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Hearst Castle
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Not to be confused with Hurst Castle, Henry VIII's Device Fort near Lymington in England.
Hearst San Simeon Estate
U.S. National Register of Historic Places
U.S. National Historic Landmark
California Historical Landmark #640[2]
The Casa Grande is the 60,645 square-foot centerpiece of Hearst Castle.
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Nearest city
San Simeon, California, United States
Area
More than 90,000 sq ft (8,400 m2)
Built
1919-1947[3]
Architect
Julia Morgan
Architectural style
Mediterranean Revival, other late 19th and 20th century Revivals
NRHP reference #
72000253[1]
CHISL #
640[2]
Significant dates
Added to NRHP
June 22, 1972
Designated CHISL
April 28, 1958
Hearst Castle is a National Historic Landmark and California Historical Landmark mansion located on the Central Coast of California, United States. Designed by architect Julia Morgan,[3] it was a residence for newspaper magnate William Randolph Hearst between 1919 and 1947. Hearst died in 1951, and it became a California State Park in 1954.[4] The site was opened to visitors in 1958.[5] Since that time, it has been operated as the Hearst San Simeon State Historical Monument, where the estate, and its considerable collection of art and antiques, is open for public tours. Despite its location far from any urban center, the site attracts "millions of travelers each year".[6]
Hearst formally named the estate "La Cuesta Encantada" ("The Enchanted Hill"), but usually called it "the ranch". Hearst Castle and grounds are also sometimes referred to as "San Simeon", without distinguishing between the Hearst property and the adjacent unincorporated area of the same name.
Contents
1 History
2 Location
2.1 Climate
3 Design
4 Size
5 Gallery
6 See also
7 References
8 Sources
9 External links
History[edit]
Invitations to Hearst Castle were highly coveted during its heyday in the 1920s and 1930s. The Hollywood and political elite often visited, usually flying into the estate's airfield or taking a private Hearst-owned train car from Los Angeles. Among Hearst's guests were Charlie Chaplin, Cary Grant, the Marx Brothers, Charles Lindbergh, Greta Garbo, Joan Crawford, Clark Gable, James Stewart, Bob Hope, Calvin Coolidge, Franklin Roosevelt, Dolores del Río, and Winston Churchill. While guests were expected to attend the formal dinners each evening, they were normally left to their own dwellings during the day while Hearst directed his business affairs. Since "the Ranch" had so many facilities, guests were rarely at a loss for things to do. The estate's theater usually screened films from Hearst's own movie studio, Cosmopolitan Productions.
Hearst Castle was the inspiration for the "Xanadu" mansion of the 1941 Orson Welles film Citizen Kane, a fictionalization of William Randolph Hearst's career.[7] Hearst Castle was not used as a location for the film, which instead used Oheka Castle in Huntington, New York, as well as buildings in San Diego's Balboa Park. Commercial filming is rare at Hearst Castle, and most requests are turned down. Since the property was donated to the state of California, only two projects have been granted permission: Stanley Kubrick's Spartacus, which used the castle to stand in as Crassus' villa, and Lady Gaga's music video for "G.U.Y."[8][9]
Spanish style guest house, designed by Julia Morgan
One condition of the Hearst Corporation's donation of the estate was that the Hearst family would be allowed to use it when they wished. Patty Hearst, a granddaughter of William Randolph, related that as a child, she hid behind statues in the Neptune Pool while tours passed by. Although the main estate is now a museum, the Hearst family continues to use an older Victorian house on the property as a retreat – the original house built by George Hearst in the late 19th century. The house is screened from tourist routes by a dense grove of eucalyptus to provide maximum privacy for the guests. In 2001, Patty Hearst hosted a Travel Channel show on the estate, and Amanda Hearst modeled for a fashion photo shoot at the estate for a Hearst Corporation magazine, Town and Country, in 2006.
Hearst Castle joined the National Register of Historic Places on June 22, 1972, and became a United States National Historic Landmark on May 11, 1976.
Hearst Castle was included as one of America's "10 Amazing Castles" by Forbes Travel.com. Forbes said, "Quite possibly the nation's most famous castle, William Randolph Hearst went to great lengths to bring back the best of European architecture – most notably ceilings from churches and monasteries – which were pieced back together in California to create his highly eclectic Central Coast getaway."[10]
Location[edit]
Zebras are a popular attraction on the Hearst ranch. They are descendants from Hearst's private zoo.
Hearst Castle is located near the unincorporated community of San Simeon, California, approximately 250 miles (400 km) from both Los Angeles and San Francisco, and 43 miles (69 km) from San Luis Obispo at the northern end of San Luis Obispo County. The estate itself is five miles (eight kilometers) inland atop a hill of the Santa Lucia Range at an altitude of 1,600 feet (490 m). The region is sparsely populated because the Santa Lucia Range abuts the Pacific Ocean, which provides dramatic seaside vistas but few opportunities for development and hampers transportation. The surrounding countryside visible from the mansion remains largely undeveloped. Its entrance is approximately five miles north of Hearst San Simeon State Park.
Hearst Castle was built on Rancho Piedra Blanca that William Randolph Hearst's father, George Hearst, originally purchased in 1865. The younger Hearst grew fond of this site over many childhood family camping trips. He inherited the ranch, which had grown to 250,000 acres (1,012 km2)[3] and 14 miles (23 km) of coastline, from his mother Phoebe Hearst in 1919.[11] Although the large ranch already had a Victorian mansion, the location selected for Hearst Castle was undeveloped, atop a steep hill whose ascent was a dirt path accessible only by foot or on horseback over 5 miles (8 km) of cutbacks.
Climate[edit]
The Hearst Castle area has a warm-summer mediterranean climate (Köppen csb) that is moderated by its relative proximity to the Pacific coastline.
hideClimate data for Hearst Castle
Month
Jan
Feb
Mar
Apr
May
Jun
Jul
Aug
Sep
Oct
Nov
Dec
Year
Average high °F (°C)
59.9
(15.5) 60.3
(15.7) 63.1
(17.3) 65.3
(18.5) 70.5
(21.4) 74.7
(23.7) 75
(23.9) 78.3
(25.7) 78.6
(25.9) 73.9
(23.3) 65.8
(18.8) 59.4
(15.2) 68.7
(20.4)
Daily mean °F (°C)
52.3
(11.3) 52.3
(11.3) 54.7
(12.6) 55.9
(13.3) 61.2
(16.2) 64.9
(18.3) 66
(18.9) 68
(20.0) 69.1
(20.6) 64
(17.8) 57.6
(14.2) 52
(11.1) 59.9
(15.5)
Average low °F (°C)
44.8
(7.1) 44.4
(6.9) 46.2
(7.9) 46.6
(8.1) 51.6
(10.9) 55.4
(13.0) 57
(13.9) 57.7
(14.3) 59.7
(15.4) 54.3
(12.4) 49.3
(9.6) 44.4
(6.9) 51.1
(10.6)
Average precipitation inches (mm)
5.5
(139.7) 5.902
(149.9) 4.5
(114.3) 1.902
(48.3) 0.701
(17.8) 0.098
(2.5) 0
(0) 0.098
(2.5) 0.299
(7.6) 1.402
(35.6) 2.799
(71.1) 5.799
(147.3) 28.902
(734.1)
Source: [12]
Design[edit]
Owner William Randolph Hearst with architect Julia Morgan in 1926. Photograph by Irvin Willat.
Hearst first approached American architect Julia Morgan[3] with ideas for a new project in April 1919, shortly after he took ownership. Hearst's original idea was to build a bungalow, according to a draftsman who worked in Morgan's office who recounted Hearst's words from the initial meeting:
I would like to build something upon the hill at San Simeon. I get tired of going up there and camping in tents. I'm getting a little too old for that. I'd like to get something that would be a little more comfortable.[13]
After approximately one month of discussion, Hearst's original idea for a modest dwelling swelled to grand proportions. Discussion for the exterior style switched from initial ideas of Japanese and Korean themes to the Spanish Revival that was gaining popularity and that Morgan had furthered with her work on the Los Angeles Herald Examiner headquarters in 1915. Hearst was fond of Spanish Revival, but dissatisfied with the crudeness of the colonial structures in California. Mexican colonial architecture had more sophistication but he objected to its profusion of ornamentation. Turning to the Iberian Peninsula for inspiration, he found Renaissance and Baroque examples in southern Spain more to his tastes. Hearst particularly admired a church in Ronda and asked Morgan to pattern the Main Building towers after it. The Panama-California Exposition of 1915 in San Diego held the closest approaches in California to the look Hearst desired. He decided to substitute a stucco exterior in place of masonry in deference to Californian traditions.
Tower of the Church of Santa María la Mayor, in Ronda, Spain, which served as inspiration for the Hearst Castle towers.
By late summer 1919, Morgan had surveyed the site, analyzed its geology, and drawn initial plans for the Main Building. Construction began in 1919 and continued through 1947 when Hearst stopped living at the estate due to ill health.[3] Morgan persuaded Hearst to begin with the guest cottages, because the smaller structures could be completed more quickly.
Hearst Castle as seen from the southwest
The estate is a pastiche of historic architectural styles that its owner admired in his travels around Europe. Hearst was a prolific buyer who did not so much purchase art and antiques to furnish his home as he did build his home to get his bulging collection out of warehouses. This led to incongruous elements, such as the private cinema, whose walls were lined with shelves of rare books. The floor plan of the Main Building is chaotic due to his habit of buying centuries-old ceilings, which dictated the proportions and decor of various rooms.

Hearst Castle featured 56 bedrooms, 61 bathrooms, 19 sitting rooms, 127 acres (0.5 km2)[3] of gardens, indoor and outdoor swimming pools, tennis courts, a movie theater, an airfield, and the world's largest private zoo. Zebras and other exotic animals still roam the grounds. Morgan, an accomplished civil engineer, devised a gravity-based water delivery system that transports water from artesian wells on the slopes of Pine Mountain, a 3,500-foot-high (1,100 m) peak 7 miles (11 km) east of Hearst Castle, to a reservoir on Rocky Butte, a 2,000-foot (610 m) knoll less than a mile southeast from Hearst Castle.[14]
The Neptune Pool at Hearst Castle was rebuilt three times to suit its owner's tastes. Its centerpiece is the façade of an ancient Roman temple Hearst imported to California.
One highlight of the estate is the outdoor Neptune Pool, located near the edge of the hilltop, which offers an expansive vista of the mountains, ocean and the main house. Due to drought conditions and leaks in the pool, it is now drained.[15][16] The Neptune Pool patio features an ancient Roman temple front, transported wholesale from Europe and reconstructed at the site. Hearst was an inveterate tinkerer, and would tear down structures and rebuild them on a whim. For example, the Neptune Pool was rebuilt three times before Hearst was satisfied. As a consequence of Hearst's persistent design changes, the estate was never completed in his lifetime.
Although Hearst Castle's ornamentation is borrowed from historic European themes, its underlying structure is primarily steel reinforced concrete. The use of modern engineering techniques reflects Morgan's background as a civil engineering graduate of the University of California, Berkeley and the first female architecture graduate of the École nationale supérieure des Beaux-Arts in Paris. Morgan designed tiles for the castle, and used several tile companies to produce them: Solon & Schemmel (San Jose), Grueby Faience Company (Boston), Batchelder (Pasadena), and California Faience (Berkeley). Albert Solon and Frank Schemmel came to Hearst Castle to perform the tile work.[17] Camille Solon, son of Marc-Louis-Emmanuel Solon and brother of Albert Solon of Solon and Schemmel Tile Company, designed the mosaics of blue-and-gold Venetian glass tile used in the property's indoor Roman Pool and he also designed and painted the murals in the Gothic Study of Casa Grande.[18]
Hearst Castle Roman Pool in September 2012
During Hearst's ownership, a private power plant supplied electricity to the remote location. Most of the estate's chandeliers have bare light bulbs, because electrical technology was so new when Hearst Castle was built.
Size[edit]
The total square footage of the buildings on the estate exceeds 90,000 square feet (8,300 m²). The area of Casa Grande, the "castle", is 60,645 square feet (5,634 m²). The areas of the guest houses on the property are:[19]
Casa del Mar: 5,875 square feet (546 m²)
Casa del Monte: 2,291 square feet (213 m²)
Casa del Sol: 2,604 square feet (242 m²)
Gallery[edit]
Egyptian Artifacts - Oldest Pieces In The Collection
Door detail, Casa del Monte
Hearst Castle dining room
Terrace and vista
See also[edit]
California portal
Hearst San Simeon State Park
Neptune Pool
Wyntoon
Xanadu (Citizen Kane)
List of largest houses in the United States
References[edit]
Jump up ^ National Park Service (2007-01-23). "National Register Information System". National Register of Historic Places. National Park Service.
Jump up ^ "Hearst San Simeon State Historical Monument". Office of Historic Preservation, California State Parks. Retrieved 2014-04-14.
^ Jump up to: a b c d e f Fodor's (21 December 2010). Fodor's Northern California 2011: With Napa, Sonoma, Yosemite, San Francisco & Lake Tahoe. Random House Digital, Inc. p. 88. ISBN 978-1-4000-0503-1. Retrieved 25 December 2011.
Jump up ^ "State Park Notes". Planning and Civic Comment. 20 (3): 27. September 1954.
Jump up ^ "State Park Notes". Planning and Civic Comment. 24 (3): 58. September 1958. The Hearst San Simeon State Historical Monument was dedicated and opened to the public June 2
Jump up ^ "Hearst Castle". San Simeon welcomes you. San Simeon Chamber of Commerce. Retrieved 2014-07-09.
Jump up ^ "The top houses from the movies". Daily Telegraph.
Jump up ^ Linn, Sarah (February 11, 2014). "Hearst Castle goes Gaga as pop diva shoots video this week at estate". The Cambrian. Archived from the original on March 31, 2014. Retrieved March 31, 2014.
Jump up ^ Dodge, Shyam (April 15, 2014). "Lady Gaga denies claim she 'wasted 356K gallons of water to fill massive pool at Hearst Castle' during worst drought in California history". MailOnline.com. Daily Mail.
Jump up ^ "News Release: Hearst Castle and Vikingsholm named by Forbes Travel.com" (PDF). California Department of Parks and Recreation. February 10, 2009. See also "In Pictures: Great American Castles". Forbes.com. December 10, 2010. p. 7.
Jump up ^ Hearst Castle history
Jump up ^ "San Simeon - Hearst Castle, California Temperature Averages". Weaterbase. Retrieved 7 October 2016.
Jump up ^ Mark A. Wilson; forward by Lynn Forney McMurray (2007). Julia Morgan: Architect of Beauty. Gibbs Smith. p. 105. ISBN 978-1-4236-0088-6. Retrieved 27 May 2009.
Jump up ^ Garden and Vistas - Tour Information Archived 2010-08-30 at the Wayback Machine. from HearstCastle.org
Jump up ^ "Drought forces Hearst Castle to pool resources". San Louis Obispo Tribune. February 4, 2014.
Jump up ^ Ami Lieu (July 2, 2014). "Drought prompts Hearst Castle to close restrooms, drain pool". Social Wanderer. KCET.
Jump up ^ Wilson: Julia Morgan: Architect of Beauty, 2007, p. xi
Jump up ^ "Guide to the Camille Solon Drawings Collection, 1900-1952".
Jump up ^ Facts and Stats from the official Hearst Castle website
Sources[edit]
This article includes a list of references, but its sources remain unclear because it has insufficient inline citations. Please help to improve this article by introducing more precise citations. (February 2014) (Learn how and when to remove this template message)
Lewis, O. (1958). Fabulous San Simeon; a history of the Hearst Castle, a Calif. state monument located on the scenic coast of Calif., together with a guide to the treasures on display. San Francisco: California Historical Society.
Collord, M., & Miller, A. (1972). Castle fare: featuring authentic recipes served in Hearst Castle. San Luis Obispo, CA: Blake Printery.
Boulian, D. M. (1972). Enchanted gardens of Hearst Castle. Cambria, Calif: Phildor Press.
Martin, C. (1977). Hearst Castle: mythology, legend, history in art. Cambria, Calif: Galatea Publications.
Coffman, T. (1985).
Morgan, J., Hearst, W. R., & Loe, N. E. (1987). San Simeon revisited: the correspondence between architect Julia Morgan and William Randolph Hearst. San Luis Obispo, Calif: Library Associates, California Polytechnic State University.
Blades, J., Nargizian, R. A., & Carr, G. (1993). The Hearst Castle collection of carpets: fine rug reproductions. Santa Barbara, Calif: Jane Freeburg.
Kastner, V. (1994). Remains to be seen: remains of Spanish ceilings at Hearst Castle. San Simeon, CA: Hearst San Simeon State Historic Monument.
Loe, N. E. (1994). Hearst Castle: an interpretive history of W.R. Hearst's San Simeon estate. [S.l.]: ARA Services.
Sullivan, J. (1996). Castle chronicles: : "sketching around Hearst Castle". Los Osos, Calif: The Bay News?.
California. (2001). Hearst Castle: Hearst San Simeon State Historical Monument. Sacramento, CA: California State Parks.
External links[edit]
Wikimedia Commons has media related to Hearst Castle.
Official website
Hearst Castle Virtual Tour
California State Parks web page
The Mosaics of Hearst Castle
Hearst Castle Press
National Geographic Theater at Hearst Castle – Featuring the Hearst Castle Experience
Coordinates: 35°41′07″N 121°10′00″W / 35.6852°N 121.1666°W / 35.6852; -121.1666
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Retrieved from "https://en.wikipedia.org/w/index.php?title=Hearst_Castle&oldid=851474774"
Categories:
State parks of California
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Fluid mosaic model
From Wikipedia, the free encyclopedia
Jump to: navigation, search
Fluid mosaic model of a cell membrane
The fluid mosaic model explains various observations regarding the structure of functional cell membranes. The model, which was devised by SJ Singer and GL Nicolson in 1972, describes the cell membrane as a two-dimensional liquid that restricts the lateral diffusion of membrane components. Such domains are defined by the existence of regions within the membrane with special lipid and protein composition that promote the formation of lipid rafts or protein and glycoprotein complexes. Another way to define membrane domains is the association of the lipid membrane with the cytoskeleton filaments and the extracellular matrix through membrane proteins.[1] The current model describes important features relevant to many cellular processes, including: cell-cell signaling, apoptosis, cell division, membrane budding, and cell fusion.
Contents
[hide]
1 Chemical Makeup
2 Experimental evidence
3 Subsequent developments
3.1 Membrane asymmetry
3.2 Non-bilayer membranes
3.3 Membrane curvature
3.4 Lipid movement within the membrane
4 Restrictions to bilayer fluidity
4.1 Lipid rafts
4.2 Protein complexes
4.3 Cytoskeletal fences (corrals) and binding to the extracellular matrix
5 Historical timeline
6 Notes and references
Chemical Makeup[edit]
Chemically a cell membrane is composed of four components: Phospholipids, Proteins, Carbohydrates, Cholesterol
Experimental evidence[edit]
The fluid property of functional biological membranes had been determined through labeling experiments, x-ray diffraction, and calorimetry. These studies showed that integral membrane proteins diffuse at rates affected by the viscosity of the lipid bilayer in which they were embedded, and demonstrated that the molecules within the cell membrane are dynamic rather than static.[2]
Previous models of biological membranes included the Robertson Unit Membrane Model and the Davidson-Danielli Tri-Layer model.[1] These models had proteins present as sheets neighboring a lipid layer, rather than incorporated into the phospholipid bilayer. Other models described repeating, regular units of protein and lipid. These models were not well supported by microscopy and thermodynamic data, and did not accommodate evidence for dynamic membrane properties.[1]
An important experiment that provided evidence supporting fluid and dynamic biological was performed by Frye and Edidin. They used Sendai virus to force human and mouse cells to fuse and form a heterokaryon. Using antibody staining, they were able to show that the mouse and human proteins remained segregated to separate halves of the heterokaryon a short time after cell fusion. However, the proteins eventually diffused and over time the border between the two halves was lost. Lowering the temperature slowed the rate of this diffusion by causing the membrane phospholipids to transition from a fluid to a gel phase.[3] Singer and Nicholson rationalized the results of these experiments using their fluid mosaic model.[2]
The fluid mosaic model explains changes in structure and behavior of cell membranes under different temperatures, as well as the association of membrane proteins with the membranes. While Singer and Nicolson had substantial evidence drawn from multiple subfields to support their model, recent advances in fluorescence microscopy and structural biology have validated the fluid mosaic nature of cell membranes.
Subsequent developments[edit]
Membrane asymmetry[edit]
Additionally, the two leaflets of biological membranes are asymmetric and divided into subdomains composed of specific proteins or lipids, allowing spatial segregation of biological processes associated with membranes. Cholesterol and cholesterol-interacting proteins can concentrate into lipid rafts and constrain cell signaling processes to only these rafts.[4] Another form of asymmetry was shown by the work of Mouritsen and Bloom in 1984, where they proposed a Mattress Model of lipid-protein interactions to address the biophysical evidence that the membrane can range in thickness and hydrophobicity of proteins.[5] Phospholipids can be asymmetric through the active translocation of a phospholipid type from one leaflet of the membrane to the other. This process is controlled by flippase and scramblase enzymes.[6] A biological example of this is the segregation of phosphatidylserine to be highly enriched on the cytosolic side of the plasma membrane. Phosphatidylserine localization to the outer leaflet can trigger immune responses.[7]
Non-bilayer membranes[edit]
The existence of non-bilayer lipid formations with important biological functions was confirmed subsequent to publication of the fluid mosaic model. These membrane structures may be useful when the cell needs to propagate a non bilayer form, which occurs during cell division and the formation of a gap junction.[8]
Membrane curvature[edit]
The membrane bilayer is not always flat. Local curvature of the membrane can be caused by the asymmetry and non-bilayer organization of lipids as discussed above. More dramatic and functional curvature is achieved through BAR domains, which bind to phosphatidylinositol on the membrane surface, assisting in vesicle formation, organelle formation and cell division.[9] Curvature development is in constant flux and contributes to the dynamic nature of biological membranes.[10]
Lipid movement within the membrane[edit]
During the decade of 1970, it was acknowledged that individual lipid molecules undergo free lateral diffusion within each of the layers of the lipid membrane.[11] Diffusion occurs at a high speed, with an average lipid molecule diffusing ~2 µm, approximately the length of a large bacterial cell, in about 1 second.[11] It has also been observed that individual lipid molecules rotate rapidly around their own axis.[11] Moreover, phospholipid molecules can, although they seldom do, migrate from one side of the lipid bilayer to the other (a process known as flip-flop). However, flip-flop might be enhanced by flippase enzymes. The processes described above influence the disordered nature of lipid molecules and interacting proteins in the lipid membranes, with consequences to membrane fluidity, signaling, trafficking and function.
Restrictions to bilayer fluidity[edit]
There are restrictions to the lateral mobility of the lipid and protein components in the fluid membrane imposed by the formation of subdomains within the lipid bilayer. These subdomains arise by several processes e.g. binding of membrane components to the extracellular matrix, nanometric membrane regions with a particular biochemical composition that promote the formation of lipid rafts and protein complexes mediated by protein-protein interactions.[1] Furthermore, protein-cytoskeleton associations mediate the formation of “cytoskeletal fences”, corrals wherein lipid and membrane proteins can diffuse freely, but that they can seldom leave.[1] Restriction on lateral diffusion rates of membrane components is very important because it allows the functional specialization of particular regions within the cell membranes.
Lipid rafts[edit]
Lipid rafts are membrane nanometric platforms with a particular lipid and protein composition that laterally diffuse, navigating on the liquid bilipid layer. Sphingolipids and cholesterol are important building blocks of the lipid rafts.[12]
Protein complexes[edit]
Cell membrane proteins and glycoproteins do not exist as single elements of the lipid membrane, as first proposed by Singer and Nicolson in 1972. Rather, they occur as diffusing complexes within the membrane.[1] The assembly of single molecules into these macromolecular complexes has important functional consequences for the cell; such as ion and metabolite transport, signaling, cell adhesion, and migration.[1]
Cytoskeletal fences (corrals) and binding to the extracellular matrix[edit]
Some proteins embedded in the bilipid layer interact with the extracellular matrix outside the cell, cytoskeleton filaments inside the cell, and septin ring-like structures. These interactions have a strong influence on shape and structure, as well as on compartmentalization. Moreover, they impose physical constraints that restrict the free lateral diffusion of proteins and at least some lipids within the bilipid layer.[1]
When integral proteins of the lipid bilayer are tethered to the extracellular matrix, they are unable to diffuse freely. Proteins with a long intracellular domain may collide with a fence formed by cytoskeleton filaments.[13] Both processes restrict the diffusion of proteins and lipids directly involved, as well as of other interacting components of the cell membranes.
S.cerevisiae septins
Septin ring-like structures (in green) can pinch cell membranes and split them into subdomains.
Septins are a family of GTP-binding proteins highly conserved among eukaryotes. Prokaryotes have similar proteins called paraseptins. They form compartmentalizing ring-like structures strongly associated with the cell membranes. Septins are involved in the formation of structures such as, cilia and flagella, dendritic spines, and yeast buds.[14]
Historical timeline[edit]
1895 – Ernest Overton hypothesized that cell membranes are made out of lipids.[15]
1925 – Evert Gorter and François Grendel found that red blood cell membranes are formed by a fatty layer two molecules thick, i.e. they described the bilipid nature of the cell membrane.[16]
1935 – Hugh Davson and James Danielli proposed that lipid membranes are layers composed by proteins and lipids with pore-like structures that allow specific permeability for certain molecules. Then, they suggested a model for the cell membrane, consisting of a lipid layer surrounded by protein layers at both sides of it.[17]
1957 – J. David Robertson, based on electron microscopy studies, establishes the "Unit Membrane Hypothesis". This, states that all membranes in the cell, i.e. plasma and organelle membranes, have the same structure: a bilayer of phospholipids with monolayers of proteins at both sides of it.[18]
1972 – SJ Singer and GL Nicolson proposed the fluid mosaic model as an explanation for the data and latest evidence regarding the structure and thermodynamics of cell membranes.[2]
Notes and references[edit]
^ Jump up to: a b c d e f g h Nicolson GL (2014). "The Fluid—Mosaic Model of Membrane Structure: Still relevant to understanding the structure, function and dynamics of biological membranes after more than 40 years". Biochimica et Biophysica Acta. 1838 (6): 1451–146. PMID 24189436. doi:10.1016/j.bbamem.2013.10.019.
^ Jump up to: a b c Singer SJ, Nicolson GL (Feb 1972). "The fluid mosaic model of the structure of cell membranes". Science. 175 (4023): 720–31. PMID 4333397. doi:10.1126/science.175.4023.720.
Jump up ^ Frye LD, Edidin M (1970). "The rapid intermixing of cell surface antigens after formation of mouse-human heterokaryons". J Cell Sci. 7 (2): 319–35. PMID 4098863.
Jump up ^ Silvius JR (2005). "Partitioning of membrane molecules between raft and non-raft domains: Insights from model-membrane studies". Biochimica et Biophysica Acta. 1746 (3): 193–202. PMID 16271405. doi:10.1016/j.bbamcr.2005.09.003.
Jump up ^ Mouritsen OG, Bloom M (1984). "Mattress model of lipid-protein interactions in membranes". Biophys J. 46 (2): 141–153. PMC 1435039  . PMID 6478029. doi:10.1016/S0006-3495(84)84007-2.
Jump up ^ Daleke DL (2003). "Regulation of transbilayer plasma membrane phospholipid asymmetry". J Lipid Res. 44 (2): 233–242. PMID 12576505. doi:10.1194/jlr.R200019-JLR200.
Jump up ^ Fadok, VA; et al. (1998). "The role of phosphatidylserine in recognition of apoptotic cells by phagocytes". Cell Death and Differ. 5 (7): 551–562. PMID 10200509. doi:10.1038/sj.cdd.4400404.
Jump up ^ van den Brink-van der Laan E; et al. (2004). "Nonbilayer lipids affect peripheral and integral membrane proteins via changes in the lateral pressure profile". Biochim Biophys Acta. 1666 (1–2): 275–88. PMID 15519321. doi:10.1016/j.bbamem.2004.06.010.
Jump up ^ Frost A; et al. (2009). "The BAR domain superfamily: membrane-molding macromolecules". Cell. 137 (2): 191–6. PMID 19379681. doi:10.1016/j.cell.2009.04.010.
Jump up ^ Rodríguez-García R; et al. (2009). "Bimodal spectrum for the curvature fluctuations of bilayer vesicles: pure bending plus hybrid curvature-dilation modes". Phys Rev Lett. 102 (12): 128101. PMID 19392326. doi:10.1103/PhysRevLett.102.128101.
^ Jump up to: a b c Alberts B, Johnson A, Lewis J, et al. (2008). Molecular Biology of the Cell (5th ed.). New York: Garland Science. pp. 621–622. ISBN 978-0-8153-4105-5.
Jump up ^ Lingwood D, Simons K (2010). "Lipid rafts as a membrane-organizing principle". Science. 327 (5961): 46–50. PMID 20044567. doi:10.1126/science.1174621.
Jump up ^ G. Vereb; et al. (2003). "Dynamic, yet structured: The cell membrane three decades after the Singer–Nicolson model". PNAS. 100 (14): 8053–8058. PMC 166180  . PMID 12832616. doi:10.1073/pnas.1332550100.
Jump up ^ Juha Saarikangas; Yves Barral (2011). "The emerging functions of septins in metazoans". EMBO Reports. 12 (11): 1118–1126. PMC 3207108  . PMID 21997296. doi:10.1038/embor.2011.193.
Jump up ^ Overton, E (1895). "Uberdie osmotischen Eigenshafter der Lebenden Pflanzen und tierzelle". Vjschr Naturf Ges Zurich. 40: 159–201.
Jump up ^ E. Gorter; F. Grendel (1925). "On Biomolecular Layers of Lipoids on the Chromocytes of the Blood". Journal of Experimental Medicine. 41: 439–443. PMC 2130960  . PMID 19868999. doi:10.1084/jem.41.4.439.
Jump up ^ James Danielli; Hugh Davson (1935). "A contribution to the theory of permeability of thin films". Journal of Cellular and Comparative Physiology. 5 (4): 495–508. doi:10.1002/jcp.1030050409.
Jump up ^ John E. Heuser (1995). "In Memory of J. David Robertson" (PDF). Newsletter of the American Society of Cell Biology.
[show]
v
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Structures of the cell / organelles (TH H1.00.01.2-3)
Endomembrane system
Cell membrane
Nucleus
Nucleolus
Endoplasmic reticulum
Golgi apparatus
Parenthesome
Autophagosome
Vesicles
Exosome
Lysosome
Endosome
Phagosome
Vacuole
Acrosome
Cytoplasmic granules
Melanosome
Microbody
Glyoxysome
Peroxisome
Weibel–Palade body
Cytoskeleton
Microfilaments
Intermediate filaments
Microtubules
Prokaryotic cytoskeleton
MTOCs
Centrosome
Centriole
Basal body
Spindle pole body
Myofibril
Endosymbionts
Mitochondrion
Plastids
Chloroplast
Chromoplast
Gerontoplast
Leucoplast
Amyloplast
Elaioplast
Proteinoplast
Tannosome
Other internal
RNA
Ribosome
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Cytoplasm
Proteasome
External
Undulipodium
Cilium
Flagellum
Axoneme
Radial spoke
Extracellular matrix
Cell wall
[show]
v
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Structures of the cell membrane
Membrane lipids
Lipid bilayer
Phospholipids
Proteolipids
Sphingolipids
Sterols
Membrane proteins
Membrane glycoproteins
Integral membrane proteins/transmembrane protein
Peripheral membrane protein/Lipid-anchored protein
Other
Caveolae/Coated pits
Cell junctions
Glycocalyx
Lipid raft/microdomains
Membrane contact sites
Membrane nanotubes
Myelin sheath
Nodes of Ranvier
Nuclear envelope
Phycobilisomes
Porosomes
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