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21 Bacteria: The Deinococci and Nonproteobacteria

Gram-Negatives

 

CHAPTER OVERVIEW

This chapter is devoted to nine of the more interesting and important eubacterial groups from volumes 1 and 5 of the 2nd edition of Bergey's Manual of Systematic Bacteriology. Though the organization and perspective of the 2nd edition is used, the description of each group in the current edition is summarized. Where appropriate the distinguishing characteristics, morphology, reproduction, physiology, metabolism, and ecology of each group is included. The taxonomy of each major group is summarized and representative species are discussed.

CHAPTER OBJECTIVES

After reading this chapter you should be able to:

CHAPTER OUTLINE

  1. Aquifex and Thermotogas
    1. Aquifex (e.g., Aquifex pyrophilus) and its relatives such as Hydrogenobacter are members of

the deepest or oldest branch of the eubacteria

      1. Hyperthermophile with a temperature optimum of 85° C and a maximum of 95° C
      2. Autotroph that generates energy by oxidizing donors such as hydrogen, thiosulfate, and sulfur with oxygen as the acceptor

B. Thermotoga - second oldest branch of the eubacteria

      1. Hyperthermophile with an optimum of 80° C and a maximum of 90° C
      2. Gram-negative rod with an outer sheath-like envelope (like a toga) that can balloon out from the ends of the cell
      3. Grows in active geothermal areas - marine hydrothermal vents and terrestrial solfataric springs
      4. Chemoheterotroph with a functional glycolytic pathway
      5. Can grow anaerobically on carbohydrates and protein digests
      6. Consists of at least seven genera in two orders

II. The Deinococci

    1. One familywith three genera
    2. Spherical or rod-shaped with distinctively different 16S rRNA
    3. Associated in pairs or tetrads
    4. Aerobic, mesophilic, catalase positive, and usually able to produce acid from few sugars
    5. They stain gram-positive but have a layered cell wall and an outer membrane like the gram-negative bacteria
    6. They have L-ornithine in their peptidoglycan; lack teichoic acid; have a plasma membrane with large amounts of palmitoleic acid rather than phosphatidylglycerol phospholipids
    7. Extraordinarily resistant to desiccation and radiation: can survive 3-5 million rad whereas 100 rad can be lethal to humans
    8. Can be isolated from ground meat, feces, air, fresh water, and other sources but their natural habitat is not known
    9. Have an unusual ability to repair chromosomal damage (even fragmentation) that probably accounts for their ability to resist radiation
  1. Photosynthetic Bacteria
    1. Three groups: purple bacteria, green bacteria, and cyanobacteria
    2. Green and purple bacteria use anoxygenic photosynthesis
      1. Use bacteriochlorophylls
      2. Use H2O and other reduced electron donors
      3. Use wavelengths of light that enable them to grow at deeper water levels

C. Cyanobacteria use oxygenic photosynthesis

      1. Use chlorophyll a
      2. Use H2O as an electron donor
      3. Grow primarily at the surface of bodies of water

D. Divided into two groups (oxygenic and anoxygenic) in the 1st edition of Bergey's Manual;

however, the 2nd edition divides them into six groups

E. The Chloroflexi

      1. Has both photosynthetic and nonphotosynthetic members
      2. Chloroflexus is the major representative of the photosynthetic green nonsulfur bacteria
      1. Filamentous, gliding, thermophilic bacteria
      2. Often isolated from neutral to alkaline hot springs
      3. Grows in the form of orange-reddish mats
      4. Metabolism is similar to that of the purple nonsulfur bacteria
      5. Can carry out anoxygenic photosynthesis with organic compounds as carbon sources or can grow aerobically as a chemoheterotroph
      6. Not closely related to any eubacterial group

F. Chlorobia

      1. Green sulfur bacteria
      2. Obilgately anaerobic photolithoautotrophs that use hydrogen sulfide, elemental sulfur and hydrogen as electron sources
      3. Elemental sulfur produced by sulfide oxidation is deposited outside the cell
      4. Photosynthetic pigments are located in a ellipsoidal vesicles called chlorosomes (chlorobium vesicles) which are attached to the plasma membrane but not continuous with it
      5. Chlorosome membrane is not a normal lipid bilayer
      6. Chlorosomes have accessory bacteriochlorophylls but the reaction center bacteriochlorophyll is located in the plasma membrane
      7. They lack flagella and are nonmotile but some species have gas vesicles to adjust their depth for adequate light and hydrogen sulfide; species without gas vesicles are found in sulfide-rich muds at the bottom of lakes and ponds
      8. Morphologically diverse; rods cocci, or vibrios; grow singly, in chains, or in clusters; grass-green or chocolate-brown in color

G. Prochloron and Cyanobacteria

      1. The cyanobacteria are the largest and most diverse group of photosynthetic bacteria
      2. The 1st edition of Bergey's Manual describes 34 genera in some detail
      3. Procaryotes but the photosynthetic system resembles that of eucaryotes
      1. They have chlorophyll a and photosystem II
      2. Carry out oxygenic photosynthesis
      1. Photosynthetic pigments are in thylakoid membranes lined with particles called
      2. phycobilisomes - contain the phycobilin pigment phycocyanin which transfers energy to

        photosystem II; some species are red-brown and contain the pigment phycoerythrin

      3. They are photolithoautotrophs but some can grow slowly in the dark as
      4. chemoheterotrophs

      5. Some species can carry out anoxygenic photosynthesis if in an anaerobic environment
      6. They have typical procaryotic structures with a gram-negative cell wall
      7. Vary greatly in shape and appearance; unicellular; colonies of many shapes; filaments
      8. called trichomes - rows of bacterial cells that are in close contact with one another

        over a large area

      9. Many filamentous cyanobacteria can fix atmospheric nitrogen in special cells called
      10. heterocysts

      11. They often use gas vesicles to move vertically in the water
      12. Filamentous cyanobacteria have a gliding motility
      13. Some marine species lack flagella but are able to move by an unknown mechanism
      14. Reproduce by binary fission, budding fragmentation and multiple fission
      15. Fragmentation generates small motile filaments called hormogonia
      16. Some species develop akinetes, which are thick-walled resting cells that are resistant to
      17. desiccation

      18. Some cyanobacteria use multiple fission to produce small reproductive cells called
      19. baeocytes

      20. Some cyanobacteria form linear filaments; others produce branches or aggregates
      21. Tolerant of environmental extremes; thermophilic species can grow at temperatures up
      22. to 75° C

      23. Successful at establishing symbiotic relationships - photosynthetic partner in most

lichen associations

17. The order Prochlorales is placed with the cyanobacteria in the 2nd edition of Bergey's

Manual

18. Prochlorales are oxygenic phototrophic procaryotes that have both chlorophyll a and

chlorophyll b but lack phycobilins

      1. Similar to, but different from, cyanobacteria
      2. Three recognized genera
      1. Prochloron - extracellular symbiont on the surface or within the cloacal
      2. cavity of marine colonial ascidan invertibrates

      3. Prochlorothrix - free living
      4. Prochlorococcus - has a modified chlorophyll a and a -carotene rather than b -carotene
  1. The Planctomycetes
    1. Order Planctomycetales
      1. Spherical or oval, budding eubacteria with distinctive crateriform structures (pits) in their
      2. walls

      3. In two genera, Gemmata and Pirullela, the nuclear body is membrane bounded,

something that is not seen in other procaryotes

3. The genus Planctomyces attaches to surfaces through a stalk and holdfast; other genera

lack stalks

4. Most of these eubacteria have life cycles in which sessile cells bud to produce motile

swarmer cells

B. Order Chlamydiales - contains a single genus Chlamydia

      1. Nonmotile, coccoid, gram-negative eubacteria
      2. Obligate intracellular parasites with a unique developmental cycle involving elementary bodies (EBs) that reorganize into reproductively specialized reticulate bodies (RBs) which then change back to infectious elementary bodies
      3. Completely dependent on the host for ATP; most metabolic activity in RBs; very little metabolic activity in EBs
      4. Gram-negative-like wall but lacks muramic acid and peptidoglycan
      5. Use cross-linking of outer membrane and, possibly, periplasmic proteins to achieve osmotic stability
      6. Found mostly in mammals and birds but have been recently isolated from spiders, clams and freshwater invertebrates
      7. Three recognized human pathogens
      1. C. trachomatis - trachoma, nongonococcal urethritis, and other diseases in humans
      2. and mice

      3. C. psittaci - causes psittacosis in humans and infects many other mammals as well;
      4. invades the respiratory and genital tracts; the placenta, developing fetuses, the eye,

        and synovial fluid of the joints

      5. C. pneumoniae - a causative agent of human pneumonia
  1. The Spirochetes
    1. Gram-negative, chemoheterotrophic, flexibly helical bacteria that exhibit a creeping
    2. (crawling) motility due to a structure called an axial filament

    3. The axial filament (a complex of periplasmic flagella) lies in a flexible outer sheath (outer
    4. membrane) outside the protoplasmic cylinder that houses the nucleoid and cytoplasm; the

      function of the sheath is essential (spirochetes will die if it is removed) but unknown

    5. Flagellar rotation is responsible for motility by an unknown mechanism presumably by
    6. rotating the outer sheath or flexing the cell for a crawling motion

    7. Can be anaerobic, facultatively anaerobic, or aerobic and can use a diverse array of organic
    8. molecules as carbon and energy sources

    9. Ecologically diverse
      1. Spirochaeta - anaerobic
      2. Leptospira - aerobic
      3. Treponema - includes the causative agent of syphillis (T. pallidum)
      4. Borrelia - includes the causative agent of Lyme disease (B. burgdorferi)

IV. The Bacteroides

    1. Obligate anaerobes, nonsporing, chemoheterotrophic
    2. Found in oral cavity and intestinal tract of humans and other animals and the rumen of ruminants
    3. Often benefit the host by degrading cellulose, pectins, and other complex carbohydrates, providing extra nutrition for the host
    4. Can constitute as much as 30% of the bacteria isolated from human feces
    5. Some species can be associated with disease

V. The Sphingobacteria

    1. Contains some of the eubacteria with gliding motility; others will be in the g -proteobacteria
    2. Contains the genera Flexibacter, Cytophaga and Sporocytophaga
      1. Differ in morphology, life cycle and physiology
      2. Cytophaga - slender rods with pointed ends
      3. Sporocytophaga - similar to Cytophaga but form spherical resting cells called microcysts
      4. Flexibacter - form long threads; unlike the other two genera, they are unable to degrade complex carbohydrates
    1. Gliding motility used when in contact with a surface; leaves a slime trail; bacteria are
    2. nonmotile when in suspension

    3. Many are capable of utilizing insoluble material as nutrient sources, encountering them during
    4. their gliding activity

    5. Can position themselves for optimal conditions of light intensity, oxygen, hydrogen sulfide,
    6. temperature, and other factors that influence growth

    7. Play a major role in the mineralization of organic matter and can damage exposed wooden
    8. structures

    9. Contribute significantly to wastewater treatment
    10. Some pathogenic species (e.g., C. columnaris causes disease in freshwater and marine fish)

 


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