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Cell structure of a bacterium, one of the two groups of prokaryotic life.

The prokaryotes (singular pronounced /proʊˈkæriət/) are a group of organisms that lack a cell nucleus (= karyon), or any other membrane-bound organelles. They differ from the eukaryotes, which have a cell nucleus. Most are unicellular, but a few prokaryotes such as myxobacteria have multicellular stages in their life cycles.[1] The word prokaryote comes from the Greek πρό- (pro-) "before" + καρυόν (karyon) "nut or kernel", referring to the cell nucleus, + suffix -ώτης (-ōtēs) (pl. -ώτες (-ōtes)). It is also spelled "procaryote".[2]

The prokaryotes are divided into two domains: the bacteria and the archaea. Archaea were recognized as a domain of life in 1990. These organisms were originally thought to live only in inhospitable conditions such as extremes of temperature, pH, and radiation but have since been found in all types of habitats.

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Relationship to eukaryotes

A distinction between prokaryotes and eukaryotes (meaning true kernel, also spelled "eucaryotes") is that eukaryotes do have "true" nuclei containing their DNA, whereas the genetic material in prokaryotes is not membrane-bound. Eukaryotic organisms may be unicellular, as in amoebae, or multicellular, as in plants and humans. The difference between the structure of prokaryotes and eukaryotes is so great that it is sometimes considered to be the most important distinction among groups of organisms. However, a criticism of this classification is that the word "prokaryote" is based on what these organisms are not (they are not eukaryotic), rather than what they are (either archaea or bacteria).[3] In 1977, Carl Woese proposed dividing prokaryotes into the Bacteria and Archaea (originally Eubacteria and Archaebacteria) because of the major differences in the structure and genetics between the two groups of organisms. This arrangement of Eukaryota (also called "Eukarya"), Bacteria, and Archaea is called the three-domain system replacing the traditional two-empire system.[4]

The cell structure of prokaryotes differs greatly from that of eukaryotes. The defining characteristic is the absence of a nucleus. The genomes of prokaryotes are held within an irregular DNA/protein complex in the cytosol called the nucleoid, which lacks a nuclear envelope.[5] Prokaryotes generally lack membrane-bound cell compartments: such as mitochondria and chloroplasts. Instead processes such as oxidative phosphorylation and photosynthesis take place across the prokaryotic plasma membrane.[6] However, prokaryotes do possess some internal structures, such as cytoskeletons,[7][8] and the bacterial order Planctomycetes have a membrane around their nucleoid and contain other membrane-bound cellular structures.[9] Both eukaryotes and prokaryotes contain large RNA/protein structures called ribosomes, which produce protein. Prokaryotes are usually much smaller than eukaryotic cells.[2]

Prokaryotes also differ from eukaryotes in that they contain only a single loop of stable chromosomal DNA stored in an area named the nucleoid, while eukaryote DNA is found on tightly bound and organized chromosomes. Although some eukaryotes have satellite DNA structures called plasmids, these are generally regarded as a prokaryote feature, and many important genes in prokaryotes are stored on plasmids.[2]

Prokaryotes have a larger surface area to volume ratio giving them a higher metabolic rate, a higher growth rate and consequently a shorter generation time compared to Eukaryotes.[2]

Sociality

While prokaryotes are still commonly imagined to be strictly unicellular, most are capable of forming stable aggregate communities. When such communities are encased in a stabilizing polymer matrix (“slime”), they may be called “biofilms”. Cells in biofilms often show distinct patterns of gene expression (phenotypic differentiation) in time and space. Also, like multicellular eukaryotes, these changes in expression appear to often result from cell-to-cell signaling, a phenomenon known as quorum sensing.

Biofilms may be highly heterogeneous and structurally complex and may attach to solid surfaces, or exist at liquid-air interfaces, or potentially even liquid-liquid interfaces. Bacterial biofilms are often made up of microcolonies (approximately dome-shaped masses of bacteria and matrix) separated by “voids” through which the medium (e.g. water) may flow relatively uninhibited. The microcolonies may join together above the substratum to form a continuous layer, closing the network of channels separating microcolonies. This structural complexity—combined with observations that oxygen limitation (a ubiquitous challenge for anything growing in size beyond the scale of diffusion) is at least partially eased by movement of medium throughout the biofilm—has led some to speculate that this may constitute a circulatory system [10].

It is not surprising that many researchers have started calling prokaryotic communities multicellular (for example [11]). Differential cell expression, collective behavior, signaling, programmed cell death, and (in some cases) discrete biological dispersal events all seem to point in this direction. However, these colonies are seldom if ever founded by a single founder (in the way that animals and plants are founded by single cells), which presents a number of theoretical issues. Most explanations of co-operation and the evolution of multicellularity have focused on high relatedness between members of a group (or colony, or whole organism). If a copy of a gene is present in all members of a group, behaviors that promote cooperation between members may permit those members to have (on average) greater fitness than a similar group of selfish individuals[12] (see inclusive fitness and Hamilton's rule). What to make of prokaryotic communities clearly founded by many (most likely unrelated) individuals, yet defined by (apparently) high levels of cooperation, communication, and coordinated behavior?

It is likely that these instances of prokaryotic sociality are the rule rather than the exception, a fact that has serious implications for the way we view prokaryotes in general and the way we deal with them in medicine. Bacterial biofilms may be 100x more resistant to antibiotics than free-living unicells and may be nearly impossible to remove from surfaces once they have colonized them[13]. Other aspects of bacterial cooperation—such as bacterial conjugation and quorum-sensing mediated pathogenicity—present additional challenges to researchers and medical professionals seeking to treat the associated diseases.

Reproduction

Bacteria and archaea reproduce through asexual reproduction, usually by binary fission or budding. Genetic exchange and recombination still occur, but this is a form of horizontal gene transfer and is not a replicative process, simply involving DNA being transferred between two cells, as in bacterial conjugation.

Structure

The sizes of prokaryotes relative to other organisms and biomolecules

Recent research indicates that all prokaryotes actually do have cytoskeletons, albeit more primitive than those of eukaryotes. Besides homologues of actin and tubulin (MreB and FtsZ) the helically arranged building block of the flagellum, flagellin, is one of the most significant cytoskeletal proteins of bacteria as it provides structural backgrounds of chemotaxis, the basic cell physiological response of bacteria. At least some prokaryotes also contain intracellular structures which can be seen as primitive organelles. Membranous organelles (a.k.a. intracellular membranes) are known in some groups of prokaryotes, such as vacuoles or membrane systems devoted to special metabolic properties, e.g. photosynthesis or chemolithotrophy. Additionally, some species also contain protein-enclosed microcompartments, which have distinct physiological roles (e.g. carboxysomes or gas vacuoles).

Most prokaryotes are between 1 µm and 10 µm, but they can vary in size from 0.2 µm to 750 µm (Thiomargarita namibiensis).

Prokaryotic cell Structure
Flagellum
Cell membrane
Cell wall (except genus Mycoplasma)
Cytoplasm
Ribosome
Nucleoid
Glycocalyx
Inclusions

Morphology of prokaryotic cells

Prokaryotic cells have various shapes; the four basic shapes are:[14]

Environment

Prokaryotes live in nearly all environments on earth where there is liquid water. Some archaea and bacteria thrive in harsh conditions, such as high temperatures (thermophiles) or high salinity (halophiles). Organisms such as these are referred to as extremophiles. Many archaea grow as plankton in the oceans. Symbiotic prokaryotes live in or on the bodies of other organisms, including humans.

Evolution of prokaryotes

Phylogenetic tree showing the diversity of prokaryotes, compared to eukaryotes.

The current model of the evolution of the first living organisms is that these were some form of prokaryotes, which may have evolved out of protobionts. The eukaryotes are generally thought to have evolved later in the history of life.[15] However, some authors have questioned this conclusion, arguing that the current set of prokaryotic species may have evolved from more complex eukaryotic ancestors through a process of simplification.[16][17][18] Others have argued that the three domains of life arose simultaneously, from a set of varied cells that formed a single a gene pool.[19] This controversy was summarized in 2005:[20]

There is no consensus among biologists concerning the position of the eukaryotes in the overall scheme of cell evolution. Current opinions on the origin and position of eukaryotes span a broad spectrum including the views that eukaryotes arose first in evolution and that prokaryotes descend from them, that eukaryotes arose contemporaneously with eubacteria and archeabacteria and hence represent a primary line of descent of equal age and rank as the prokaryotes, that eukaryotes arose through a symbiotic event entailing an endosymbiotic origin of the nucleus, that eukaryotes arose without endosymbiosis, and that eukaryotes arose through a symbiotic event entailing a simultaneous endosymbiotic origin of the flagellum and the nucleus, in addition to many other models, which have been reviewed and summarized elsewhere.

The oldest known fossilized prokaryotes were laid down approximately 3.5 billion years ago, only about 1 billion years after the formation of the Earth's crust. Even today, prokaryotes are perhaps the most successful and abundant life forms. Eukaryotes only appear in the fossil record later, and may have formed from endosymbiosis of multiple prokaryote ancestors. The oldest known fossil eukaryotes are about 1.7 billion years old. However, some genetic evidence suggests eukaryotes appeared as early as 3 billion years ago.[21]

While Earth is the only place in the universe where life is known to exist, some have suggested that there is evidence on Mars of fossil or living prokaryotes;[22][23] but this possibility remains the subject of considerable debate and skepticism.[24][25]

Prokaryotes have diversified greatly throughout their long existence. The metabolism of prokaryotes is far more varied than that of eukaryotes, leading to many highly distinct prokaryotic types. For example, in addition to using photosynthesis or organic compounds for energy, as eukaryotes do, prokaryotes may obtain energy from inorganic compounds such as hydrogen sulfide. This enables prokaryotes to thrive in harsh environments as cold as the snow surface of Antarctica, and as hot as undersea hydrothermal vents and land-based hot springs.

See also

References

  1. ^ Kaiser D (October 2003). "Coupling cell movement to multicellular development in myxobacteria". Nat. Rev. Microbiol. 1 (1): 45–54. doi:10.1038/nrmicro733. PMID 15040179.  
  2. ^ a b c d Campbell, N. "Biology:Concepts & Connections". Pearson Education. San Francisco: 2003.
  3. ^ Sapp J (June 2005). "The prokaryote-eukaryote dichotomy: meanings and mythology". Microbiol. Mol. Biol. Rev. 69 (2): 292–305. doi:10.1128/MMBR.69.2.292-305.2005. PMID 15944457. PMC 1197417. http://mmbr.asm.org/cgi/pmidlookup?view=long&pmid=15944457.  
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  6. ^ Harold F (1 June 1972). "Conservation and transformation of energy by bacterial membranes". Bacteriol Rev 36 (2): 172–230. PMID 4261111.  
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  8. ^ Michie KA, Löwe J (2006). "Dynamic filaments of the bacterial cytoskeleton". Annu. Rev. Biochem. 75: 467–92. doi:10.1146/annurev.biochem.75.103004.142452. PMID 16756499. http://www2.mrc-lmb.cam.ac.uk/SS/Lowe_J/group/PDF/annrev2006.pdf.  
  9. ^ Fuerst J (2005). "Intracellular compartmentation in planctomycetes". Annu Rev Microbiol 59: 299–328. doi:10.1146/annurev.micro.59.030804.121258. PMID 15910279.  
  10. ^ Costerton JW, Lewandowski Z, Caldwell DE, Korber DR, Lappin-Scott HM (1995). "Microbial biofilms". Annu. Rev. Microbiol. 49: 711–45. doi:10.1146/annurev.mi.49.100195.003431. PMID 8561477. http://arjournals.annualreviews.org/doi/abs/10.1146/annurev.mi.49.100195.003431.  
  11. ^ Shapiro JA (1998). "Thinking about bacterial populations as multicellular organisms". Annu. Rev. Microbiol. 52: 81–104. doi:10.1146/annurev.micro.52.1.81. PMID 9891794. http://arjournals.annualreviews.org/doi/abs/10.1146/annurev.micro.52.1.81.  
  12. ^ Hamilton WD (July 1964). "The genetical evolution of social behaviour. II". J. Theor. Biol. 7 (1): 17–52. doi:10.1016/0022-5193(64)90039-6. PMID 5875340. http://linkinghub.elsevier.com/retrieve/pii/0022-5193(64)90039-6.  
  13. ^ Costerton JW, Stewart PS, Greenberg EP (May 1999). "Bacterial biofilms: a common cause of persistent infections". Science 284 (5418): 1318–22. doi:10.1126/science.284.5418.1318. PMID 10334980. http://www.sciencemag.org/cgi/pmidlookup?view=long&pmid=10334980.  
  14. ^ Bauman, Robert W.; Tizard, Ian R.; Machunis-Masouka, Elizabeth (2006). Microbiology. San Francisco: Pearson Benjamin Cummings. ISBN 0-8053-7693-3.  
  15. ^ Zimmer C (August 2009). "Origins. On the origin of eukaryotes". Science 325 (5941): 666–8. doi:10.1126/science.325_666. PMID 19661396.  
  16. ^ Brown, J.R. (February 2003). "Ancient Horizontal Gene Transfer". Nature Reviews Genetics 4 (2): 121–132. doi:10.1038/nrg1000. PMID 12560809.  
  17. ^ Forterre P, Philippe H (October 1999). "Where is the root of the universal tree of life?". Bioessays 21 (10): 871–9. doi:10.1002/(SICI)1521-1878(199910)21:10<871::AID-BIES10>3.0.CO;2-Q. PMID 10497338.  
  18. ^ Poole, Anthony, Jeffares, Daniel, Penny, David (September 1999). "Early evolution: prokaryotes, the new kids on the block". Bioessays 21 (10): 880–9. doi:10.1002/(SICI)1521-1878(199910)21:10<880::AID-BIES11>3.0.CO;2-P. PMID 10497339.  
  19. ^ Woese C (June 1998). "The universal ancestor". Proc. Natl. Acad. Sci. U.S.A. 95 (12): 6854–9. doi:10.1073/pnas.95.12.6854. PMID 9618502.  
  20. ^ Martin, William. Woe is the Tree of Life. In Microbial Phylogeny and Evolution: Concepts and Controversies (ed. Jan Sapp). Oxford: Oxford University Press; 2005: 139.
  21. ^ Carl Woese, J Peter Gogarten, "When did eukaryotic cells (cells with nuclei and other internal organelles) first evolve? What do we know about how they evolved from earlier life-forms?" Scientific American, October 21, 1999.
  22. ^ McSween HY (1997). "Evidence for life in a martian meteorite?". GSA Today 7 (7): 1–7. PMID 11541665.  
  23. ^ McKay DS, Gibson EK, Thomas-Keprta KL, et al (August 1996). "Search for past life on Mars: possible relic biogenic activity in martian meteorite ALH84001". Science 273 (5277): 924–30. doi:10.1126/science.273.5277.924. PMID 8688069. http://www.sciencemag.org/cgi/pmidlookup?view=long&pmid=8688069.  
  24. ^ Crenson, Matt (2006-08-06). "After 10 years, few believe life on Mars". Associated Press (on space.com. http://www.space.com/scienceastronomy/ap_060806_mars_rock.html. Retrieved 2006-08-06.  
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External links

PD-icon.svg This article incorporates public domain material from the NCBI document "Science Primer".


Simple English

File:Prokaryote cell
Structure of a prokaryotic bacteria cell

Prokaryotes (or monera) are the simplest living things:[1] bacteria and archaea. They generally do not have a cell nucleus, nor cell organelles, however a small number of exceptions have been found.

The word describes a type of cell. The name comes from Greek pro- (meaning before) and karion, meaning nut or kernel. Prokaryotes are cells that do not have a cell nucleus, and lack other things eukaryotes (cells with a true nucleus) have. Prokaryotes do not have membranes inside the cell. This means that there are no vacuoles, Golgi apparatus, endoplasmic reticula or other organelles inside the cell.

Prokaryotes are unicellular. They are either bacteria or archaea. The Archaea include simple organisms, that were first discovered in extreme environments. Most of them can survive at very high or very low temperatures. Some of them can also survive in highly salty, acidic or alkaline water. Some have been found in geysers, black smokers or oil wells.

Cells with a nucleus are called Eukaryotes. Eukaryote cells include organelles which were once free-living prokaryotes. These are like cells but they do not have any organelles, they actually infect in the cells, a virus infects a cell by attaching to it and injecting viral nucleic acid. Eventually the nucleic acid directs the cell to make more viral nucleic acid and protein coats.

References

  1. Excluding viruses








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