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Biomedical subjects

L Margulis

Publications and source records attributed to L Margulis.

At least 37 records · Page 2Linked to original sources

The microbial community of Ophrydium versatile colonies: endosymbionts, residents, and tenants.

Ophrydium versatile is a sessile peritrichous ciliate (Kingdom Protoctista, class Oligohymenophora, order Peritrichida, suborder Sessilina) that forms green, gelatinous colonies. Chlorophyll a and b impart a green color to Ophrydium masses due to 400-500 Chlorella-like endosymbionts in each peritrich. Ophrydium colonies, collected from two bog wetlands (Hawley and Leverett, Massachusetts) were analyzed for their gel inhabitants. Other protists include ciliates, mastigotes, euglenids, chlorophytes, and heliozoa. Routine constituents include from 50-100,000 Nitzschia per ml of gel and at least four other diatom genera (Navicula, Pinnularia, Gyrosigma, Cymbella) that may participate in synthesis of the gel matrix. Among the prokaryotes are filamentous and coccoid cyanobacteria, large rod-shaped bacteria, at least three types of spirochetes and one unidentified Saprospira-like organism. Endosymbiotic methanogenic bacteria, observed using fluorescence microscopy, were present in unidentified hypotrichous ciliates. Animals found inside the gel include rotifers, nematodes, and occasional copepods. The latter were observed in the water reservoir of larger Ophrydium masses. From 30-46% of incident visible radiation could be attenuated by Ophrydium green jelly masses in laboratory observations. Protargol staining was used to visualize the elongate macronuclei and small micronucleus of O. versatile zooids and symbiotic algal nuclei. Electron microscopic analysis of the wall of the Chlorella-like symbiont suggests that although the Ophrydium zooids from British Columbia harbor Chlorella vulgaris, those from Hawley Bog contain Graesiella sp. The growth habit in the photic zone and loose level of individuation of macroscopic Ophrydium masses are interpretable as extant analogs of certain Ediacaran biota: colonial protists in the Vendian fossil record.

Animals↗

Life as a planetary phenomenon: the colonization of Mars.

Life is a planet-wide phenomenon in which its components incessantly move and interact. Life imperatively recycles its parts at the surface of the Earth in a chemical transformation and physical transport that depends utterly on the energy from a recent star, the Sun. Humanity, entirely dependent on other beings, plays a recent and relatively small part in the great phenomenon of life that transports and transforms the surface of the Earth. Our species accelerates but does not dominate the metabolism of the Earth system. Ironically, during the Apollo days of the sixties, fears were rampant that Martian or other extraterrestrial "germs" might "contaminate" our planet. After Viking, such fears are seen as the manifestation of cultural paranoia. The Viking missions complemented ground-based astronomical observation and yielded definitive evidence for the lack of life on the red planet. The Gaia hypothesis states that the surface temperature, composition of the reactive gases, oxidation state, alkalinity-acidity on today's Earth are kept homeorrhetically at values set by the sum of the activities of the current biota. Life, in other words, not only produces and maintains its immediate environment, but appears on Earth only as a planetary phenomenon. Since the natural tendency of all life is to grow exponentially to fill proximal volume, the question now "can life ecopoietically expand to Mars?" is entirely equivalent to the query of "can Gaia reproduce?"

Earth, Planet↗

Do prokaryotes contain microtubules?

In eukaryotic cells, microtubules are 24-nm-diameter tubular structures composed of a class of conserved proteins called tubulin. They are involved in numerous cell functions including ciliary motility, nerve cell elongation, pigment migration, centrosome formation, and chromosome movement. Although cytoplasmic tubules and fibers have been observed in bacteria, some with diameters similar to those of eukaryotes, no homologies to eukaryotic microtubules have been established. Certain groups of bacteria including azotobacters, cyanobacteria, enteric bacteria, and spirochetes have been frequently observed to possess microtubule-like structures, and others, including archaebacteria, have been shown to be sensitive to drugs that inhibit the polymerization of microtubules. Although little biochemical or molecular biological information is available, the differences observed among these prokaryotic structures suggest that their composition generally differs among themselves as well as from that of eukaryotes. We review the distribution of cytoplasmic tubules in prokaryotes, even though, in all cases, their functions remain unknown. At least some tend to occur in cells that are large, elongate, and motile, suggesting that they may be involved in cytoskeletal functions, intracellular motility, or transport activities comparable to those performed by eukaryotic microtubules. In Escherichia coli, the FtsZ protein is associated with the formation of a ring in the division zone between the newly forming offspring cells. Like tubulin, FtsZ is a GTPase and shares with tubulin a 7-amino-acid motif, making it a promising candidate in which to seek the origin of tubulins.

Animals↗

Harold Kirby's symbionts of termites: karyomastigont reproduction and calonymphid taxonomy.

Harold Kirby's brilliant principle of mastigont multiplicity is published here posthumously more than 40 years after it was written. He applies this principle to large multinucleate protist symbionts of termites in establishing the taxonomy of Calonymphids (Family Calonymphidae in Phylum Zoomastigina, Kingdom Protoctista). The nuclei and kinetosomes in these heterotrophic cells are organized into trichomonad-style mastigont units which reproduce independently of cytokinesis to generate nine new Calonympha and nineteen new Stephanonympha species. The total of six genera (Calonympha, Coronympha, Diplonympha, Metacoronympha, Snyderella and Stephanonympha, all symbionts of dry-wood-eating termites, Kalotermitidae) are recognized. With the aid of Michael Yamin, the distribution of all twenty-eight of Kirby's Calonympha and Stephanonympha species are tabulated. In italic type I have annotated this paper to be comprehensible to a wide readership of cell biologists, protistologists and those interested in insect symbionts. Although this extremely original and careful work was not finished when Kirby died suddenly in 1952, I deemed it important and complete enough to finally publish it so that it would not be lost to scientific posterity.

Animals↗

Composite, large spirochetes from microbial mats: spirochete structure review.

Phenomena previously unknown in free-living spirochetes are reported: large-sized cells with variable diameter (length to 100 microns, width between 0.4 and 3.0 microns), composite structure (smaller spirochetes inside larger ones), and positive phototropic behavior. These bacteria, Spirosymplokos, are compared with all other spirochete genera. The large spirochete, grown in mixed culture, was studied live and by transmission EM. The protoplasmic cylinder was replete with spherical granules 20-32 nm in diameter, and three to six periplasmic 26-nm flagella were inserted subterminally. Comparably granulated and flagellated small spirochetes were located inside the protoplasmic cylinder and in the periplasm of the large ones. When exposed to air, movement became erratic, protoplasmic cylinders retracted to lie folded inside the outer membrane, and refractile membranous structures formed. From one to four structures per still-moving spirochete were seen. Spirosymplokos was enriched from laboratory samples exposed to oxygen-rich and desiccating, but not dry, conditions for at least 4 mo after removal of microbial mat from the field.

Microscopy, Electron↗

Spirosymplokos deltaeiberi nov. gen., nov. sp.: variable-diameter composite spirochete from microbial mats.

Large (up to 100 micrometers long), loosely coiled, free-living spirochetes with variable diameters (from 0.4 to 3 micrometers in the same cell) were seen at least 40 times between August 1990 and January 1993. These spirochetes were observed in mud water and enrichment media from highly specific habitats in intertidal evaporite flats at three disjunct localities, one in Spain and two in Mexico. All three are sites of commercial saltworks. Associated with Microcoleus chthonoplastes the large spirochetes from Spain display phototaxis and a composite organization. Shorter and smaller-diameter spirochetes are seen inside both healthy and spent periplasm of larger ones. Small spirochetes attached to large ones have been observed live. From two to twelve spirochete protoplasmic cylinders were seen inside a single common outer membrane. A distinctive granulated cytoplasm in which the granules are of similar diameter (20-32 nanometers) to that of the flagella (26 nanometers) was present. Granule diameters were measured in thin section and in negatively-stained whole-mount preparations. Based on their ultrastructure, large size, variable diameter, number of flagella (3 to 6), and phototactic behavior these unique spirochetes are formally named Spirosymplokos deltaeiberi. Under anoxic (or low oxygen) conditions they formed blooms in mixed culture in media selective for spirochetes. Cellobiose was the major carbon source in 80% seawater, the antibiotic rifampicin was added, mat from the original field site was present and tubes were incubated in the light at from 18-31 degrees C. Within 1-2 weeks populations of the large spirochete developed at 25 degrees C but they could not be transferred to fresh medium.

Environmental Microbiology↗

Origins of species: acquired genomes and individuality.

Entire genomes with their accompanying protein synthetic systems are transferred throughout the biosphere primarily as bacteria and protists which become symbionts as they irreversibly integrate into pre-existing organisms to form more complex individuals. Individualization is stabilized by simultaneous transmission of once-separate heterologous genetic systems. The origin of new species is hypothesized to correlate with the acquisition, integration and subsequent inheritance of such acquired microbial genomes. These processes were recognized by Mereschkovsky ("Symbiogenesis" in Russian, 1909) and by Wallin ("Symbionticism", see p. 181, this issue).

Animals↗

The 'tubulin-like' S1 protein of Spirochaeta is a member of the hsp65 stress protein family.

A 65-kDa protein (called S1) from Spirochaeta bajacaliforniensis was identified as 'tubulin-like' because it cross-reacted with at least four different antisera raised against tubulin and was isolated, with a co-polymerizing 45-kDa protein, by warm-cold cycling procedures used to purify tubulin from mammalian brain. Furthermore, at least three genera of non-cultivable symbiotic spirochetes (Pillotina, Diplocalyx, and Hollandina) that contain conspicuous 24-nm cytoplasmic tubules displayed a strong fluorescence in situ when treated with polyclonal antisera raised against tubulin. Here we summarize results that lead to the conclusion that this 65-kDa protein has no homology to tubulin. S1 is an hsp65 stress protein homologue. Hsp65 is a highly immunogenic family of hsp60 proteins which includes the 65-kDa antigens of Mycobacterium tuberculosis (an active component of Freund's complete adjuvant), Borrelia, Treponema, Chlamydia, Legionella, and Salmonella. The hsp60s, also known as chaperonins, include E. coli GroEL, mitochondrial and chloroplast chaperonins, the pea aphid 'symbionin' and many other proteins involved in protein folding and the stress response.

Amino Acid Sequence↗

Gaia and the colonization of Mars.

The Gaia hypothesis states that the atmosphere, hydrosphere, surface sediments, and life on Earth behave dynamically as a single integrated physiological system. What has been traditionally viewed as the passive environment is a highly active, integral part of the gaian system. Aspects of the surface temperature and chemistry are regulated by the sum of life, the biota. Formulated first by James E. Lovelock, in the late 1960s, the Gaia hypothesis has been in the scientific literature for more than 25 years. Because of its properties of exponential growth and propagation, life is a powerful geologic force. A useful aspect of the Gaia idea is that it requires integration of scientific disciplines for the study of Earth. The recently touted Earth system science is broadly parallel with the gaian concept of the physiochemical regulation of Earth's surface. We discuss here, in a gaian context, the colonization of Mars by Earth organisms. Although colonizing Mars may be impossible, its accomplishment would be exactly equivalent to "the reproduction of Gaia by budding."

Atmosphere↗

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↗

Kingdoms in turmoil.

How should the world's living organisms be classified? Into how many kingdoms should they be grouped? Scientists have been grappling with these questions since the time of Aristotle, drawing on a broad base of biological characteristics for clues. The fossil record, visible traits of living organisms and, more recently, results from cell biology have all shaped theories of biological classification. But last year a new and controversial concept emerged: a classification of life based solely on molecular traits. The focal point of the controversy is a tree of life, or "phylogeny", devised by Carl Woese of the University of Illinois, Otto Kandler of the University of Munich and Mark Wheelis of the University of California. The tree is unusual because, unlike all previous schemes, it is constructed solely from biochemical data such as DNA sequences rather than a range of different organism characteristics. But that is not all. The scheme also challenges the idea that life on Earth is best divided into five kingdoms, with the main split being between bacteria and all other organisms. Woese and his colleagues create three main groupings by dividing the bacteria in two and unifying all other organisms.

Animals↗

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↗

Cristispira from oyster styles: complex morphology of large symbiotic spirochetes.

Crystalline styles (digestive organs) of bivalve mollusks provide the habitat for highly motile bacteria. Styles from freshly-collected oysters, Crassostrea virginica, were studied by electron microscopy; Cristispira spirochetes were abundant in these organs. Detailed study reveals these spirochetes to be among the most complex prokaryotic cells known. More than 600 periplasmic flagella and an adhering outer lipoprotein membrane (e.g., a 270 degrees sillon) form the ultrastructural basis for the "crista," first described by light microscopy. Unique rosette structures corresponding to the "chambers" or "ovoid inclusions" of light microscopy were detected at the periphery of all protoplasmic cylinders. Polar organelles and linearly aligned flagellar insertions are conspicuous. In size and complexity, Cristispira more resembles Pillotina, Diplocalyx, Clevelandina and Hollandina (large spirochetes symbiotic in termites) than it does Treponema. Cristispira pectinis (Gross, 1910), the type species; Spirillum ostrea (Noguchi, 1921); and another, less frequent bacterial symbiont are the predominant inhabitants of the dense style matrix. The ultrastructure of the spirillum and an electron micrograph of the third bacterium are shown.

Animals↗

Mobilifilum chasei: morphology and ecology of a spirochete from an intertidal stratified microbial mat community.

Spirochetes were found in the lower anoxiphototrophic layer of a stratified microbial mat (North Pond, Laguna Figueroa, Baja California, Mexico). Ultra-structural analysis of thin sections of field samples revealed spirochetes approximately 0.25 micrometer in diameter with 10 or more periplasmic flagella, leading to the interpretation that these spirochetes bear 10 flagellar insertions on each end. Morphometric study showed these free-living spirochetes greatly resemble certain symbiotic ones, i.e., Borrelia and certain termite spirochetes, the transverse sections of which are presented here. The ultrastructure of this spirochete also resembles Hollandina and Diplocalyx (spirochetes symbiotic in arthropods) more than it does Spirochaeta, the well known genus of mud-dwelling spirochetes. The new spirochete was detected in mat material collected both in 1985 and in 1987. Unique morphology (i.e., conspicuous outer coat of inner membrane, large number of periplasmic flagella) and ecology prompt us to name a new free-living spirochete.

Animals↗

Kingdom Animalia: the zoological malaise from a microbial perspective.

Pain and cognitive dissonance abounds amongst biologists: the plant-animal, botany-zoology wound has nearly healed and the new gash--revealed by department and budget reorganizations--is "molecular" vs. "organismic" biology. Here I contend that resolution of these tensions within zoology requires that an autopoietic-gaian view replace a mechanical-neodarwinian perspective; in the interest of brevity and since many points have been discussed elsewhere, rather than develop detailed arguments I must make staccato statements and refer to a burgeoning literature. The first central concept is that animals, all organisms developing from blastular embryos, evolved from single protist cells that were unable to reproduce their undulipodia. The second points to the usefulness of recognizing the analogy between cyclically established symbioses and meiotic sexuality.

Animal Population Groups↗