(q23), and translocations between chromosomes 1 and 8 or 7 and 17. In addition we also observed gain of ploidy and monosomy 21. These results suggest that chromosomal alterations during evolution of the disease include special chromosome gains or abnormalities of chromosomes 1, 7, 8, 11 and 17 with involvement of ETV-1, Hox-A9, Pax 4, MLL genes besides a putative gene mapped at 17q25. We also applied the International Prognostic Scoring System (IPSS) to 114 patients, excluding those submitted to allogeneic bone marrow transplant. Our patients were classified into four distinct risk groups. The analysis of risk groups presented by 27 patients who showed evolution of the disease revealed 18 at the high risk group and four at the intermediate-2 group. From the intermediate-1 risk group only five patients showed evolution of the disease. Three of these patients evolved from RA to RAEB with gain of a del(11)(q23) or an expansion of a del(12)(p12) clone. Our results suggest that some chromosomal alterations are responsible for each step in the evolution of the disease. As the pathway of evolution is not unique it has been very difficult to define what genetic alteration comes first. However from several results in the literature and our own, it seems that some chromosomal alterations may predict the evolution of the disease and are correlated with short survival, as for example the trisomy of chromosome 8, and might be incorporated in the high risk group in the IPSS. This score system has been proved to be useful for predicting survival and evolution from MDS to AML.

Adolescent↗

Long-term evolution of gut inflammation in patients with spondyloarthropathy.

BACKGROUND & AIMS: Intestinal inflammation has been observed in patients with spondyloarthropathy (SpA). This prospective study reports the evolution of the intestinal inflammation observed in patients with SpA. METHODS: One hundred twenty-three patients with SpA who had undergone initial endoscopy were clinically reassessed. Intestinal evolution was evaluated by ileocolonoscopy and the histological study of biopsy specimens in 49 patients. RESULTS: Articular remission rates were independent of initial gut inflammation and associated with endoscopic and histological remission. Persistent gut inflammation was observed in active joint disease. Gut inflammation rarely disappeared, despite the persistence of articular complaints. Initial chronic gut inflammation implied a high risk of evolution to ankylosing spondylitis. Evolution to inflammatory bowel disease (IBD) was observed in 7% of patients. Mainly patients with initial chronic inflammation and mild complaints of diarrhea were at risk. Sulfasalazine was more frequently needed in the treatment of patients with gut inflammation with a beneficial effect on articular and intestinal evolution but did not prevent evolution to IBD. CONCLUSIONS: This study supports the etiopathogenetic role of the gut in SpA. Presence of chronic gut inflammation and mild complaints of diarrhea implies a high risk of evolution to ankylosing spondylitis and IBD. Sulfasalazine has a beneficial effect on articular activity by controlling gut inflammation, but it cannot prevent evolution to overt IBD.

Adolescent↗

Effects of Temperature on Photosynthesis and CO(2) Evolution in Light and Darkness by Green Leaves.

Using an open and a closed system of gas analysis, it was found that CO(2) evolution in light and in darkness from plant leaves (sunflower, soybean, watermelon, eggplant, and jackbean) have a different response to temperature. While the rate of CO(2) evolution in light increased with increasing temperature from 17 to 35 degrees and then declined, the rate of CO(2) evolution in darkness increased continuously up to 40 degrees . The rate of CO(2) evolution in light was affected by light intensity. At 1800 ft-c and below 35 degrees the rate of CO(2) evolution in light was greater than in darkness, but above 35 degrees it became lower than in darkness. The Q(10) for CO(2) evolution in light was consistently lower than that in darkness.Apparent photosynthesis decreased with increasing temperature, from 20 to 40 degrees and its rate was affected by both light intensity and oxygen concentration. In leaves of dicotyledonous plants studied the decrease in apparent photosynthesis between 20 to 30 degrees at 21% O(2) was shown to be due primarily to an increase in CO(2) evolution in light with relatively little effect on photosynthesis.In corn which does not evolve CO(2) during illumination there was little effect of increasing temperature on the rate of apparent photosynthesis.The different response to temperature of CO(2) evolution in light and in darkness support the earlier conclusion that these are 2 different processes.

Journal Article↗

Correlation of absorbance changes and thylakoid fusion with the induction of oxygen evolution in bean leaves greened by brief flashes.

Dark-grown bean leaves (Phaseolus vulgaris) which had been greened for several days in a repetitive series of brief xenon flashes were studied during the initial induction period when O(2) evolution first appears. The induction of O(2) evolution requires actinic irradiation (e.g. 2 mw/cm(2) of red light) and goes to completion in about 8 minutes with a half-time just under 3 minutes. Absorbance measurements on the intact leaves showed that a change of a carotenoid pigment, monitored at 505 nm, was closely correlated with the rate of O(2) evolution during the induction period. Inhibitor studies, however, showed that the absorbance change persisted in the presence of a number of inhibitors which blocked O(2) evolution. Electron microscopy revealed that the primary thylakoids which were unfused in the flashed leaves before induction became fused in pairs or groups of three during the 8-minute induction period. It is postulated that the 505-nm absorbance change of the carotenoid pigment is correlated more directly with the fusion process than with O(2) evolution. Heat treatment (45 C for 5 min) or infiltration with 0.8 m tris, which prevented the fusion process, also prevented the absorbance change.If the leaves were preilluminated for 8 minutes with very weak red light (20 muw/cm(2)) which induced no O(2) evolution, absorbance change, or thylakoid fusion, there was an immediate burst of O(2) evolution at the onset of actinic irradiation and the induction period, as noted by O(2) evolution or by the 505-nm absorbance change, was reduced to 2 minutes (half-time of 40 seconds). It is concluded that the electron transport system in the flashed leaves is blocked at the Mn site between water and photosystem II and that the photoactivation of Mn into the thylakoid membranes occurs during the low light, photoactivation process. After the electron transport chain is thus repaired, ion-pumping mechanisms driven by actinic light may lead to steady-state photosynthesis as well as to thylakoid fusion.

Journal Article↗

The evolution of genomic base composition in bacteria.

Guanine plus cytosine (GC) content ranges broadly among bacterial genomes. In this study, we explore the use of a Brownian-motion model for the evolution of GC content over time. This model assumes that GC content varies over time in a continuous and homogeneous manner. Using this model and a maximum-likelihood approach, we analyzed the evolution of GC content across several bacterial phylogenies. Using three independent tests, we found that the observed divergence in GC content was consistent with a homogeneous Brownian-motion model. For example, similar rates of GC content evolution were inferred in several different bacterial subclades, indicating that there is relatively little rate heterogeneity in GC content evolution over broad evolutionary time scales. We thus argue that the homogeneous Brownian-motion model provides a good working model for GC content evolution. We then use this model to determine the overall rate of GC content evolution among eubacteria. We also determine the time frame over which GC content remains similar in related taxa, using a flexible definition for "similarity" in GC content so that, depending on the context, more or less stringent criteria may be applied. Our results have implications for models of sequence evolution, including those used for phylogenetic reconstruction and for inferring unusual changes in GC content.

Bacteria↗

General theory of evolution.

The biological, neural, cultural and technical evolutions and their phenomena have been explored, and on the basis of our findings the formation of a general theory of evolution has been undertaken. In each of the systems studied, the presence of structural building units, excitable structures and an energy-flow going through the system can be observed. Under the organizing effect of this energy-flow, the spontaneous generation of the replicative information begins and the structures of the system establish functional relations with each other. It can be demonstrated that the evolution of structures has a replicative character. The evolution goes through a phase of non-identical replication, and reaches the phase of identical replication. The parts of the system become separated, that is, compartments develop within it. The replicative information becomes compartmentalized and it converges. As a consequence of the convergence, the compartments compose new structural units which is tantamount to the development of new evolutional levels. The direction of evolution is determined by the growth of replicative information, and this process is concluded when the total system becomes one replicative unit. In the last part of the paper a few of the basic principles of evolution concerning matter, energy and information are drawn up.

Animals↗

Molecular evolution of growth hormone (GH) in Cetartiodactyla: cloning and characterization of the gene encoding GH from a primitive ruminant, the chevrotain (Tragulus javanicus).

In mammals the sequence of pituitary growth hormone (GH) is generally strongly conserved, indicating a slow basal rate of molecular evolution. However, on two occasions, during the evolution of primates and that of cetartiodactyls, the rate of evolution has increased dramatically (25 to 50-fold) so that the sequences of human and ruminant GHs differ markedly from those of other mammalian GHs. To define further the burst of GH evolution that occurred in cetartiodactyls, the GH gene of the chevrotain (Tragulus javanicus) has been cloned and characterized by use of genomic DNA and a polymerase chain reaction technique. Two very similar gene sequences, which probably reflect allelic variation, were isolated. The deduced sequence for the mature chevrotain GH differs from that of the bovine or red deer GH at only two to three residues, and phylogenetic analysis shows that the burst of rapid evolution of GH that occurred in the Cetartiodactyla must have been completed before the divergence of the Tragulidae and the advanced ruminants (Pecora). The rate of evolution during this burst must therefore have been greater than previously estimated. In other aspects (including signal sequence, 5' upstream sequence, and synonymous substitutions in the coding sequence), the chevrotain GH gene differs considerably from the GH genes of other ruminants and here there is no evidence for the period of accelerated evolution that is seen for GH itself.

Amino Acid Sequence↗

Ubiquitins revisited: further examples of within- and between-locus concerted evolution.

Ubiquitin genes provide a model for studying the effects of concerted evolution on the evolution of a family of short repeated sequences. Previous work has demonstrated the occurrence of within-locus concerted evolution and raised the question of the effectiveness of between-locus concerted evolution for ubiquitin repeats. In this study comparative analysis of additional nucleotide sequences of ubiquitin tandem repeats provides further details of within-locus concerted evolution. Moreover, the availability of multiple polyubiquitin loci and ubiquitin fusion loci within a species makes possible the detection of between-locus concerted evolution. These data indicate that concerted evolution is an effective force for homogenizing repeats between, as well as within, loci.

Animals↗

Molecular evolution of ruminant lysozymes.

The evolution of a new digestive enzyme, stomach lysozyme, from an antibacterial host defense enzyme provides a link between molecular evolution and organismal evolution. Lysozymes have been recruited at least three times (twice from a conventional lysozyme c and once from a calcium-binding lysozyme c) in vertebrates for functioning in the stomach. The recruitment of lysozyme for its new biological function involved many molecular changes, beyond those required to adapt the protein to function in the stomach. The evolution of the stomach lysozyme gene has been extensively studied in ruminant artiodactyls. In ruminants, the lysozyme c gene has duplicated to yield a family of about ten genes. These duplications allowed: (1) specialization of gene function and (2) increased levels of expression. The ruminant stomach lysozyme genes have evolved in an episodic fashion - there was a period of rapid adaptive sequence evolution, driven by positive selection in the early ruminant, that was followed by an increase in purifying selection upon the well-adapted stomach lysozyme sequence among modern species. Recombination of small portions (exons) of the genes between members of the lysozyme gene family may have aided in adaptive evolution. Evolution to a stomach lysozyme is not irreversible; at least one member of the ruminant stomach lysozyme gene family appears to have reverted to a more ancestral function, yet retains hallmarks of its history as a stomach lysozyme.

Animals↗

Molecular pathways to parallel evolution: I. Gene nexuses and their morphological correlates.

Aspects of the regulatory interactions among genes are probably as old as most genes are themselves. Correspondingly, similar predispositions to changes in such interactions must have existed for long evolutionary periods. Features of the structure and the evolution of the system of gene regulation furnish the background necessary for a molecular understanding of parallel evolution. Patently "unrelated" organs, such as the fat body of a fly and the liver of a mammal, can exhibit fractional homology, a fraction expected to become subject to quantitation. This also seems to hold for different organs in the same organism, such as wings and legs of a fly. In informational macromolecules, on the other hand, homology is indeed all or none. In the quite different case of organs, analogy is expected usually to represent attenuated homology. Many instances of putative convergence are likely to turn out to be predominantly parallel evolution, presumably including the case of the vertebrate and cephalopod eyes. Homology in morphological features reflects a similarity in networks of active genes. Similar nexuses of active genes can be established in cells of different embryological origins. Thus, parallel development can be considered a counterpart to parallel evolution. Specific macromolecular interactions leading to the regulation of the c-fos gene are given as an example of a "controller node" defined as a regulatory unit. Quantitative changes in gene control are distinguished from relational changes, and frequent parallelism in quantitative changes is noted in Drosophila enzymes. Evolutionary reversions in quantitative gene expression are also expected. The evolution of relational patterns is attributed to several distinct mechanisms, notably the shuffling of protein domains. The growth of such patterns may in part be brought about by a particular process of compensation for "controller gene diseases," a process that would spontaneously tend to lead to increased regulatory and organismal complexity. Despite the inferred increase in gene interaction complexity, whose course over evolutionary time is unknown, the number of homology groups for the functional and structural protein units designated as domains has probably remained rather constant, even as, in some of its branches, evolution moved toward "higher" organisms. In connection with this process, the question is raised of parallel evolution within the purview of activating and repressing master switches and in regard to the number of levels into which the hierarchies of genic master switches will eventually be resolved.

Animals↗

Chemical evolution of photosynthesis.

The principles of biological evolution of photosynthesis are established, but the ways of chemical evolution are unclear yet. The model systems will help to elucidate the problem. Every type of photosynthesis requires photoreceptor absorbing solar radiation. We studied as photoreceptors inorganic components of Earth crust, some coenzymes and porphyrins of abiogenic and biogenic origin. By the aid of inorganic photosensitizers (TiO2, ZnO) the models of photosystems I and II were constructed. Photochemical activation of some coenzymes may serve as an intermediate step from heterotrophic 'dark' to 'light' metabolism. The further evolution led to the separation of catalytic and photosensitizing functions. Porphin, chlorin and bacteriochlorin were formed by abiogenic synthesis. Magnesium complexes of porphyrins are active being excited by light. They are capable to reversible acceptance or donation of an electron to partner molecule. Excited Mg-complexes of porphyrins (P) are capable to transfer an electron from electron-donor (D) to electron-acceptor (A) accompanied by conversion of light quanta energy into potential chemical energy. The primary electron transfer unit (D-P-A) was incorporated into primary membrane. The transition from random to anisotropic arrangement of (D-P-A) in the membrane was plausable as a step of evolution; charge translocation appeared. (D-P-A) units created in the period of chemical evolution were probably used in the course of biological evolution. The (D-P-A) units were coupled with noncyclic and cyclic electron transfer resulting in ATP formation; coupling of two (D-P-A) units led to H2O oxidation and NADP reduction in photosynthetic organisms. The improvement of pigments biosynthesis created the phenomenon of excitation energy migration from the bulk of the pigment to (D-P-A) unit, being reactive center. The models described points the plausible steps of chemical evolution; the real sequence of events will be probably disclosed in the studies of precambrian rocks and space exploration.

Biological Evolution↗

[Historic and functional biology: the inadequacy of a system theory of evolution].

In the first half of the 20th century neo-Kantianism in a broad sense proved itself the main conceptual and methodological background of the central European biology. As such it contributed much to the victory on the typological, idealistic-morphological and psycho-vitalistic interpretations of life. On the other hand it could not give tools to the biologists for working out a strictly darwinian evolution theory. Kant's theory of organism was conceived without evolution as a theory of the internal functionality of the organism. There was only some 'play' with the evolutionary differentiation of the species. Since then the disputes around the work of August Weismann, a synthetical evolution theory which is now behind time, arose. This theory developed from coinciding claims, elaborated by geneticists, mathematicians, and by biologists studying development, natural history and systematics. This was done under a strong influence of marxist ideas. Through the interweaving of such different approaches it was possible for this evolutionary synthesis to influence successfully the development of evolution research during more than 40 years. Philosophically speaking modern evolution theory means therefore an aversion, even a positive abolition of Kantian positions. A number of biologists however--as L. von Bertalanffy--refused to adhere to a misinterpreted Kantian methodology and oriented themselves to an approach via system theory, which obtained a place in evolution research. In fact this is a Kantian approach as well. They only repeated the Kantian dilemma of the evolution which can also be found in Lamarck and Hegel. The system theory of the functionality of the organism never reaches to the level of the evolving species, but remains always on the level of epigenetic thinking, because of its philosophical origin. This paper points out the consequences of this still current dilemma. At the same time an all-enclosing reflection on the methodological, epistemological and the important historical questions of evolutionary biology in its scientific context is recommended.

Animals↗

Constrained evolution with respect to gene overlap of hepatitis B virus.

With the aim of elucidating the evolution of a hepadnavirus family, we constructed molecular phylogenetic trees for 27 strains of hepatitis B virus (HBV) using both the unweighted pair-grouping and neighbor-joining methods. All five gene regions, P, C, S, X, and preS, were used to construct the phylogenetic trees. Using the phylogenetic trees obtained, we classified these strains into five major groups in which the strains were closely related to each other. Our classification reinforced our previous view that genetic classification is not always compatible with conventional classification determined by serological subtypes. Moreover, constraints on the evolutionary process of HBV were analyzed for amino-acid-altering (nonsynonymous) and silent (synonymous) substitutions, because two-thirds of the open reading frame (ORF), P, contains alternating overlapping ORFs. In our unique analysis of this interesting gene structure of HBV, the most frequent synonymous substitutions were observed in the nonoverlapped parts of the P and C genes. On the other hand, the number of synonymous substitutions per nucleotide site for the S gene was quite low and appeared a strongly constrained evolution. Because the P gene overlaps the S gene in a different frame, the low rate of synonymous substitution for the S gene can be explained by the evolutionary constraints which are imposed on the overlapping gene region. In other words, synonymous substitutions in the S gene can cause amino acid changes in its overlapping region in a different frame. Thus, the evolution of HBV is constrained evolutionarily by the overlapping genes. We propose calling this mode of viral evolution "constrained evolution." The evolution of HBV represents a typical constrained evolution.

Amino Acid Sequence↗

The modern theory of biological evolution: an expanded synthesis.

In 1858, two naturalists, Charles Darwin and Alfred Russel Wallace, independently proposed natural selection as the basic mechanism responsible for the origin of new phenotypic variants and, ultimately, new species. A large body of evidence for this hypothesis was published in Darwin's Origin of Species one year later, the appearance of which provoked other leading scientists like August Weismann to adopt and amplify Darwin's perspective. Weismann's neo-Darwinian theory of evolution was further elaborated, most notably in a series of books by Theodosius Dobzhansky, Ernst Mayr, Julian Huxley and others. In this article we first summarize the history of life on Earth and provide recent evidence demonstrating that Darwin's dilemma (the apparent missing Precambrian record of life) has been resolved. Next, the historical development and structure of the "modern synthesis" is described within the context of the following topics: paleobiology and rates of evolution, mass extinctions and species selection, macroevolution and punctuated equilibrium, sexual reproduction and recombination, sexual selection and altruism, endosymbiosis and eukaryotic cell evolution, evolutionary developmental biology, phenotypic plasticity, epigenetic inheritance and molecular evolution, experimental bacterial evolution, and computer simulations (in silico evolution of digital organisms). In addition, we discuss the expansion of the modern synthesis, embracing all branches of scientific disciplines. It is concluded that the basic tenets of the synthetic theory have survived, but in modified form. These sub-theories require continued elaboration, particularly in light of molecular biology, to answer open-ended questions concerning the mechanisms of evolution in all five kingdoms of life.

Animals↗