Prokaryotes: Thriving Everywhere!

1. Chapter 27 Prokaryotes

2. Overview: They’re (Almost) Everywhere! • Most prokaryotes are microscopic, but what they lack in size they make up for in numbers • There are more in a handful of fertile soil than the number of people who ever lived • Prokaryotes thrive almost everywhere, including places too acidic, too salty, too cold, or too hot for most other organisms • They have an astonishing genetic diversity

4. Concept 27.1: Structural, functional, and genetic adaptations contribute to prokaryotic success • Most prokaryotes are unicellular, although some species form colonies • Prokaryotic cells have a variety of shapes • The three most common of which are spheres (cocci), rods (bacilli), and spirals Video: Tubeworms

5. LE 27-2 1 µm 2 µm 5 µm Spiral Spherical (cocci) Rod-shaped (bacilli)

6. Cell-Surface Structures • An important feature of nearly all prokaryotic cells is their cell wall, which maintains cell shape, provides physical protection, and prevents the cell from bursting in a hypotonic environment • Using the Gram stain, scientists classify many bacterial species into groups based on cell wall composition, Gram-positive and Gram-negative

7. LE 27-3 Lipopolysaccharide Outer membrane Pepridoglycan layer Cell wall Cell wall Pepridoglycan layer Plasma membrane Plasma membrane Protein Protein Gram- positive bacteria Gram- negative bacteria 20 µm Gram-negative Gram-positive

8. The cell wall of many prokaryotes is covered by a capsule, a sticky layer of polysaccharide or protein

9. LE 27-4 200 nm Capsule

10. Some prokaryotes have fimbriae and pili, which allow them to stick to their substrate or other individuals in a colony

11. LE 27-5 Fimbriae 200 nm

12. Motility • Most motile bacteria propel themselves by flagella that are structurally and functionally different from eukaryotic flagella • In a heterogeneous environment, many bacteria exhibit taxis, the ability to move toward or away from certain stimuli Video: Prokaryotic Flagella (Salmonella typhimurium)

13. LE 27-6 Flagellum Filament 50 nm Cell wall Hook Basal apparatus Plasma membrane

14. Internal and Genomic Organization • Prokaryotic cells usually lack complex compartmentalization • Some prokaryotes do have specialized membranes that perform metabolic functions

15. LE 27-7 1 µm 0.2 µm Respiratory membrane Thylakoid membranes Photosynthetic prokaryote Aerobic prokaryote

16. The typical prokaryotic genome is a ring of DNA that is not surrounded by a membrane and that is located in a nucleoid region

17. LE 27-8 Chromosome 1 µm

18. Some species of bacteria also have smaller rings of DNA called plasmids

19. Reproduction and Adaptation • Prokaryotes reproduce quickly by binary fission and can divide every 1–3 hours • Many prokaryotes form endospores, which can remain viable in harsh conditions for centuries

20. LE 27-9 Endospore 0.3 µm

21. Rapid reproduction and horizontal gene transfer facilitate the evolution of prokaryotes in changing environments

22. Concept 27.2: A great diversity of nutritional and metabolic adaptations have evolved in prokaryotes • All four models of nutrition are found among prokaryotes: • Photoautotrophy • Chemoautotrophy • Photoheterotrophy • Chemoheterotrophy

24. Metabolic Relationships to Oxygen • Prokaryotic metabolism varies with respect to oxygen: • Obligate aerobes require oxygen • Facultative anaerobes can survive with or without oxygen • Obligate anaerobes are poisoned by oxygen

25. Nitrogen Metabolism • Prokaryotes can metabolize nitrogen in a variety of ways • In nitrogen fixation, some prokaryotes convert atmospheric nitrogen to ammonia

26. Metabolic Cooperation • Cooperation between prokaryotes allows them to use environmental resources they could not use as individual cells • In the cyanobacterium Anabaena, photosynthetic cells and nitrogen-fixing cells exchange metabolic products Video: Cyanobacteria (Oscillatoria)

27. LE 27-10 Photosynthetic cells Heterocyte 20 µm

28. In some prokaryotic species, metabolic cooperation occurs in surface-coating colonies called biofilms

30. Concept 27.3: Molecular systematics is illuminating prokaryotic phylogeny • Until the late 20th century, systematists based prokaryotic taxonomy on phenotypic criteria • Applying molecular systematics to the investigation of prokaryotic phylogeny has produced dramatic results

31. Lessons from Molecular Systematics • Molecular systematics is leading to a phylogenetic classification of prokaryotes • It allows systematists to identify major new clades

32. A tentative phylogeny of some of the major taxa of prokaryotes based on molecular systematics

33. LE 27-12 Domain Eukarya Domain Archaea Domain Bacteria Proteobacteria Delta Epsilon Alpha Gram-positive bacteria Beta Chlamydias Korarchaeotes Euryarchaeotes Crenarcaeotes Nanoarchaeotes Gamma Cyanobacteria Eukaryotes Spirochetes Universal ancestor

34. Bacteria • Diverse nutritional types are scattered among the major groups of bacteria • The two largest groups are the proteobacteria and the Gram-positive bacteria

35. LE 27-13 PROTEOBACTERIA CHLAMYDIAS Subgroup: Alpha Proteobacteria 2.5 µm 2.5 µm Chlamydia (arrows) Rhizobium (arrows) SPIROCHETES Subgroup: Beta Proteobacteria 5 µm 1 µm Leptospira Nitrosomonas GRAM-POSITIVE BACTERIA Subgroup: Gamma Proteobacteria 1 µm 5 µm 0.5 µm Mycoplasmas covering a human fibroblast cell Streptomyces Chromatium CYANOBACTERIA Subgroup: Delta Proteobacteria 50 µm 5 µm 10 µm Oscillatoria Bdellovibrio bacteriophorus Chrondromyces crocatus Subgroup: Epsilon Proteobacteria 2 µm Heliocobacter pylori

36. Archaea • Archaea share certain traits with bacteria and other traits with eukaryotes

38. Some archaea live in extreme environments • Extreme thermophiles thrive in very hot environments • Extreme halophiles live in high saline environments

40. Methanogens live in swamps and marshes and produce methane as a waste product

41. Concept 27.4: Prokaryotes play crucial roles in the biosphere • Prokaryotes are so important to the biosphere that if they were to disappear, the prospects for any other life surviving would be dim

42. Chemical Recycling • Prokaryotes play a major role in the continual recycling of chemical elements between the living and nonliving components of ecosystems • Chemoheterotrophic prokaryotes function as decomposers, breaking down corpses, dead vegetation, and waste products • Nitrogen-fixing prokaryotes add usable nitrogen to the environment

43. Symbiotic Relationships • Many prokaryotes live with other organisms in symbiotic relationships • In mutualism, both symbiotic organisms benefit • In commensalism, one organism benefits while neither harming nor helping the other in any significant way • In parasitism, one organism, called a parasite, benefits at the expense of the host

45. Concept 27.5: Prokaryotes have both harmful and beneficial impacts on humans • Some prokaryotes are human pathogens, but others have positive interactions with humans

46. Pathogenic Prokaryotes • Prokaryotes cause about half of all human diseases • Lyme disease is an example

48. Pathogenic prokaryotes typically cause disease by releasing exotoxins or endotoxins • Exotoxins cause disease even if the prokaryotes that produce them are not present • Endotoxins are released only when bacteria die and their cell walls break down • Many pathogenic bacteria are potential weapons of bioterrorism

49. Prokaryotes in Research and Technology • Experiments using prokaryotes have led to important advances in DNA technology • Prokaryotes are the principal agents in bioremediation, the use of organisms to remove pollutants from the environment