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Prokaryote
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Diagram of a typical prokaryotic bacteria cell
A prokaryote is a unicellular organism that lacks a membrane-bound nucleus, mitochondria, or any other membrane-bound organelle.[1] The word prokaryote comes from the Greek πρό (pro) "before" and κάρυον (karyon) "nut or kernel".[2][3] Prokaryotes are divided into two domains, Archaea and Bacteria. In contrast, species with nuclei and organelles are placed in the third domain, Eukaryota.[4]. Prokaryotes reproduce without fusion of gametes. The first living organisms are thought to have been prokaryotes.
In the prokaryotes, all the intracellular water-soluble components (proteins, DNA and metabolites) are located together in the cytoplasm enclosed by the cell membrane, rather than in separate cellular compartments. Bacteria, however, do possess protein-based bacterial microcompartments, which are thought to act as primitive organelles enclosed in protein shells.[5][6] Some prokaryotes, such as cyanobacteria may form large colonies. Others, such as myxobacteria, have multicellular stages in their life cycles.[7]
Molecular studies have provided insight into the evolution and interrelationships of the three domains of biological species.[8] Eukaryotes are organisms, including humans, whose cells have a well defined membrane-bound nucleus (containing chromosomal DNA) and organelles. The division between prokaryotes and eukaryotes reflects the existence of two very different levels of cellular organization. Distinctive types of prokaryotes include extremophiles and methanogens; these are common in some extreme environments.[1]
Contents
[hide]
1 Structure
2 Morphology
3 Reproduction
4 DNA transfer
5 Sociality
6 Environment
7 Classification
8 Evolution
9 Relationship to eukaryotes
10 See also
11 References
12 External links
Structure[edit]
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Also see: Human timeline and Nature timeline
Prokaryotes have a prokaryotic cytoskeleton, albeit more primitive than that of the 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 that can be seen as primitive organelles. Membranous organelles (or intracellular membranes) are known in some groups of prokaryotes, such as vacuoles or membrane systems devoted to special metabolic properties, such as photosynthesis or chemolithotrophy. In addition, some species also contain carbohydrate-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 (Mycoplasma genitalium) to 750 µm (Thiomargarita namibiensis).
Prokaryotic cell structure
Flagellum (only in some types of prokaryotes)[which?]
Long, whip-like protrusion that aids cellular locomotion used by both gram positive and gram negative organisms.
Cell membrane
Surrounds the cell's cytoplasm and regulates the flow of substances in and out of the cell.
Cell wall (except genera Mycoplasma and Thermoplasma)
Outer covering of most cells that protects the bacterial cell and gives it shape.
Cytoplasm
A gel-like substance composed mainly of water that also contains enzymes, salts, cell components, and various organic molecules.
Ribosome
Cell structures responsible for protein production.
Nucleoid
Area of the cytoplasm that contains the prokaryote's single DNA molecule.
Glycocalyx (only in some types of prokaryotes)
A glycoprotein-polysaccharide covering that surrounds the cell membranes.
Inclusions
It contains the inclusion bodies like ribosomes and larger masses scattered in the cytoplasmic matrix.
Morphology[edit]
Prokaryotic cells have various shapes; the four basic shapes of bacteria are:[9]
Cocci – spherical
Bacilli – rod-shaped
Spirochaete – spiral-shaped
Vibrio – comma-shaped
The archaeon Haloquadratum has flat square-shaped cells.[10]
Reproduction[edit]
Bacteria and archaea reproduce through asexual reproduction, usually by binary fission. Genetic exchange and recombination still occur, but this is a form of horizontal gene transfer and is not a replicative process, simply involving the transference of DNA between two cells, as in bacterial conjugation.
DNA transfer[edit]
DNA transfer between prokaryotic cells occurs in bacteria and archaea, although it has been mainly studied in bacteria. In bacteria, gene transfer occurs by three processes. These are (1) bacterial virus (bacteriophage)-mediated transduction, (2) plasmid-mediated conjugation, and (3) natural transformation. Transduction of bacterial genes by bacteriophage appears to reflect an occasional error during intracellular assembly of virus particles, rather than an adaptation of the host bacteria. The transfer of bacterial DNA is under the control of the bacteriophage’s genes rather than bacterial genes. Conjugation in the well-studied E. coli system is controlled by plasmid genes, and is an adaptation for distributing copies of a plasmid from one bacterial host to another. Infrequently during this process, a plasmid may integrate into the host bacterial chromosome, and subsequently transfer part of the host bacterial DNA to another bacterium. Plasmid mediated transfer of host bacterial DNA (conjugation) also appears to be an accidental process rather than a bacterial adaptation.
Play media
3D animation of a prokaryotic cell that shows all the elements that compose it
Natural bacterial transformation involves the transfer of DNA from one bacterium to another through the intervening medium. Unlike transduction and conjugation, transformation is clearly a bacterial adaptation for DNA transfer, because it depends on numerous bacterial gene products that specifically interact to perform this complex process.[11] For a bacterium to bind, take up and recombine donor DNA into its own chromosome, it must first enter a special physiological state called competence. About 40 genes are required in Bacillus subtilis for the development of competence.[12] The length of DNA transferred during B. subtilis transformation can be as much as a third to the whole chromosome.[13][14] Transformation is a common mode of DNA transfer, and 67 prokaryotic species are thus far known to be naturally competent for transformation.[15]
Among archaea, Halobacterium volcanii forms cytoplasmic bridges between cells that appear to be used for transfer of DNA from one cell to another.[16] Another archaeon, Sulfolobus solfataricus, transfers DNA between cells by direct contact. Frols et al.[17] found that exposure of S. solfataricus to DNA damaging agents induces cellular aggregation, and suggested that cellular aggregation may enhance DNA transfer among cells to provide increased repair of damaged DNA via homologous recombination.
Sociality[edit]
While prokaryotes are considered strictly unicellular, most can form stable aggregate communities.[18] When such communities are encased in a stabilizing polymer matrix ("slime"), they may be called "biofilms".[19] Cells in biofilms often show distinct patterns of gene expression (phenotypic differentiation) in time and space. Also, as with multicellular eukaryotes, these changes in expression often appear to 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 easily. 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 [20] and many researchers have started calling prokaryotic communities multicellular (for example [21]). Differential cell expression, collective behavior, signaling, programmed cell death, and (in some cases) discrete biological dispersal[22] 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[23] (see inclusive fitness and Hamilton's rule).
Should these instances of prokaryotic sociality prove to be the rule rather than the exception, it would have serious implications for the way we view prokaryotes in general, and the way we deal with them in medicine.[24] Bacterial biofilms may be 100 times more resistant to antibiotics than free-living unicells and may be nearly impossible to remove from surfaces once they have colonized them.[25] 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.
Environment[edit]
Phylogenetic ring showing the diversity of prokaryotes, and symbiogenetic origins of eukaryotes
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, studied in cryobiology or as hot as undersea hydrothermal vents and land-based hot springs.
Prokaryotes live in nearly all environments on Earth. Some archaea and bacteria are extremophiles, thriving in harsh conditions, such as high temperatures (thermophiles) or high salinity (halophiles).[26] Many archaea grow as plankton in the oceans. Symbiotic prokaryotes live in or on the bodies of other organisms, including humans.
Classification[edit]
Phylogenetic and symbiogenetic tree of living organisms, showing the origins of eukaryotes and prokaryotes
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. Archaea were originally thought to be extremophiles, living only in inhospitable conditions such as extremes of temperature, pH, and radiation but have since been found in all types of habitats. The resulting arrangement of Eukaryota (also called "Eucarya"), Bacteria, and Archaea is called the three-domain system, replacing the traditional two-empire system.[27][28]
Evolution[edit]
Main article: Molecular evolution
Diagram of the origin of life with the Eukaryotes appearing early, not derived from Prokaryotes, as proposed by Richard Egel in 2012. This view, one of many on the relative positions of Prokaryotes and Eukaryotes, implies that the universal common ancestor was relatively large and complex.[29]
A widespread current model of the evolution of the first living organisms is that these were some form of prokaryotes, which may have evolved out of protocells, while the eukaryotes evolved later in the history of life.[30] 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.[31][32][33] Others have argued that the three domains of life arose simultaneously, from a set of varied cells that formed a single gene pool.[34] This controversy was summarized in 2005:[35]
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. 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.[36]
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.[37][38] However, this possibility remains the subject of considerable debate and skepticism.[39][40]
Relationship to eukaryotes[edit]
Comparison of eukaryotes vs. prokaryotes
The division between prokaryotes and eukaryotes is usually considered the most important distinction or difference among organisms. The distinction is that eukaryotic cells have a "true" nucleus containing their DNA, whereas prokaryotic cells do not have a nucleus. Both eukaryotes and prokaryotes contain large RNA/protein structures called ribosomes, which produce protein.
Another difference is that ribosomes in prokaryotes are smaller than in eukaryotes. However, two organelles found in many eukaryotic cells, mitochondria and chloroplasts, contain ribosomes similar in size and makeup to those found in prokaryotes.[41] This is one of many pieces of evidence that mitochondria and chloroplasts are themselves descended from free-living bacteria. This theory holds that early eukaryotic cells took in primitive prokaryotic cells by phagocytosis and adapted themselves to incorporate their structures, leading to the mitochondria we see today.
The genome in a prokaryote is held within a DNA/protein complex in the cytosol called the nucleoid, which lacks a nuclear envelope.[42] The complex contains a single, cyclic, double-stranded molecule of stable chromosomal DNA, in contrast to the multiple linear, compact, highly organized chromosomes found in eukaryotic cells. In addition, many important genes of prokaryotes are stored in separate circular DNA structures called plasmids.[2]
Prokaryotes lack mitochondria and chloroplasts. Instead, processes such as oxidative phosphorylation and photosynthesis take place across the prokaryotic cell membrane.[43] However, prokaryotes do possess some internal structures, such as prokaryotic cytoskeletons.[44][45] It has been suggested that the bacterial order Planctomycetes have a membrane around their nucleoid and contain other membrane-bound cellular structures.[46] However, further investigation revealed that Planctomycetes cells are not compartmentalized or nucleated and like the other bacterial membrane systems are all interconnected.[47]
Prokaryotic cells are usually much smaller than eukaryotic cells.[2] Therefore, prokaryotes have a larger surface-area-to-volume ratio, giving them a higher metabolic rate, a higher growth rate, and as a consequence, a shorter generation time than eukaryotes.[2]
See also[edit]
Molecular and cell biology portal
Biology portal
Bacterial cell structure
Evolution of sexual reproduction
List of sequenced archaeal genomes
List of sequenced bacterial genomes
Monera, an obsolete kingdom including Archaea and Bacteria
Nanobacterium
Nanobe
Parakaryon
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External links[edit]
Wikimedia Commons has media related to Procaryota.
Prokaryote versus eukaryote, BioMineWiki
The Taxonomic Outline of Bacteria and Archaea
The Prokaryote-Eukaryote Dichotomy: Meanings and Mythology
Quiz on prokaryote anatomy
TOLWEB page on Eukaryote-Prokaryote phylogeny
This article incorporates public domain material from the NCBI document "Science Primer".
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Prokaryotes: Bacteria classification (phyla and orders)
Domain
Archaea
Bacteria
Eukaryota
(Supergroup
Plant
Hacrobia
Heterokont
Alveolata
Rhizaria
Excavata
Amoebozoa
Opisthokonta
Animal
Fungi)
G-/
OM
Terra-/
Glidobacteria
(BV1)
Eobacteria
Deinococcus–Thermus
Deinococcales
Thermales
Chloroflexi
Anaerolineales
Caldilineales
Chloroflexales
Herpetosiphonales
Dehalococcoidales
Ktedonobacterales
Thermogemmatisporales
Thermomicrobiales
Sphaerobacterales
other glidobacteria
Thermodesulfobacteria
thermophiles
Aquificae
Thermotogae
Cyanobacteria
Proteobacteria
(BV2)
Alpha
Caulobacterales
Kiloniellales
Kordiimonadales
Magnetococcales
Parvularculales
Rhizobiales
Rhodobacterales
Rhodospirillales
Rickettsiales
Sneathiellales
Sphingomonadales
Beta
Burkholderiales
Hydrogenophilales
Methylophilales
Neisseriales
Nitrosomonadales
Procabacteriales
Rhodocyclales
Gamma
Acidithiobacillales
Aeromonadales
Alteromonadales
Cardiobacteriales
Chromatiales
Enterobacteriales
Legionellales
Methylococcales
Oceanospirillales
Orbales
Pasteurellales
Pseudomonadales
Salinisphaerales
Thiotrichales
Vibrionales
Xanthomonadales
Delta
Bdellovibrionales
Desulfarculales
Desulfobacterales
Desulfovibrionales
Desulfurellales
Desulfuromonadales
Myxococcales
Syntrophobacterales
Syntrophorhabdales
Epsilon
Campylobacterales
Nautiliales
Zeta
Mariprofundales
BV4
Spirochaetes
Spirochaetes
Sphingobacteria
(FCB group)
Fibrobacteres
Chlorobi
Chlorobiales
Ignavibacteriales
Bacteroidetes
Bacteroidales
Cytophagales
Flavobacteriales
Sphingobacteriales
Planctobacteria/
(PVC group)
Chlamydiae
Lentisphaerae
Lentisphaerales
Oligosphaerales
Victivallales
Planctomycetes
Phycisphaerales
Planctomycetales
Verrucomicrobia
Puniceicoccales
Opitutales
Chthoniobacterales
Verrucomicrobiales
"Poribacteria"
Other GN
Acidobacteria
Acidobacteriales
Acanthopleuribacterales
Holophagales
Solibacterales
Armatimonadetes
Armatimonadales
Chthonomonadales
Fimbriimonadales
Caldiserica
Chrysiogenetes
Deferribacteres
Dictyoglomi
Elusimicrobia
Fusobacteria
Gemmatimonadetes
Nitrospirae
Synergistetes
G+/
no OM
Firmicutes
(BV3)
Bacilli
Bacillales
Lactobacillales
Clostridia
Clostridiales
Halanaerobiales
Thermoanaerobacteriales
Natranaerobiales
Erysipelotrichia
Erysipelotrichiales
Thermolithobacteria
Thermolithobacterales
Tenericutes/
Mollicutes
Mycoplasmatales
Entomoplasmatales
Anaeroplasmatales
Acholeplasmatales
Haloplasmatales
Negativicutes
Selenomonadales
Actinobacteria
(BV5)
Actinobacteria
Actinomycetales
Bifidobacteriales
Acidimicrobiia
Acidimicrobiales
Coriobacteriidae
Coriobacteriales
Nitriliruptoria
Euzebyales
Nitriliruptorales
Rubrobacteria
Gaiellales
Rubrobacterales
Thermoleophilales
Solirubrobacterales
Incertae
sedis
†Archaeosphaeroides
†Eobacterium
†Leptotrichites
Source: Bergey's Manual (2001–2012). Alternative views: Wikispecies.
[show]
v
t
e
Prokaryotes: Archaea classification
Domain
Archaea
Bacteria
Eukaryota
(Supergroup
Plant
Hacrobia
Heterokont
Alveolata
Rhizaria
Excavata
Amoebozoa
Opisthokonta
Animal
Fungi)
Euryarchaeota
Euryarchaeota
Archaeoglobi
Archaeoglobales
Halobacteria
Halobacteriales
Methanobacteria
Methanobacteriales
Methanococci
Methanococcales
Methanomicrobia
Methanocellales
Methanomicrobiales
Methanosarcinales
Methanopyri
Methanopyrales
Thermococci
Thermococcales
Thermoplasmata
Thermoplasmatales
DPANN
Diapherotrites
"Ca. Iainarchaeum"
Parvarchaeota
"Ca. Micrarchaeum"・ "Ca. Parvarchaeum"
Aenigmarchaeota
"Ca. Aenigmarchaeum"
Nanohaloarchaea
"Ca. Nanosalina"・ "Ca. Nanosalinarum"
Nanoarchaeota
Nanoarchaeum
TACK+ASGARD
TACK
Crenarchaeota
Thermoprotei
Acidilobales
Desulfurococcales
Fervidicoccales
Sulfolobales
Thermoproteales
Thaumarchaeota
Nitrososphaeria
Cenarchaeales
"Nitrosopumilales"
Nitrososphaerales
Korarchaeota
"Ca. Korarchaeum cryptofilum"
Aigarchaeota
"Ca. Caldiarchaeum subterraneum"
Geoarchaeota
Geoarchaeota
Bathyarchaeota
Bathyarchaeota
ASGARD
"Ca. Lokiarchaeum"
Odinarchaeota
Thorarchaeota
Heimdallarchaeota
Authority control
GND: 4201051-2
Retrieved from "https://en.wikipedia.org/w/index.php?title=Prokaryote&oldid=827075611"
Categories:
Prokaryotes
Bacteriology
Biochemistry
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Temple (anatomy)
From Wikipedia, the free encyclopedia
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This article needs additional citations for verification. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed. (July 2016) (Learn how and when to remove this template message)
Temple (anatomy)
Location of temple
Human skull. Temporal bone is orange, and the temple overlies the temporal bone as well as overlying the sphenoid bone.
Details
Artery
superficial temporal artery
Vein
superficial temporal vein
Identifiers
Latin
Tempus
TA
A01.1.00.004
FMA
46450
Anatomical terminology
[edit on Wikidata]
Temple indicates the side of the head behind the eyes. The bone beneath is the temporal bone as well as part of the sphenoid bone.
Cladists classify land vertebrates based on the presence of an upper hole, a lower hole, both, or neither in the cover of dermal bone that formerly covered the temporalis muscle, whose origin is the temple and whose insertion is the jaw. The brain has a lobe called the temporal lobe.
Contents
[hide]
1 Etymology
2 References
3 See also
4 External links
Etymology[edit]
The word "templar" as used in anatomy has a separate etymology from the word temple, meaning "place of worship". Both come from Latin, but the word for the place of worship comes from templum, whereas the word for the part of the head comes from Vulgar Latin *tempula, modified from tempora, plural form ("both temples") of tempus, a word that meant both "time" and the part of the head. Due to the common source with the word for time, the adjective for both is "temporal" (both "pertaining to time" and "pertaining to the anatomical temple").
Regarding the temporalis muscle: in English, this muscle’s name is the time muscle. The word temporalis comes from the Latin word tempus meaning “time.” The muscle covers the temporal bone, or time bone, which received its name because the hair of the head covering this bone is often the first hair to turn gray during the aging process. Therefore, this region is the first to show the effects of aging.[1]
References[edit]
Jump up ^ http://www.anatomyexpert.com/app/structure/5007/4/
See also[edit]
pterion, the weakest part of the skull
External links[edit]
Media related to Temple (anatomy) at Wikimedia Commons
[hide]
v
t
e
Human regional anatomy
Head
Ear
Face
Cheek
Chin
Eye
Mouth
Nose
Forehead
Jaw
Occiput
Scalp
Temple
Neck
Adam's apple
Throat
Trunk
Abdomen
Waist
Midriff
Navel
Back
Thorax
Breast
Pelvis
Sex organs
Limbs
Arm
Shoulder
Axilla
Brachium
Elbow
Forearm
Wrist
Hand
Finger
Thumb
Index
Middle
Ring
Little
Leg
Buttocks
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Thigh
Knee
Calf
Foot
Ankle
Heel
Sole
Toe
Other
Écorché
General anatomy: systems and organs, regional anatomy, planes and lines, superficial axial anatomy, superficial anatomy of limbs
Retrieved from "https://en.wikipedia.org/w/index.php?title=Temple_(anatomy)&oldid=825195114"
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Human head and neck
Vertebrate anatomy
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UNESCO member states
UNESCO member state dependent territory with separate NOC
UNESCO associates
UNESCO observers
UNESCO member states with intention to leave
As of November 2015[update], UNESCO members include 195 member states and 10 associate members.[1][2] Some members have additional National Organizing Committees (NOCs) for some of their dependent territories.[3] The associate members are non-independent states.
Three UNESCO member states are not UN member states: Cook Islands, Niue, and Palestine (Palestine is a non-member observer State of the United Nations General Assembly since 29 November 2012), while one UN member state (Liechtenstein) is not a UNESCO member. Kosovo was approved for membership by UNESCO's executive board in 2015,[4] but the proposal did not receive the required 2/3 of votes in favour at the general conference.[5]
Contents
[hide]
1 Member States
2 Associate members
3 Observers
4 See also
5 Notes
6 References
Member States[edit]
The current 195 UNESCO member states with the date on which they became members:[6][7][1][3]
Afghanistan (4 May 1948)
Albania (16 October 1958)
Algeria (15 October 1962)
Andorra (20 October 1993)
Angola (11 March 1977)
Antigua and Barbuda (15 July 1982)
Argentina (15 September 1948)
Armenia (9 June 1992)
Australia (4 November 1946)
Austria (13 August 1948)
Azerbaijan (3 June 1992)
Bahamas (23 April 1981)
Bahrain (18 January 1972)
Bangladesh (27 October 1972)
Barbados (24 October 1968)
Belarus (12 May 1954)
Belgium (29 November 1946)
Belize (10 May 1982)
Benin (18 October 1960)
Bhutan (13 April 1982)
Bolivia (13 November 1946)
Bosnia and Herzegovina (2 June 1993)
Botswana (16 January 1980)
Brazil (4 November 1946)
Brunei Darussalam (17 March 2005)
Bulgaria (17 May 1956)
Burkina Faso (14 November 1960)
Burundi (16 November 1962)
Cambodia (3 July 1951)
Cameroon (11 November 1960)
Canada (4 November 1946)
Cape Verde (15 February 1978)
Central African Republic (11 November 1960)
Chad (19 December 1960)
Chile (7 July 1953)
China[a][b] (4 November 1946)
including separate NOC for Hong Kong
Colombia (31 October 1947)
Comoros (22 March 1977)
Congo (24 October 1960)
Cook Islands (25 October 1989)
Costa Rica (19 May 1950)
Côte d'Ivoire (27 October 1960)
Croatia (1 June 1992)
Cuba (29 August 1947)
Cyprus (6 February 1961)
Czech Republic (22 February 1993)
Democratic Republic of the Congo (25 November 1960)
Denmark[a] (4 November 1946)
Djibouti (31 August 1989)
Dominica (9 January 1979)
Dominican Republic (4 November 1946)
Ecuador (22 January 1947)
Egypt (4 November 1946)
El Salvador (28 April 1948)
Equatorial Guinea (29 November 1979)
Eritrea (2 September 1993)
Estonia (14 October 1991)
Ethiopia (1 July 1955)
Fiji (14 July 1983)
Finland (10 October 1956)
France (4 November 1946)
Gabon (16 November 1960)
Gambia (1 August 1973)
Georgia (7 October 1992)
Germany (11 July 1951)
Ghana (11 April 1958)
Greece (4 November 1946)
Grenada (17 February 1975)
Guatemala (2 January 1950)
Guinea (2 February 1960)
Guinea-Bissau (1 November 1974)
Guyana (21 March 1967)
Haiti (18 November 1946)
Honduras (16 December 1947)
Hungary (14 September 1948)
Iceland (8 June 1964)
India (4 November 1946)
Indonesia (27 May 1950)
Iran (6 September 1948)
Iraq (21 October 1948)
Ireland (3 October 1961)
Israel (16 September 1949)
Italy (27 January 1948)
Jamaica (7 November 1962)
Japan (2 July 1951)
Jordan (14 June 1950)
Kazakhstan (22 May 1992)
Kenya (7 April 1964)
Kiribati (24 October 1989)
North Korea (18 October 1974)
South Korea (14 June 1950)
Kuwait (18 November 1960)
Kyrgyzstan (2 June 1992)
Lao People's Democratic Republic (9 July 1951)
Latvia (14 October 1991)
Lebanon (4 November 1946)
Lesotho (29 September 1967)
Liberia (6 March 1947)
Libya (27 June 1953)
Lithuania (7 October 1991)
Luxembourg (27 October 1947)
Republic of Macedonia (28 June 1993)
Madagascar (10 November 1960)
Malawi (27 October 1964)
Malaysia (16 June 1958)
Maldives (18 July 1980)
Mali (7 November 1960)
Malta (10 February 1965)
Marshall Islands (30 June 1995)
Mauritania (10 January 1962)
Mauritius (25 October 1968)
Mexico (4 November 1946)
Federated States of Micronesia (19 October 1999)
Moldova (27 May 1992)
Monaco (6 July 1949)
Mongolia (1 November 1962)
Montenegro (1 March 2007)[c]
Morocco (7 November 1956)
Mozambique (11 October 1976)
Myanmar (27 June 1949)
Namibia (2 November 1978)
Nauru (17 October 1996)
Nepal (1 May 1953)
Netherlands[a] (1 January 1947)
New Zealand[a] (4 November 1946)
Nicaragua (22 February 1952)
Niger (10 November 1960)
Nigeria (14 November 1960)
Niue (26 October 1993)[d]
Norway (4 November 1946)
Oman (10 February 1972)
Pakistan (14 September 1949)
Palau (20 September 1999)
Palestine (23 November 2011)[e]
Panama (10 January 1950)
Papua New Guinea (4 October 1976)
Paraguay (20 June 1955)
Peru (21 November 1946)
Philippines (21 November 1946)
Poland (6 November 1946)
Portugal (11 September 1974)[f]
Qatar (27 January 1972)
Romania (27 July 1956)
Russian Federation (21 April 1954)[g]
Rwanda (7 November 1962)
Saint Kitts and Nevis (26 October 1983)
Saint Lucia (6 March 1980)
Saint Vincent and the Grenadines (14 January 1983)
Samoa (3 April 1981)
San Marino (12 November 1974)
São Tomé and Príncipe (22 January 1980)
Saudi Arabia (4 November 1946)
Senegal (10 November 1960)
Serbia (20 December 2000)[c]
Seychelles (18 October 1976)
Sierra Leone (28 March 1962)
Singapore (8 October 2007)[h]
Slovakia (9 February 1993)
Slovenia (27 May 1992)
Solomon Islands (7 September 1993)
Somalia (15 November 1960)
South Africa (12 December 1994)[i]
South Sudan (27 October 2011)[9]
Spain (30 January 1953)
Sri Lanka (14 November 1949)
Sudan (26 November 1956)
Suriname (16 July 1976)
Swaziland (25 January 1978)
Sweden (23 January 1950)
Switzerland (28 January 1949)
Syrian Arab Republic (16 November 1946)
Tajikistan (6 April 1993)
United Republic of Tanzania (6 March 1962)
Thailand (1 January 1949)
Timor-Leste (5 June 2003)
Togo (17 November 1960)
Tonga (29 September 1980)
Trinidad and Tobago (2 November 1962)
Tunisia (8 November 1956)
Turkey (4 November 1946)
Turkmenistan (17 August 1993)
Tuvalu (21 October 1991)
Uganda (9 November 1962)
Ukraine (12 May 1954)
United Arab Emirates (20 April 1972)
United Kingdom of Great Britain and Northern Ireland[a] (1 July 1997)[j]
including separate NOC for Bermuda
United States of America (1 October 2003)[k][l]
including separate NOCs for: Guam, American Samoa, Puerto Rico and U.S. Virgin Islands
Uruguay (8 November 1947)
Uzbekistan (26 October 1993)
Vanuatu (10 February 1994)
Venezuela (25 November 1946)
Vietnam (6 July 1951)
Yemen (2 April 1962)
Zambia (9 November 1964)
Zimbabwe (22 September 1980)
Currently Liechtenstein is not a member of UNESCO, but they have a NOC.[3]
The United States announced its intention to withdraw from UNESCO on 12 October 2017, accusing it of "anti-Israel bias”.[11] Hours later on the same day, Israel announced it would also be leaving UNESCO.[11]
Associate members[edit]
The 10 associate members of UNESCO and the date on which they became associate members:[1]
Anguilla (5 November 2013)
Aruba (20 October 1987)
British Virgin Islands (24 November 1983)
Cayman Islands (30 October 1999)
Curaçao (25 October 2011)[m]
Faroes (12 October 2009)
Macao (25 October 1995)[n]
Montserrat (3 November 2015)
Sint Maarten (25 October 2011)[m]
Tokelau (15 October 2001)
Observers[edit]
There are 2 Permanent Observers and 10 intergovernmental organizations with Permanent Observer Missions to UNESCO.[12]
Non-member States[13]
Holy See
Entities[13]
Sovereign Military Order of Malta
Intergovernmental organizations[13]
African Union
Arab League Educational, Cultural and Scientific Organization
Council of Europe
European Union
Inter-American Development Bank
Islamic Educational, Scientific and Cultural Organization
Latin American Faculty of Social Sciences
Latin Union
League of Arab States
Organization of Ibero-American States for Education, Science and Culture
In addition, there is a liaison office of the United Nations University at UNESCO.[14]
See also[edit]
United Nations portal
Member states of the United Nations
Notes[edit]
^ Jump up to: a b c d e Some of the dependent territories of China, Denmark, Netherlands, New Zealand, United Kingdom are associate members of UNESCO.
Jump up ^ Membership was founded as the Republic of China to 1971.
^ Jump up to: a b The former Socialist Federal Republic of Yugoslavia (SFRY) became a Member State on 31 March 1950. The participation of Yugoslavia in meetings of governing bodies and conferences of UNESCO was suspended following Resolution 47/1 adopted by the General Assembly of the United Nations on 22 September 1992, which stated that the Federal Republic of Yugoslavia (Serbia and Montenegro) could not continue automatically the membership of the former SFRY. Accordingly, the Federal Republic of Yugoslavia, which became a Member State on 20 December 2000, could not automatically succeed the former SFRY as a member of the Organization. Following the adoption of the Constitutional Charter of Serbia and Montenegro by the Assembly of the Federal Republic of Yugoslavia on 4 February 2003, the name of the State of the Federal Republic of Yugoslavia has been changed to "Serbia and Montenegro". Further to the Declaration of Independence adopted by the Parliament of Montenegro on 3 June 2006, UNESCO has been informed that the membership of the State Union Serbia and Montenegro in UNESCO is continued by the Republic of Serbia on the basis of Article 60 of the Constitutional Charter of Serbia and Montenegro.
Jump up ^ Niue does not have a National Organizing Committee established.[3]
Jump up ^ On 31 October 2011, the UNESCO General Conference in Paris admitted Palestine as a UNESCO member state, with 107 votes in favour of admission and 14 votes against, with 52 abstentions (a two-thirds majority vote in favour by member states is required). The decision took effect on 23 November 2011 when Palestine ratified UNESCO’s constitution.[2]
Jump up ^ Previously a Member State from 11 March 1965 to 31 December 1972.
Jump up ^ Membership was founded as the Soviet Union to 1991.
Jump up ^ Previously a Member State from 28 October 1965 to 31 December 1985.[8]
Jump up ^ Previously a Member State from 4 November 1946 (founding member) to 31 December 1956.
Jump up ^ Previously a Member State from 4 November 1946 (founding member) to 31 December 1985.
Jump up ^ Previously a Member State from 4 November 1946 (founding member) to 31 December 1984.
Jump up ^ Withdrawal notified on 12 October 2017, to take effect on 31 December 2018.[10]
^ Jump up to: a b The Netherlands Antilles became associate member of UNESCO on 26 October 1983. On Sunday 10 October 2010, a new status of the Netherlands Antilles came into effect within the Kingdom of the Netherlands, by which the country-status Dutch: land of the Netherlands Antilles ceased to exist. Under the new status, Curacao and Sint Maarten become countries (enjoying internal self-government within the Kingdom), joining Aruba, which gained similar status in 1986. Bonaire, Sint Eustatius and Saba have become part of the country "the Netherlands". The Netherlands retains responsibility for e.g. defence and foreign policy.
Jump up ^ As Macau before 1999
References[edit]
^ Jump up to: a b c UNESCO official site: List of the 195 Member States (and the 10 Associate Members) of UNESCO and the date on which they became members (or Associate Members) of the Organization, as of 5 November 2013
^ Jump up to: a b "General Conference admits Palestine as UNESCO Member". UNESCO. 31 October 2011. Retrieved 25 October 2017.
^ Jump up to: a b c d WADA. "Summary update on Government progress to become a State Party to the UNESCO International Convention against Doping in Sport" (PDF). p. 2. Archived from the original (PDF) on 16 January 2013. Retrieved 28 July 2009.
Jump up ^ "Kosovo Moves Closer To UNESCO Membership". Radio Free Europe/Radio Liberty. 21 October 2015. Retrieved 21 October 2015.
Jump up ^ "Kosovo's UNESCO Membership Bid Fails". 9 November 2015. Retrieved 8 December 2015.
Jump up ^ "UK Depositary Status List - Constitution of the United Nations Educational, Scientific and Cultural Organization (UNESCO)" (PDF). Foreign and Commonwealth Office. April 2016. Retrieved 2017-10-15.
Jump up ^ "Constitution of the United Nations Educational, Scientific and Cultural Organisation [as later amended]". Foreign and Commonwealth Office. Retrieved 2017-10-15.
Jump up ^ UNESCO. "UNESCO - Singapore - official relations". Retrieved 1 November 2011. [permanent dead link]
Jump up ^ UNESCO. "South Sudan – United Nations Educational, Scientific and Cultural Organization". Retrieved 19 August 2013.
Jump up ^ Nauert, Heather (12 October 2017). "The United States Withdraws From UNESCO". United States Department of State. Retrieved 25 October 2017.
^ Jump up to: a b "Israel to join US in quitting Unesco". BBC News. 12 October 2017. Retrieved 12 October 2017.
Jump up ^ UNESCO. "UNESCO Permanent delegations". Retrieved 1 November 2011.
^ Jump up to: a b c UNESCO. "Directory of Permanenent Delegations and Permanent Observer Missions to UNESCO". Retrieved 1 November 2011.
Jump up ^ UNESCO. "Directory of Permanent Delegations to UNESCO". Retrieved 1 November 2011.
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