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Biomedical subjects

L Margulis

Publications and source records attributed to L Margulis.

At least 19 recordsLinked to original sources

Biodiversity: molecular biological domains, symbiosis and kingdom origins.

The number of extant species of organisms is estimated to be from fewer than 3 to more than 30 x 10(6) (May, 1992). Molecular biology, comparative genetics and ultrastructural analyses provide new insights into evolutionary relationships between these species, including increasingly precise ideas of how species and higher taxa have evolved from common ancestors. Accumulation of random mutations and large macromolecular sequence change in all organisms since the Proterozoic Eon has been importantly supplemented by acquisition of inherited genomes ('symbiogenesis'). Karyotypic alterations (polyploidization and karyotypic fissioning) have been added to these other mechanisms of species origin in plants and animals during the Phanerozoic Eon. The new evolution concepts (coupled with current rapid rates of species extinction and ignorance of the extent of biodiversity) prompted this analysis of the field of systematic biology and its role in the reorganization of extant species into higher taxa. Two superkingdoms (= Domains: Prokaryotae and Eukaryotae) and five kingdoms (Monera = Procaryotae or Bacteria; Protoctista: algae, amoebae, ciliates, foraminifera, oomycetes, slime molds, etc.; Mychota: 'true' fungi; Plantae: one phylum (division) of bryophytes and nine phyla of tracheophytes; and Animalia) are recognized. Two subkingdoms comprise the monera: the great diverse lineages are Archaebacteria and Eubacteria. The criteria for classification using molecular, ultrastructural and genetic data for this scheme are mentioned. For the first time since the nineteenth century, logical, technical definitions for each group are given with their time of appearance as inferred from the fossil record in the primary scientific literature. This classification scheme, which most closely reflects the evolutionary history, molecular biology, genetics and ultrastructure of extant life, requires changes in social organization of biologists, many of whom as botanists and zoologists, still behave as if there were only two important kingdoms (plants and animals).

Animal Population Groups

Search for eukaryotic motility proteins in spirochetes: immunological detection of a tektin-like protein in Spirochaeta halophila.

The serial endosymbiotic theory (SET) for the spirochete origin of undulipodia (cilia and eukaryotic flagella) predicts that a greater number of axonemal proteins will show homology to spirochete than to other bacterial proteins. To continue testing, the SET proteins associated with eukaryotic motility (tektin, centrin and calmodulin) were sought in Spirochaeta halophila. Western blot immunological detection techniques (for tektin and centrin) and two-dimensional gel analysis (for calmodulin) were used. Tektins are filamentous proteins extending the length of the axoneme in sperm tails and other undulipodia. Whole cell extracts of S. halophila were probed with antibodies made against three sea urchin (Lytechinus pictus) sperm axonemal tektins (tektins A, B, and C). In the spirochetes, one tektin-like protein was detected as a band on Western blots (a C tektin.) An antibody made against Lytechinus pictus sperm tail axonemes, affinity-purified against the C tektin of another sea urchin, Stronglyocentrotus purpuratus, bound to a 30 kDa protein from Spirochaeta halophila. The C tektin epitope was not detected in Escherichia coli. Both the poly- and monoclonal anti-centrin antibodies cross-reacted with multiple proteins in the control alga (Tetraselmis striata) and in the putatively negative control bacterium E. coli. No cross reaction was seen between any anti-centrin antibody and S. halophila. Neither did a two-dimensional gel analysis reveal the presence of calmodulin in these spirochetes or in the two other prokaryotes tested (Spiroplasma citri, Acholeplasma laidlawii). Although neither centrin nor calmodulin were detected, a 30 kDa tektin-like protein apparently is present in these spirochetes.

Biological Evolution

The evolutionary transition from RNA to DNA in early cells.

The evolution of genetic material can be divided into at least three major phases: first, genomes of "nucleic acid-like" molecules; secondly, genomes of RNA; and finally, double-stranded DNA genomes such as those present in all contemporary cells. Using properties of nucleic acid molecules, we attempt to explain the evolutionary transition from RNA alone as a cellular informational macromolecule prior to the evolution of cell systems based on double-stranded DNA. The idea that ribonucleic acid-based cellular genomes preceded DNA is based on the following: (1) protein synthesis can occur in the absence of DNA but not of RNA; (2) RNA molecules have some catalytic properties; (3) the ubiquity of purine and pyridine nucleotide coenzymes as well as other similar ribonucleotide cofactors in metabolic pathways; and (4) the fact that the biosynthesis of deoxyribonucleotides always proceeds via the enzymatic reduction of ribonucleotides. The "RNA prior to DNA" hypothesis can be further developed by understanding the selective pressures that led to the biosynthesis of deoxyribose, thymine, and proofreading DNA polymerases. Taken together these observations suggest to us that DNA was selected as an informational molecule in cells to stabilize earlier RNA-protein replicating systems.(ABSTRACT TRUNCATED AT 250 WORDS)

Biological Evolution

Heliobacterium and the origin of chrysoplasts.

Chrysoplasts, golden-yellow and brown photosynthetic membrane-bounded plastids, photosynthetic organelles of algae such as phaeophytes (brown seaweeds), bacillariophytes (diatoms) and chrysophytes (golden-yellow algae including silicoflagellates), are hypothesized to have originated from brownish photoheterotrophic bacteria such as the newly discovered anaerobic nitrogen-fixing Heliobacterium. The consequences of this hypothesis as well as the data required to verify or disprove it are presented.

Biological Evolution

Desiccation resistance and contamination as mechanisms of gaia.

The gaia hypothesis, formulated by J.E. Lovelock, asserts the composition of the reactive gases, the oxidation-reduction state and the temperature of the lower atmosphere of the planet Earth are actively regulated by the biota. Lovelock and Watson, using highly simplified mathematical models, have shown that the modulation of atmospheric temperature can be achieved by exponentially growing populations of differently colored organisms ("dark and light daisies"). It is more likely that the modulation of atmospheric gas composition is based on the colligative properties of exponentially growing mixed populations of microorganisms rather than on "daisies". Exponential growth of one population of microorganisms leads to gaseous and other metabolic products released to the environment, which favor the exponential growth of different populations, each with their own unique emissions. Extremely high densities of mixed populations of microorganisms ensue. These populations form structured microbial communities composed of members in varying states of activity. Growth potential of metabolically diverse populations most likely provides the basis for the responsiveness of the biota to changing environments. We have attempted to measure an aspect of the growth potential and diversity of one microbial community, that from a flat laminated microbial mat dominated by the cyanobacterium, Microcoleus. Microbial mat samples collected at yearly intervals between 1977 and 1982 were allowed to dry. Subsamples were revived under laboratory conditions by rewetting, and the resulting complex microbial populations were analyzed. Greater than 10(4) viable organisms per ml were estimated to be present in the desiccated samples. Only a portion of the diverse community could be characterized. There were at least 115 different types of desiccation resistant microorganisms present in these samples, primarily bacteria. However, more than a dozen types of rather uncommon fungi and protoctists were removed from naturally desiccated material. Several did not fit descriptions of previously known species or strains. Neither animals nor plants were recovered from these tiny samples. We present minimal diversity estimates for microorganisms in both the desiccated samples and the corresponding fresh laminated microbial mat community from which they were taken. We found that many organisms in laboratory samples capable of immediate growth from a desiccated state were not common components of growing mats in the field.(ABSTRACT TRUNCATED AT 400 WORDS)

Atmosphere

Order amidst animalcules: the Protoctista kingdom and its undulipodiated cells.

The eukaryotic microorganisms have always been studied and described in the context of Zoology (as tiny animals), Botany (as tiny plants), Mycology (as water molds) or Microbiology (as disease agents). Because their extent (as a group of about 200,000 species) and differences from the rest of life (worthy of Protoctista kingdom status) only recently has been recognized, their classification within the Plant and Animal kingdoms remains in scandalous disarray. Admitting this problem, this paper represents a step toward achieving nomenclatural order based on ultrastructural, biochemical and evolutionary considerations.

Animals

Karyotypic fission theory and the evolution of old world monkeys and apes.

The karyotypes of living catarrhines are correlated with the current concepts of their fossil record and systematic classification. A phylogeny, beginning at the base of the Oligocene, for those animals and their chromosome numbers is presented. Todd's (1970) theory of karyotypic fissioning is applied to this case - three fissioning events are hypothesized. A late Eocene event (the primary catarrhine fissioning) is hypothesized to underlie the diversification of the infraorder Catarrhini into its extant families, the second fissioning underlies the radiation of the pongidae/Hominidae in the Miocene and the third accounts for the high chromosome numbers (54 - 72) and the Neogene(Miocene-Pliocene-Pleistocene) radiation of members of the genus Cercopithecus. Published catarrhine chromosome data, including that for "marked" chromosomes (those with a large achromatic region that is the site for ribosomal RNA genes) are tabulated and analysed. The ancestral X chromosome is always retained in the unfissioned metacentric state. The Pongidae/Hominidae have 15 pairs of mediocentric chromosomes that survived the second fissioning whereas the other chromosomes (besides the X) are thought to be fission-derived acrocentrics. Both the detailed karyology and the trend from low to high numbers is best interpreted to support Todd's concept of adaptive radiations correlated with karyotypic fissioning in ancestral populations.

Animals

Hydra viridis: transfer of metabolites between Hydra and symbiotic algae.

"Back transfer" of metabolites from food to endosymbiotic algae in the digestive cells of Hydra viridis was demonstrated. Brine shrimp nauplii labeled with tritiated precursors of protein and nucleic acids (DNA and RNA) were fed to light and dark grown hydras. The fate of the label after a single feeding with radioactive material in hydra and algal fractions was followed by scintillation counting and autoradiographic techniques. Labeled thymidine was incorporated into DNA in both light- and dark-grown hydras. Although the symbiosis persists indefinitely in hydras in darkness (7-10 days) the number of algae per cell is reduced. Tritiated orotic acid and tritiated uridine, RNA precursors, were incorporated into peptides and proteins, and to a lesser extent into simple sugars, oligosaccharides, and oligonucleotides in hydra and algal fractions. Thus the metabolites of the brine shrimp food are available to both partners. A decrease over time in label introduced as 3H-orotic acid and 3H-uridine and incorporated into hydra RNA is compensated for by an increase in label in the algae, implying competition for constant quantities of metabolites from the single feeding. Although food availability, light, number of algae per cell, and other factors influence the quantity and rate of nutrient transfer between the partners, in both light and dark grown hydras the amount of "back transfer" of metabolites to the symbiotic algae is impressive.

Animals

A red Beneckea from Laguna Figueroa, Baja California.

A new bacterium (nitrate-respiring, prodigiosin-producing, marine curved rod with a sheathed flagellum) has been isolated from anaerobic mud underlying a microbial meat. This brightly pigmented red bacterium, referred to as strain BV1 (Baja California vibrio, isolate 1) was taken from a closed, hypersaline basin at Laguna Figueroa (or Laguna Mormona), Baja California de Norte, Mexico. It is closely related to the recently described Beneckea gazogenes (Harwood, 1978), which was isolated from an estuarine habitat, the Sippewissett salt marsh at Woods Hole, Massachusetts, U.S.A. Strain BV1 and B-gazogenes are both oxidase positive facultative anaerobic curved rods which bear a single polar flagellum, and synthesize the red-orange tri-pyrrole pigment prodigiosin. The bacterium, which fluoresces green when excited with UV light (lambda = 455 nm), deposits pigment extracellularly in copious quantities. The extracellular pigment deposits fluoresce red-yellow. Both BV1 and B. gazogenes are able to grow utilizing xylose, cellobiose or arabinose, products of plant biosynthesis, as sole carbon sources. BV1 differs from B. gazogenes in cell size, pattern of pigment production, nutritional characteristics, the ability to perform anaerobic respiration using nitrate as a terminal electron acceptor, sensitivity to a newly discovered lytic phage and to the antibiotic vibriostat O/129.

Anaerobiosis

Bacterial resistance to ultraviolet irradiation under anaerobiosis: implications for pre-phanerozoic evolution.

The concept that low concentrations of atmospheric oxygen and consequent unattenuated ultraviolet irradiation limited the emergence of Phanerozoic life, the Berkner-Marshall hypothesis, is no longer tenable. Anaerobic bacteria, which probably evolved far earlier than Metazoa, were irradiated in a special chamber under strictly anaerobic conditions. Both intrinsic resistance and photoreactivation by visible light were discovered in obligately and facultatively anaeroboc microbes. Atmospheric scientists have shown that small amounts of oxygen would have limied pre-Phanerozoic surface ultraviolet irradiation to fluxes well below those used in the anaerobic experiments described. Since adequate ultraviolet protection mechanisms evolved early, the late Proterozoic appearance of Metazoa probably was not related to high fluxes of solar ultraviolet radiation.

Anaerobiosis

The symbiotic microbial community of the Sonoran Desert termite: Pterotermes occidentis.

Pterotermes occidentis is a large, obscure, very primitive dry wood termite limited to the Sonoran desert of North America. Its development, caste system and behavior are discussed in relation to aspects of its ecology. What appear to be single wood-digesting individual termites are in fact insect hosts which harbor extensive microbial communities estimated to contain more than 40 interacting species. All healthy, wood-eating Pterotermes contain these densely populated communities of highly motile symbionts suggesting that species diversity is necessary for termite survival. A morphological catalogue of the major hindgut microbes is presented here. From 10(7) to 10(11) bacteria and from 10(3) to 10(6) protists/ml of hindgut fluid are found in an average nymph or larva. The species of four wood-ingesting mastigotes (genera Trichonympha, Metadevescovina, Tricercomitus and Microrhopalodina) and nearly 30 morphologically distinct types of bacteria are described. The description of this morphological diversity observed with light, scanning and transmission electron microscopy is a prerequisite to functional analyses of the symbioses. Since Pterotermes is easily maintained in the laboratory and is so large it is hoped that it will provide a useful model for symbiosis research.

Animals

Evolutionary prerequisites for early Phanerozoic calcareous skeletons.

This paper develops the concept that the dramatic appearance of calcareous skeletons in the Lower Cambrian is directly related to the origin of refined mechanisms of intracellular modulation of calcium ion concentration. An homologous family of calcium modulated proteins has recently been discovered. These proteins contain "EF hands", involved in the maintenance of low concentrations of intracellular calcium and the informational use of calcium ion flow (Kretsinger, 1977). The evolution of this specialized calcium physiology, especially in muscle systems, coupled with natural selection by predators are identified as some of the preadaptations for the impressive Tommotian diversification of calcified metazoans. The distribution of calcium biominerals in the phyla of the five kingdoms and the time of first appearance of calcareous mineralization in the fossil record are tabulated. Macroscopic calcified hard parts apparently required the prior evolution of certain cell, tissue and organ system physiologies which are briefly discussed here.

Animals

Undulipodia, flagella and cilia.

The term flagella is ambiguous. It refers to bacterial structures composed of flagellin protein and to eukaryotic structures composed of microtubule proteins and ATPase (tubulin and dynein). The fact that cilia are nearly identical to eukaryotic flagella and have nothing in common with prokaryotic flagella is not apparent from the terminology. It is proposed that the 30-year old suggestion of Smagina and reiterated by Kuznicki and others, be adopted: that cilia and eukaryotic flagella be called "undulipodia." The term flagella ought to be restricted to prokaryotic organelles, bacterial flagella and spirochaete axial filaments: solid structures composed of flagellin which protrude through the plasma membrane and lack intrinsic motility throughout their length. Undulipodia are defined as intrinsically motile intracellular structures showing a 9-fold symmetry in the pattern of arrangement of 24 nm diameter microtubules. They are limited to eukaryotes, members of the protoctist, animal and plant kingdoms.

Animals

Possible evolutionary significance of spirochaetes.

Large symbiotic spirochaetes of the family Pillotaceae (e.g. pillotinas) are found in dry wood and subterranean termites (Hollande & Garagozlou 1967). These morphologically distinctive spirochaetes comprise several genera. Some of them contain microtubules within their protoplasmic cylinders. They demonstrate a variety of relations with their termite and protist hosts. Some are free-living within the lumen of the intestine, some tend to be associated with filamentous and other bacteria, some are found regularly coursing between the numerous undulipodia ( = eukaryotic flagella, cilia, and other (9 + 2) organelles of motility) of hypermastigotes and polymastigotes. Still other smaller termite spirochaetes are regularly attached to protists via specialized attachment sites. Some even form motility symbiosis with their host protists. The analogy between the behaviour of host-associated spirochaetes and the possible steps in the origin of the undulipodia and mitotic system of eukaryotes is discussed briefly.

Biological Evolution

An ultraviolet light induced bacteriophage in Beneckea gazogenes.

An ultraviolet light induced prophage has been discovered in the red pigmented marine vibrio Beneckea gazogenes. Two spontaneously derived pigment mutants, one forming pink colonies and one lacking pigment and forming white colonies, were also irradiated. The presence of pigment was not related to phage induction; uv-induced cell lysis occurred in wildtype and mutant strains at the same dosages. Lysis was not prevented or retarded by exposure after irradiation to visible light indicating the phenomenon was not photoreactivable. Electron micrographs of the 'T-like' B. gazogenes phage are shown. A second beneckea was isolated form the anaerobic zone of cyanobacterial mats growing in the hypersaline environment of Laguna Mormona, Baja California. The Baja beneckea does not harbor a uv inducible prophage and is resistant to the B. gazogenes phage under all conditions tested.

Bacteriolysis

The Viking Mission: implications for life on Mars.

The results of the Viking Biology experiments are best explained by non-biological phenomena: The interaction of the reagents with the materials comprising the regolith. Conditions of water activity, temperature, availability of carbon sources and others in most regions of the planet are too extreme for survival and growth of any known Earth microorganisms. Although the possibility persists that some very unusual form of life is somewhere on that planet the evidence is best interpreted as negative. Even though there is no evidence for current life on Mars, whether or not life ever originated there is not known.

Carbon