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The molecular evolution of G protein-coupled receptors: focus on 5-hydroxytryptamine receptors.

Phylogenetic comparisons between homologous proteins can provide information on the rates of molecular evolution of the proteins. G protein-coupled receptors are a "superfamily" of proteins which exist in species ranging from yeast to man. Based on an analysis of the percentage of amino acid homology between various species, the rate of molecular evolution of G protein-coupled receptors can be estimated at approx 1% per 10 million years. Based on this assumption, the primordial 5-HT receptor must have evolved more than 700-800 million years ago since the 3 major classes of G protein-coupled 5-HT receptors (i.e. 5-HT1, 5-HT2 and 5-HT6 receptors) are less than 25% homologous. 5-HT5, 5-HT7, 5-HTsnail, 5-HTdro and 5-HT1A receptors differentiated approx 600-700 million years ago, the time period during which vertebrates diverged from invertebrates. The mammalian 5-HT receptor subtypes have differentiated over the past 90 million years. Thus, although a recent flurry of "new" 5-HT receptors have appeared in the literature, the first "primordial" 5-HT receptor evolved over 750 million years ago, a date which likely predates the evolution of muscarinic, dopaminergic and adrenergic receptor systems. This analysis also predicts that a significant number of both mammalian and invertebrate G protein-coupled 5-HT receptor subtypes remain to be identified.

Animals↗

Molecular evolution of eastern equine encephalomyelitis virus in North America.

We examined the rate and spatial pattern of eastern equine encephalomyelitis virus (EEEV) evolution in North America using primer-extension sequencing of viral RNA. Nucleotide sequences of the entire 26 S structural gene region of four EEEV strains revealed remarkable conservation between 1933 and 1985, with an estimated 0.7% divergence or 1.4 x 10(-4) nucleotide substitutions per site per year. Sequences from smaller 26 S regions of nine additional strains suggested that EEEV evolves in North America in a single lineage, with genetic exchange regularly occurring among enzootic transmission foci. In these limited 26 S genome regions, only synonymous nucleotide substitutions became fixed between 1933 and 1988, implying a high degree of conservation in protein structure. Short nucleotide sequences from a Panamanian, South American variety isolate revealed a relatively distant relationship to North American serotype viruses. This suggested genetic divergence between antigenic varieties, and independent evolution of EEEV in North and South America. Factors related to replication and epidemiology of EEEV, which may constrain its evolution in nature, are discussed. Possible mechanisms of genetic exchange among enzootic foci are also considered.

Amino Acid Sequence↗

Evolution of human longevity: a critical overview.

Evolution of longevity of the ungulates, carnivores and primates is reviewed. Special emphasis is focused on recent evolutionary history of longevity along the hominid ancestral-descendant sequence leading to modern man. Maximum life span potential (MLP) or the change in MLP is predicted in extinct species by (1) a phylogenetic analysis of the MLP of present living species and (2) an empirical equation using brain and body weight estimates from fossils. Both of these methods indicate MLP generally increased during mammalian evolution and at an extremely fast rate during the appearance of the hominid species. These results suggest that relatively few genetic alterations were necessary during the recent evolutionary history of man to significantly extend his innate ability to maintain mental and physical health. Much evidence indicates these genetic alterations principally involve regulatory genes, which control a conserved set of structural genes. Evolution of longevity in man could therefore be a result of simple changes in temporal and quantitative expression. Whether these genetic alterations result from mutational changes and/or chromosomal rearrangement cannot yet be evaluated.

Aging↗

A computer model of the evolution of specific maximum lifespan.

To answer the question of why organisms have evolved finite and specific maximum lifespans, I have built and experimentally studied a discrete-event simulation model of the evolution of lifespan. Through natural selection, the model evolves an apparent plateau in maximum lifespan, the height of which is a decreasing function of both the intensity of niche fluctuations and specific fecundity. Evolved lifespan is therefore finite (small and essentially constant over accessible time intervals) and specific. Experiments demonstrate that the plateau is not due to group selection. Instead, it occurs because the rate of increase of maximum lifespan by natural selection - in an environment presenting a finite probability that death will occur prior to reaching the genetically specified maximum - is a decreasing function of maximum lifespan itself and asymptotically approaches zero. This supports in part a class of existing hypotheses that finite lifespan is due to an equilibrium between weak selection, as in the model, and various lifespan-decreasing processes, which however were not simulated in the present experiments. Although the model shows that such counter processes are not strictly necessary for the evolution of finite and specific maximum lifespan, my interpretation of the model's correspondence to organic evolution does imply a counter process, a bias in random genetic drift toward shorter lifespan, that is more general than those previously hypothesized.

Biological Evolution↗

Evolution escapes rugged fitness landscapes by gene or genome doubling: the blessing of higher dimensionality.

Evolution on rugged landscapes ceases when a local maximum is attained. This poses the problem of how evolution could approach or attain a global maximum, especially for large genomes for which quasispecies are ineffective. I show how increasing the dimensionality of the landscape, which occurs every time there is a gene or higher order duplication (up to polyploidy), may solve this problem. Epistasis or complementarity between the duplicated genes provides an all uphill pathway towards the global maximum. The evolution of hemoglobin and other dimeric and tetrameric proteins provides a testable case, since fitness is readily defined.

Animals↗

Immunoproteins in evolution.

All vertebrates respond to antigenic challenge by specific cellular reactions and by producing circulating antibodies, and they also contain recognition molecules that are evolutionary relics of primitive non-immune recognition. One such ancient recognition molecule is C-reactive protein (CRP), a member of the pentraxin family, that has homologs occurring in species as diverse as vertebrates, tunicates and the horseshoe crab (an ancient arachnoid). This molecule has lectin-like properties, can act as an opsonin and interact with complement and cells in a manner paralleling immunoglobulins (Igs). The horseshoe crab lectin and CRP, although unrelated to Igs, share functional idiotopes with classical antibodies. This finding reflects either an evolutionary convergence or a mechanism of "mini gene insertion" that allows molecules of distinct evolutionary histories to react to the same ligands. Serum antibodies of all vertebrates are polydisperse in charge and are composed of polypeptide chains comparable in mass to those of mammalian light and heavy chains. Molecular genetic studies of placoderm derived vertebrates are incomplete but are sufficient to allow the conclusion that Igs of these species are specified by variable (V), joining (J) and constant (C) gene segments and that rearrangement are an essential feature for the generation of antibody diversity. Here we present new evidence following from the use of antibodies directed against synthetic joining region peptides as probes in the study of rearranging Igs in evolution, the use of recombinant DNA technology to study T cell receptor V beta genes in a goldfish genomic library and the isolation and characterization of a gene fragment specifying sandbar shark light chain C region. We reached the following conclusions: (1) J region segments are the most conserved in evolution, and this most probably reflects the essential requirements for these gene segments in the formation of intact Ig genes by rearrangement; (2) the framework segments of V regions are highly conserved in vertebrate evolution for both T cell receptors and classical Igs; and (3) although C region segments of light chains of lower vertebrates are homologous to their mammalian counterparts, the degree of conservation of C region structure in phylogeny is apparently less than that for V regions. Essentially, phylogenetic trees can be built using C regions but not V regions. We have identified a molecule of approximate mass of 26 kDa in the hemolymph of the tunicate, Boltenia ovipera, that is serologically cross-reactive with shark heavy chain and with J region peptides.(ABSTRACT TRUNCATED AT 400 WORDS)

Amino Acid Sequence↗

The evolution of the dorsal thalamus of jawed vertebrates, including mammals: cladistic analysis and a new hypothesis.

The evolution of the dorsal thalamus in various vertebrate lineages of jawed vertebrates has been an enigma, partly due to two prevalent misconceptions: the belief that the multitude of nuclei in the dorsal thalamus of mammals could be meaningfully compared neither with the relatively few nuclei in the dorsal thalamus of anamniotes nor with the intermediate number of dorsal thalamic nuclei of other amniotes and a definition of the dorsal thalamus that too narrowly focused on the features of the dorsal thalamus of mammals. The cladistic analysis carried out here allows us to recognize which features are plesiomorphic and which apomorphic for the dorsal thalamus of jawed vertebrates and to then reconstruct the major changes that have occurred in the dorsal thalamus over evolution. Embryological data examined in the context of Von Baerian theory (embryos of later-descendant species resemble the embryos of earlier-descendant species to the point of their divergence) supports a new 'Dual Elaboration Hypothesis' of dorsal thalamic evolution generated from this cladistic analysis. From the morphotype for an early stage in the embryological development of the dorsal thalamus of jawed vertebrates, the divergent, sequential stages of the development of the dorsal thalamus are derived for each major radiation and compared. The new hypothesis holds that the dorsal thalamus comprises two basic divisions--the collothalamus and the lemnothalamus--that receive their predominant input from the midbrain roof and (plesiomorphically) from lemniscal pathways, including the optic tract, respectively. Where present, the collothalamic, midbrain-sensory relay nuclei are homologous to each other in all vertebrate radiations as discrete nuclei. Within the lemnothalamus, the dorsal lateral geniculate nucleus of mammals and the dorsal lateral optic nucleus of non-synapsid amniotes (diapsid reptiles, birds and turtles) are homologous as discrete nuclei; most or all of the ventral nuclear group of mammals is homologous as a field to the lemniscal somatosensory relay and motor feedback nuclei of non-synapsid amniotes; the anterior, intralaminar and medial nuclear groups of mammals are collectively homologous as a field to both the dorsomedial and dorsolateral (including perirotundal) nuclei of non-synapsid amniotes; the anterior, intralaminar, medial and ventral nuclear groups and the dorsal lateral geniculate nucleus of mammals are collectively homologous as a field to the nucleus anterior of anamniotes, as are their homologues in non-synapsid amniotes. In the captorhinomorph ancestors of extant land vertebrates, both divisions of the dorsal thalamus were elaborated to some extent due to an increase in proliferation and lateral migration of neurons during development.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Chromosome evolution in retinoblastoma.

An attempt to reconstruct the chromosome evolution in 19 primary retinoblastoma tumors was made, taking into consideration the frequencies of the rearrangements for each case. The rearrangements were classified as early or late according to their frequency. Early chromosome rearrangements were +1q, +6p, -13/del(13q), -16/del(16q), -17/del(17p), and late rearrangements were (in decreasing order) -8, -17/del(17p), -22, +3/+3q, -4, -19, +1q, +7/+7q, -14, +21. Chromosome evolution was reconstructed by the analysis of parsimony using programs of phylogenetic inference, which belong to the computer package PHYLIP. Results of the reconstruction of chromosome evolution suggest that there are two types of retinoblastoma tumors which can be distinguished based on chromosome imbalance, and correlate with the presence or absence of additional copies of the short arm (p) of chromosome 6. Tumors with additional 6p chromosome had marked heterogeneity, with a high frequency of early as well as late chromosome changes. By contrast, tumors with a normal 6p complement had the fewest chromosome changes and, in particular, a lower frequency of late changes.

Biological Evolution↗

Emergence of adaptable systems and evolution of a translation device.

An over-all organizational framework for the origin of life is outlined and attempts for realization are given. Evolution can be described as a process resulting in an increase of "knowledge" where knowledge is the number of carriers of genetic information discarded, on the average, until the evolutionary state under consideration is reached. A model for the evolution of a translation device, a crucial event in the origin of life, is described in detail. Aggregates of short polynucleotide strands in a hairpin conformation play a major role in this model. Experimental evidence for the selectivity of aggregation supports the idea of aggregates as error filters. Chromatographic separation as selection process during chemical evolution supports the model of the early translation device leading to the origin of the genetic code.

Amino Acids↗

Evolution of anticodons.

Anticodons are trinucleotides in transfer RNA (tRNA) molecules. The latter carry amino acids for insertion into the polypeptide sequences of proteins during the translation of messenger RNA (mRNA) molecules. Messenger RNA molecules are transcribed from genes. Evolution of tRNA molecules has resulted in a set of anticodons for the 20 amino acids that are used in protein synthesis. This set of anticodons is slightly different in mitochondrial codes from the set that used in the nuclear "universal" code. Theories for the evolution of the code include frozen accident, doublet expansion, repeating triplets and coevolutionary distribution. The number of codons has always been fixed at 64 by mathematical rules, but because an anticodon may pair with more than one codon, the number of anticodons is only 54 in the universal code, is smaller in mitochondrial codes, and was probably even smaller in archetypal primitive codes. Evidence of anticodon evolution can be seen by comparing mitochondrial codes with the universal code. Codes used by very primitive organisms that are now extinct might have specified fewer amino acids than are now used.

Amino Acids↗

The origin of adaptation and dyssymmetry in the evolution of autocatalytic systems.

The evolution of open systems that include several autocatalytic processes in parallel or in series is mathematically analyzed. For the case of two reactions in parallel, such systems progressively and finally select the reaction pathway which involves the better autocatalyst. The effect of parameters influencing the rate of this evolution is discussed. Where catalysts are strictly equivalent, e.g. enantiomers in a symmetrical surrounding, the evolution is amplified by fluctuations and retains finally only one reaction pathway, if the autocatalytic rates of reactions are more than proportional to the catalyst concentrations. When including two reactions in series, these open systems are also able to give oscillations.

Catalysis↗

Insects as test systems for assessing the potential role of microgravity in biological development and evolution.

Gravity and radiation are undoubtedly the two major environmental factors altered in space. Gravity is a weak force, which creates a permanent potential field acting on the mass of biological systems and their cellular components, strongly reduced in space flights. Developmental systems, particularly at very early stages, provide the larger cellular compartments known, where the effects of alterations in the size of the gravity vector on living organisms can be more effectively tested. The insects, one of the more highly evolved classes of animals in which early development occurs in a syncytial embryo, are systems particularly well suited to test these effects and the specific developmental mechanisms affected. Furthermore, they share some basic features such as small size, short life cycles, relatively high radio-resistance, etc. and show a diversity of developmental strategies and tempos advantageous in experiments of this type in space. Drosophila melanogaster, the current biological paradigm to study development, with so much genetic and evolutionary background available, is clearly the reference organism for these studies. The current evidence on the effects of the physical parameters altered in space flights on insect development indicate a surprising correlation between effects seen on the fast developing and relatively small Drosophila embryo and the more slowly developing and large Carausius morosus system. In relation to the issue of the importance of developmental and environmental constraints in biological evolution, still the missing link in current evolutionary thinking, insects and space facilities for long-term experiments could provide useful experimental settings where to critically assess how development and evolution may be interconnected. Finally, it has to be pointed out that since there are experimental data indicating a possible synergism between microgravity and space radiation, possible effects of space radiation should be taken into account in the planning and evaluation of experiments designed to test the potential role of microgravity on biological developmental and evolution.

Animals↗

Mitosis and early meiosis in Tetrahymena pyriformis and the evolution of mitosis in the phylum Ciliophora.

The micronuclear mitotic spindle of Tetrahymena pyriformis contains several distinctive elements. The 150 or so continuous microtubules (MTs) form a peripheral sheath just inside the inner nuclear membrane while the kinetochore bundles traverse the center of the nucleoplasm. A new set of microtubules, the separation spindle, appears during the 10-fold nuclear elongation which occurs during the late anaphase. Both the separation spindle and peripheral sheath MTs are present in the macronucleus during macronuclear division but there are no definite kinetochore MTs. In the cresent stage of meiotic prophase both peripheral sheath and kinetochore MTs are present in the micronucleus. By using our own, and other workers', data in conjunction with the phylogenetic scheme for the ciliates which has been designed by Corliss (1974, 1975), we have attempted to trace out the evolutionary history of the various elements of the ciliate mitotic spindle. For example, the micronuclear separation spindle can be followed, from its point of origin within the primitive gymnostomes, throughout the phylum. The separation spindle of the macronucleus, by contrast, is lost at the level of the heterotrichs. Similarly, the anaphase breakdown and reconstitution of the nuclear envelope, which occurs in the primitive Loxodes magnus, can be followed up some phyletic branches, such as the heterotrichs, but in the suctorians and some other groups this feature seems to have been lost. Tracing the evolution of the ciliate mitotic spindle is made very difficult both by the incompleteness of the data and by what appear to be a number of cases of secondary reduction and parallel evolution. In general, however, the evolution of mitosis correlates well with the phylogeny of the ciliates constructed by Corliss and we consider this an independent substantiation of the general correctness of his phylogeny.

Biological Evolution↗

Symbiosis and the evolution of prokaryotes.

It is postulated, with support from kinetic modelling, that a succession of symbioses was the major process of evolution during the early stages of life. The process became less effective with the passage of time, while evolution by the natural selection of variants became more effective. The postulate may contribute usefully to discussions on the evolution of biochemical complexity and the structure of cells.

Biological Evolution↗

Stepwise molecular evolution of bacterial photosynthetic energy conversion.

The concept of continuity in molecular evolution implies a stepwise formation of metabolic systems and processes. In this manner, chemical and biological evolution have given rise, step by step, to such complicated systems as the photosynthetic apparatus and thus, such elaborate processes as photosynthesis in the living cell. Among currently living organisms, the bacteria contain a much less complex photosynthetic system than the algae and higher plants, which uniquely are capable fo splitting H2O. But also the bacterial system is a very highly evolved and sophisticated, membrane-bound apparatus for the transformation of light energy to other biologically useful energy forms. The study of its molecular evolution is here undertaken by the method of attempting to break down the system into its main components and functions in order to elucidate how they had originated and evolved, and how, by divergent and convergent evolutionary steps, the stage was set for the arrival of bacterial photophosphorylation.

Adenosine Triphosphatases↗

A test of the karyotypic fissioning theory of primate evolution.

Karyotypic fissioning theory has been put forward by a number of researchers as a possible driving force of mammalian evolution. Most recently, Giusto and Margulis (BioSystems, 13 (1981) 267-302) hypothesized that karyotypic fissioning best explains the evolution of Old World monkeys, apes, and humans. According to their hypothesis, hominoid karyotypes were derived from the monkey chromosome complement by just such a fissioning event. That hypothesis is tested here by comparing the G-banded chromosomes of humans and great apes with eight species of Old World monkeys. Five submetacentric chromosomes between apes and monkeys have identical banding patterns and nine chromosomes share the same pericentric inversion. Such extensive karyological similarities are not in accordance with, or predicted by karyotypic fissioning. Apparently, karyotypic fissioning is an extremely uneconomical model of chromosomal evolution. The strong conservation of banding patterns sometimes involving the retention of identical chromosomes indicates that ancient linkages of genes have probably been maintained through many speciation events.

Animals↗

Somatic kinetics or paroral membrane: which came first in ciliate evolution?

The ciliate species which lack a distinctive oral ciliature are considered to represent an ancestral state in ciliate evolution. Consequently, the somatic kineties composed of kinetids (kinetosomes plus cilia and associated fibrillar systems) are thought to be the ancestral ciliature. Results on stomatogenesis in 'gymnostomial ciliates' have shown that these ciliates probably have evolved from ancestors already equipped with an oral ciliature. Thus instead of the somatic, the oral ciliature may be regarded an ancestral. Based on these ideas a hypothesis on the evolution of the ciliate kinetome (assembly of all kinetids covering the body of a given ciliate) is presented. The first step in the evolution of the kinetome was the formation of a paroral membrane, a compound ciliary organelle lying along the right side of the oral area which historically but falsely is termed membrane. It was composed of kinetosomal dyads (dikinetids), derived from the kinetid of a dinoflagellate-like ancestor. From the beginning the paroral membrane was responsible for locomotion, ingestion and for the formation of a cytopharyngeal tube which the first ciliate probably had inherited from its flagellate ancestor. In the second step a first somatic kinety was formed from the right row of kinetosomes of the paroral membrane as a result of a longitudinal splitting of the paroral membrane and a subsequent migration of the forming kinety to the right into the somatic cortex. To increase the number of somatic kineties this process was repeated until the kinety produced first reached the left border of the oral area. By this step the locomotive and the nutritional functions were differentiated between somatic and oral structures. In a third step the adoral organelles were formed from somatic kinetids left of the oral area. The primitive type of stomatogenesis was a buccokinetal one derived from the mode the flagellate ancestor used to distribute its replicated kinetosomes to the offspring cells (buccokinetal means that at least parts of the oral anlage for the posterior offspring cell has its origin in the parental oral apparatus). This hypothesis, based on comparative studies on ciliate morphogenesis, is corroborated by molecular data from other laboratories.

Animals↗

Behavior, archetypes and the irreversibility of evolution.

There is a tendency in modern evolutionary theory to treat organisms as organized by genetic and developmental interactions, in such a way that these play a major role determining the direction of morphological evolution. Alternatively, in this paper behavior is considered as the most fundamental cohesive factor in the morphological evolution of animals. Some phenomena, such as the existence of evolutionary trends, the maintenance of architectural types in certain taxa, and the irreversibility of evolution are discussed in this light.

Adaptation, Physiological↗