text:
Line segment
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Not to be confused with arc (geometry).
The geometric definition of a closed line segment: the intersection of all points at or to the right of A with all points at or to the left of B
historical image – create a line segment (1699)
In geometry, a line segment is a part of a line that is bounded by two distinct end points, and contains every point on the line between its endpoints. A closed line segment includes both endpoints, while an open line segment excludes both endpoints; a half-open line segment includes exactly one of the endpoints.
Examples of line segments include the sides of a triangle or square. More generally, when both of the segment's end points are vertices of a polygon or polyhedron, the line segment is either an edge (of that polygon or polyhedron) if they are adjacent vertices, or otherwise a diagonal. When the end points both lie on a curve such as a circle, a line segment is called a chord (of that curve).
Contents
[hide]
1 In real or complex vector spaces
2 Properties
3 In proofs
4 As a degenerate ellipse
5 In other geometric shapes
5.1 Triangles
5.2 Quadrilaterals
5.3 Circles and ellipses
6 Directed line segment
7 Generalizations
8 See also
9 References
10 External links
In real or complex vector spaces[edit]
If V is a vector space over R {\displaystyle \mathbb {R} } or C {\displaystyle \mathbb {C} } , and L is a subset of V, then L is a line segment if L can be parameterized as
L = { u + t v ∣ t ∈ [ 0 , 1 ] } {\displaystyle L=\{\mathbf {u} +t\mathbf {v} \mid t\in [0,1]\}}
for some vectors u , v ∈ V {\displaystyle \mathbf {u} ,\mathbf {v} \in V\,\!} , in which case the vectors u and u + v are called the end points of L.
Sometimes one needs to distinguish between "open" and "closed" line segments. Then one defines a closed line segment as above, and an open line segment as a subset L that can be parametrized as
L = { u + t v ∣ t ∈ ( 0 , 1 ) } {\displaystyle L=\{\mathbf {u} +t\mathbf {v} \mid t\in (0,1)\}}
for some vectors u , v ∈ V {\displaystyle \mathbf {u} ,\mathbf {v} \in V\,\!} .
Equivalently, a line segment is the convex hull of two points. Thus, the line segment can be expressed as a convex combination of the segment's two end points.
In geometry, it is sometimes defined that a point B is between two other points A and C, if the distance AB added to the distance BC is equal to the distance AC. Thus in R 2 {\displaystyle \mathbb {R} ^{2}} the line segment with endpoints A = (ax, ay) and C = (cx, cy) is the following collection of points:
{ ( x , y ) | ( x − c x ) 2 + ( y − c y ) 2 + ( x − a x ) 2 + ( y − a y ) 2 = ( c x − a x ) 2 + ( c y − a y ) 2 } {\displaystyle \{(x,y)|{\sqrt {(x-c_{x})^{2}+(y-c_{y})^{2}}}+{\sqrt {(x-a_{x})^{2}+(y-a_{y})^{2}}}={\sqrt {(c_{x}-a_{x})^{2}+(c_{y}-a_{y})^{2}}}\}} .
Properties[edit]
A line segment is a connected, non-empty set.
If V is a topological vector space, then a closed line segment is a closed set in V. However, an open line segment is an open set in V if and only if V is one-dimensional.
More generally than above, the concept of a line segment can be defined in an ordered geometry.
A pair of line segments can be any one of the following: intersecting, parallel, skew, or none of these. The last possibility is a way that line segments differ from lines: if two nonparallel lines are in the same Euclidean plane they must cross each other, but that need not be true of segments.
In proofs[edit]
In an axiomatic treatment of geometry, the notion of betweenness is either assumed to satisfy a certain number of axioms, or else be defined in terms of an isometry of a line (used as a coordinate system).
Segments play an important role in other theories. For example, a set is convex if the segment that joins any two points of the set is contained in the set. This is important because it transforms some of the analysis of convex sets to the analysis of a line segment. The Segment Addition Postulate can be used to add congruent segment or segments with equal lengths and consequently substitute other segments into another statement to make segments congruent.
As a degenerate ellipse[edit]
A line segment can be viewed as a degenerate case of an ellipse in which the semiminor axis goes to zero, the foci go to the endpoints, and the eccentricity goes to one. A standard definition of an ellipse is the set of points for which the sum of a point's distances to two foci is a constant; if this constant equals the distance between the foci, the line segment is the result. A complete orbit of this ellipse traverses the line segment twice. As a degenerate orbit this is a radial elliptic trajectory.
In other geometric shapes[edit]
In addition to appearing as the edges and diagonals of polygons and polyhedra, line segments appear in numerous other locations relative to other geometric shapes.
Triangles[edit]
Some very frequently considered segments in a triangle include the three altitudes (each perpendicularly connecting a side or its extension to the opposite vertex), the three medians (each connecting a side's midpoint to the opposite vertex), the perpendicular bisectors of the sides (perpendicularly connecting the midpoint of a side to one of the other sides), and the internal angle bisectors (each connecting a vertex to the opposite side). In each case there are various equalities relating these segment lengths to others (discussed in the articles on the various types of segment) as well as various inequalities.
Other segments of interest in a triangle include those connecting various triangle centers to each other, most notably the incenter, the circumcenter, the nine-point center, the centroid, and the orthocenter.
Quadrilaterals[edit]
In addition to the sides and diagonals of a quadrilateral, some important segments are the two bimedians (connecting the midpoints of opposite sides) and the four maltitudes (each perpendicularly connecting one side to the midpoint of the opposite side).
Circles and ellipses[edit]
Any straight line segment connecting two points on a circle or ellipse is called a chord. Any chord in a circle which has no longer chord is called a diameter, and any segment connecting the circle's center (the midpoint of a diameter) to a point on the circle is called a radius.
In an ellipse, the longest chord, which is also the longest diameter, is called the major axis, and a segment from the midpoint of the major axis (the ellipse's center) to either endpoint of the major axis is called a semi-major axis. Similarly, the shortest diameter of an ellipse is called the minor axis, and the segment from its midpoint (the ellipse's center) to either of its endpoints is called a semi-minor axis. The chords of an ellipse which are perpendicular to the major axis and pass through one of its foci are called the latera recta of the ellipse. The interfocal segment connects the two foci.
Directed line segment[edit]
When a line segment is given an orientation (direction) it suggests a translation or perhaps a force tending to make a translation. The magnitude and direction are indicative of a potential change. This suggestion has been absorbed into mathematical physics through the concept of a Euclidean vector.[1][2] The collection of all directed line segments is usually reduced by making "equivalent" any pair having the same length and orientation.[3] This application of an equivalence relation dates from Giusto Bellavitis’s introduction of the concept of equipollence of directed line segments in 1835.
Generalizations[edit]
Analogous to straight line segments above, one can define arcs as segments of a curve.
See also[edit]
Broken line
Interval (mathematics)
Line (geometry)
Line segment intersection, the algorithmic problem of finding intersecting pairs in a collection of line segments
Spirangle
Segment addition postulate
References[edit]
Jump up ^ Harry F. Davis & Arthur David Snider (1988) Introduction to Vector Analysis, 5th edition, page 1, Wm. C. Brown Publishers ISBN 0-697-06814-5
Jump up ^ Matiur Rahman & Isaac Mulolani (2001) Applied Vector Analysis, pages 9 & 10, CRC Press ISBN 0-8493-1088-1
Jump up ^ Eutiquio C. Young (1978) Vector and Tensor Analysis, pages 2 & 3, Marcel Dekker ISBN 0-8247-6671-7
David Hilbert The Foundations of Geometry. The Open Court Publishing Company 1950, p. 4
External links[edit]
Wikimedia Commons has media related to Line segment.
Look up line segment in Wiktionary, the free dictionary.
Weisstein, Eric W. "Line segment". MathWorld.
Line Segment at PlanetMath
Definition of line segment With interactive animation
Copying a line segment with compass and straightedge
Dividing a line segment into N equal parts with compass and straightedge Animated demonstration
This article incorporates material from Line segment on PlanetMath, which is licensed under the Creative Commons Attribution/Share-Alike License.
Retrieved from "https://en.wikipedia.org/w/index.php?title=Line_segment&oldid=837955182"
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text:
List of World Heritage Sites in Russia
From Wikipedia, the free encyclopedia
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This a list of UNESCO World Heritage Sites in Russia. Currently, there are 28 World Heritage Sites in Russia.[1] 17 properties are cultural and 11 are natural. 24 more properties are on UNESCO's tentative list.[2]
Contents
[hide]
1 List of World Heritage Sites
2 Tentative list
3 See also
4 References
List of World Heritage Sites[edit]
Komi Forests
Lake Baikal
Volcanoes of Kamchatka
Altai Mountains
Western Caucasus
Sikhote-Alin
Uvs Nuur Basin
Wrangel Island
Putorana Plateau
Lena Pillars
Saint Petersburg
Kizhi Pogost
Moscow Kremlin and Red Square, Novodevichy Convent, Kolomenskoye
Veliky Novgorod
Solovetsky Islands
Vladimir and Suzdal
Trinity Sergius Lavra
Kazan Kremlin
Ferapontov Monastery
Curonian Spit
Derbent
Yaroslavl
Struve Geodetic Arc
Bolgar
Sviyazhsk
Location of UNESCO World Heritage Sites within Russia
Legend: World Cultural Heritage Site; World Natural Heritage Site
The table lists information about each World Heritage Site:
Name: as listed by the World Heritage Committee[1]
Location: city (surrounding or nearest, if applicable), Federal subject, coordinates
Period: time period of significance, typically of construction
Description: brief description of the site
Year listed: the year the site was inscribed on the World Heritage List
Criteria: the criteria the site was listed under
Reference number: UNESCO site reference number
Name
Image
Location
Year
listed
Criteria
Ref.
num.
Period
Description
Natural
Virgin Komi Forests Komi Republic
65°4′0″N 60°9′0″E / 65.06667°N 60.15000°E / 65.06667; 60.15000 (Virgin Komi Forests) 1995 vii, ix 719 – The Virgin Komi Forests cover 3.28 million ha of tundra and mountain tundra in the Urals, as well as one of the most extensive areas of virgin boreal forest remaining in Europe. This vast area of conifers, aspens, birches, peat bogs, rivers and natural lakes has been monitored and studied for over 50 years. It provides valuable evidence of the natural processes affecting biodiversity in the taiga.
Lake Baikal Irkutsk Oblast and Buryatia
53°10′25″N 107°39′45″E / 53.17361°N 107.66250°E / 53.17361; 107.66250 (Lake Baikal) 1996 vii, viii, ix, x 754 – Situated in south-east Siberia, the 3.15-million-ha Lake Baikal is the oldest (25 million years) and deepest (1,700 m) lake in the world. It contains 20% of the world's total unfrozen freshwater reserve. Known as the 'Galapagos of Russia', its age and isolation have produced one of the world's richest and most unusual freshwater faunas, which is of exceptional value to evolutionary science.
Volcanoes of Kamchatka Kamchatka Krai
56°20′0″N 158°30′0″E / 56.33333°N 158.50000°E / 56.33333; 158.50000 (Volcanoes of Kamchatka) 1996 vii, viii, ix, x 765 – This is one of the most outstanding volcanic regions in the world, with a high density of active volcanoes, a variety of types, and a wide range of related features. The six sites included in the serial designation group together the majority of volcanic features of the Kamchatka peninsula. The interplay of active volcanoes and glaciers forms a dynamic landscape of great beauty. The sites contain great species diversity, including the world's largest known variety of salmonoid fish and exceptional concentrations of sea otter, brown bear and Steller's sea eagle.
Golden Mountains of Altai Altai Republic
50°28′0″N 86°0′0″E / 50.46667°N 86.00000°E / 50.46667; 86.00000 (Golden Mountains of Altai) 1998 x 768 – The Altai mountains in southern Siberia form the major mountain range in the western Siberia biogeographic region and provide the source of its greatest rivers – the Ob and the Irtysh. Three separate areas are inscribed: Altaisky Zapovednik and a buffer zone around Lake Teletskoye; Katunsky Zapovednik and a buffer zone around Mount Belukha; and the Ukok Quiet Zone on the Ukok plateau. The total area covers 1,611,457 ha. The region represents the most complete sequence of altitudinal vegetation zones in central Siberia, from steppe, forest-steppe, mixed forest, subalpine vegetation to alpine vegetation. The site is also an important habitat for endangered animal species such as the snow leopard.
Western Caucasus Krasnodar Krai and Adygea
44°0′0″N 40°0′0″E / 44.00000°N 40.00000°E / 44.00000; 40.00000 (Western Caucasus) 1999 ix, x 900 – The Western Caucasus, extending over 275,000 ha of the extreme western end of the Caucasus mountains and located 50 km north-east of the Black Sea, is one of the few large mountain areas of Europe that has not experienced significant human impact. Its subalpine and alpine pastures have only been grazed by wild animals, and its extensive tracts of undisturbed mountain forests, extending from the lowlands to the subalpine zone, are unique in Europe. The site has a great diversity of ecosystems, with important endemic plants and wildlife, and is the place of origin and reintroduction of the mountain subspecies of the European bison.
Central Sikhote-Alin Primorsky Krai
45°20′0″N 136°10′0″E / 45.33333°N 136.16667°E / 45.33333; 136.16667 (Sikhote-Alin) 2001 x 766 – The Sikhote-Alin mountain range contains one the richest and most unusual temperate forests of the world. In this mixed zone between taiga and subtropics, southern species such as the tiger and Himalayan bear cohabit with northern species such as the brown bear and lynx. The site stretches from the peaks of Sikhote-Alin to the Sea of Japan and is important for the survival of many endangered species such as the Amur tiger.
Uvs Nuur Basin [n 1] Tuva
50°16′30″N 92°43′11″E / 50.27500°N 92.71972°E / 50.27500; 92.71972 (Uvs Nuur Basin) 2003 ix, x 769 – The Uvs Nuur Basin (1,068,853 ha), is the northernmost of the enclosed basins of Central Asia. It takes its name from Uvs Nuur Lake, a large, shallow and very saline lake, important for migrating birds, waterfowl and seabirds. The site is made up of twelve protected areas representing the major biomes of eastern Eurasia. The steppe ecosystem supports a rich diversity of birds and the desert is home to a number of rare gerbil, jerboas and the marbled polecat. The mountains are an important refuge for the globally endangered snow leopard, mountain sheep (argali) and the Asiatic ibex.
Natural System of Wrangel Island Reserve Chukotka Autonomous Okrug
71°11′20″N 179°42′55″W / 71.18889°N 179.71528°W / 71.18889; -179.71528 (Wrangel Island) 2004 ix, x 1023 – Located well above the Arctic Circle, the site includes the mountainous Wrangel Island (7,608 km2), Herald Island (11 km2) and surrounding waters. Wrangel was not glaciated during the Quaternary Ice Age, resulting in exceptionally high levels of biodiversity for this region. The island boasts the world’s largest population of Pacific walrus and the highest density of ancestral polar bear dens. It is a major feeding ground for the grey whale migrating from Mexico and the northernmost nesting ground for 100 migratory bird species, many endangered. Currently, 417 species and subspecies of vascular plants have been identified on the island, double that of any other Arctic tundra territory of comparable size and more than any other Arctic island. Some species are derivative of widespread continental forms, others are the result of recent hybridization, and 23 are endemic.
Putorana Plateau Krasnoyarsk Krai
69°2′49″N 94°9′29″E / 69.04694°N 94.15806°E / 69.04694; 94.15806 (Putorana Plateau) 2010 vii, ix 1234 – This site coincides with the area of the Putoransky State Nature Reserve, and is located in the central part of the Putorana Plateau in northern Central Siberia. It is situated about 100 km north of the Arctic Circle. The part of the plateau inscribed on the World Heritage List harbours a complete set of subarctic and arctic ecosystems in an isolated mountain range, including pristine taiga, forest tundra, tundra and arctic desert systems, as well as untouched cold-water lake and river systems. A major reindeer migration route crosses the property, which represents an exceptional, large-scale and increasingly rare natural phenomenon.
Lena Pillars Nature Park Sakha Republic
60°40′0″N 127°0′0″E / 60.66667°N 127.00000°E / 60.66667; 127.00000 (Lena Pillars) 2012 viii 1299 – Lena Pillars Nature Park is marked by spectacular rock pillars that reach a height of approximately 100 m along the banks of the Lena River in the central part of the Sakha Republic (Yakutia). They were produced by the region's extreme continental climate with an annual temperature range of almost 100 degrees Celsius (from –60 °C in winter to +40 °C in summer). The pillars form rocky buttresses isolated from each other by deep and steep gullies developed by frost shattering directed along intervening joints. Penetration of water from the surface has facilitated cryogenic processes (freeze-thaw action), which have widened gullies between pillars leading to their isolation. Fluvial processes are also critical to the pillars. The site also contains a wealth of Cambrian fossil remains of numerous species, some of them unique.
Cultural
Historic Centre of Saint Petersburg and Related Groups of Monuments Saint Petersburg
59°57′0″N 30°19′6″E / 59.95000°N 30.31833°E / 59.95000; 30.31833 (Saint Petersburg) 1990 i, ii, iv, vi 540 18th century on The 'Venice of the North', with its numerous canals and more than 400 bridges, is the result of a vast urban project begun in 1703 under Peter the Great. Later known as Leningrad (in the former USSR), the city is closely associated with the October Revolution. Its architectural heritage reconciles the very different Baroque and pure neoclassical styles, as can be seen in the Admiralty, the Winter Palace, the Marble Palace and the Hermitage.
Kizhi Pogost Republic of Karelia
62°4′17″N 35°13′39″E / 62.07139°N 35.22750°E / 62.07139; 35.22750 (Kizhi Pogost) 1990 i, iv, v 544 18th–19th century The pogost of Kizhi (i.e. the Kizhi enclosure) is located on one of the many islands in Lake Onega, in Karelia. Two 18th-century wooden churches, and an octagonal clock tower, also in wood and built in 1862, can be seen there. These unusual constructions, in which carpenters created a bold visionary architecture, perpetuate an ancient model of parish space and are in harmony with the surrounding landscape.
Moscow Kremlin and Red Square Moscow
55°44′45″N 37°37′47″E / 55.74583°N 37.62972°E / 55.74583; 37.62972 (Moscow Kremlin and Red Square) 1990 i, ii, iv, vi 545 14th–20th century Inextricably linked to all the most important historical and political events in Russia since the 13th century, the Kremlin (built between the 14th and 17th centuries by outstanding Russian and foreign architects) was the residence of the Great Prince and also a religious centre. At the foot of its ramparts, on Red Square, St Basil's Basilica is one of the most beautiful Russian Orthodox monuments.
Historic Monuments of Novgorod and Surroundings Novgorod Oblast
58°32′0″N 31°17′0″E / 58.53333°N 31.28333°E / 58.53333; 31.28333 (Veliky Novgorod) 1992 ii, iv, vi 604 11th–17th century Situated on the ancient trade route between Central Asia and northern Europe, Novgorod was Russia's first capital in the 9th century. Surrounded by churches and monasteries, it was a centre for Orthodox spirituality as well as Russian architecture. Its medieval monuments and the 14th-century frescoes of Theophanes the Greek (Andrei Rublev's teacher) illustrate the development of its remarkable architecture and cultural creativity.
Cultural and Historic Ensemble of the Solovetsky Islands Solovetsky Islands,
Arkhangelsk Oblast
65°5′0″N 35°40′0″E / 65.08333°N 35.66667°E / 65.08333; 35.66667 (Solovetsky Islands) 1992 iv 632 16th-17th century The Solovetsky archipelago comprises six islands in the western part of the White Sea, covering 300 km2 . They have been inhabited since the 5th century B.C. and important traces of a human presence from as far back as the 5th millennium B.C. can be found there. The archipelago has been the site of fervent monastic activity since the 15th century, and there are several churches dating from the 16th to the 19th century.
White Monuments of Vladimir and Suzdal Vladimir Oblast
56°9′0″N 40°25′0″E / 56.15000°N 40.41667°E / 56.15000; 40.41667 (Vladimir)
56°25′0″N 40°26′0″E / 56.41667°N 40.43333°E / 56.41667; 40.43333 (Suzdal) 1992 i, ii, iv 633 12th–13th century These two artistic centres in central Russia hold an important place in the country's architectural history. There are a number of magnificent 12th- and 13th-century public and religious buildings, above all the masterpieces of the Collegiate Church of St Demetrios and the Cathedral of the Assumption of the Virgin.
Architectural Ensemble of the Trinity Sergius Lavra Sergiyev Posad, Moscow Oblast
56°18′37.26″N 38°7′52.32″E / 56.3103500°N 38.1312000°E / 56.3103500; 38.1312000 (Trinity Sergius Lavra) 1993 ii, iv 657 15th century on This is a fine example of a working Orthodox monastery, with military features that are typical of the 15th to the 18th century, the period during which it developed. The main church of the Lavra, the Cathedral of the Assumption (echoing the Kremlin Cathedral of the same name), contains the tomb of Boris Godunov. Among the treasures of the Lavra is the famous icon, The Trinity, by Andrei Rublev.
Church of the Ascension, Kolomenskoye Kolomenskoye, Moscow
55°39′20″N 37°40′26″E / 55.65556°N 37.67389°E / 55.65556; 37.67389 (Kolomenskoye) 1994 ii 634 16th century The Church of the Ascension was built in 1532 on the imperial estate of Kolomenskoye, near Moscow, to celebrate the birth of the prince who was to become Tsar Ivan IV ('the Terrible'). One of the earliest examples of a traditional wooden tent-roofed church on a stone and brick substructure, it had a great influence on the development of Russian ecclesiastical architecture.
Historic and Architectural Complex of the Kazan Kremlin Kazan,
Tatarstan
55°47′28″N 49°5′42″E / 55.79111°N 49.09500°E / 55.79111; 49.09500 (Kazan Kremlin) 2000 ii, iii, iv 980 16th century on Built on an ancient site, the Kazan Kremlin dates from the Muslim period of the Golden Horde and the Kazan Khanate. It was conquered by Ivan the Terrible in 1552 and became the Christian See of the Volga Land. The only surviving Tatar fortress in Russia and an important place of pilgrimage, the Kazan Kremlin consists of an outstanding group of historic buildings dating from the 16th to 19th centuries, integrating remains of earlier structures of the 10th to 16th centuries.
Ensemble of the Ferapontov Monastery Vologda Oblast
59°57′0″N 38°34′0″E / 59.95000°N 38.56667°E / 59.95000; 38.56667 (Ferapontov Monastery) 2000 i, iv 982 15th-17th century The Ferapontov Monastery, in the Vologda region in northern Russia, is an exceptionally well-preserved and complete example of a Russian Orthodox monastic complex of the 15th-17th centuries, a period of great significance in the development of the unified Russian state and its culture. The architecture of the monastery is outstanding in its inventiveness and purity. The interior is graced by the magnificent wall paintings of Dionisy, the greatest Russian artist of the end of the 15th century.
Curonian Spit [n 2] Kaliningrad Oblast
55°16′28.488″N 20°57′44.604″E / 55.27458000°N 20.96239000°E / 55.27458000; 20.96239000 (Curonian Spit) 2000 v 994 19th century on Human habitation of this elongated sand dune peninsula, 98 km long and 0.4–4 km wide, dates back to prehistoric times. Throughout this period it has been threatened by the natural forces of wind and waves. Its survival to the present day has been made possible only as a result of ceaseless human efforts to combat the erosion of the Spit, dramatically illustrated by continuing stabilisation and reforestation projects.
Citadel, Ancient City and Fortress Buildings of Derbent Derbent,
Dagestan
42°3′10.7″N 48°17′49.9″E / 42.052972°N 48.297194°E / 42.052972; 48.297194 (Derbent) 2003 iii, iv 1070 6th century on The Citadel, Ancient City and Fortress Buildings of Derbent were part of the northern lines of the Sasanian Persian Empire, which extended east and west of the Caspian Sea. The fortification was built in stone. It consisted of two parallel walls that formed a barrier from the seashore up to the mountain. The town of Derbent was built between these two walls, and has retained part of its medieval fabric. The site continued to be of great strategic importance until the 19th century.
Ensemble of the Novodevichy Convent Moscow
55°43′34″N 37°33′18.3″E / 55.72611°N 37.555083°E / 55.72611; 37.555083 (Novodevichy Convent) 2004 i, iv, vi 1097 16th-17th century The Novodevichy Convent, in south-western Moscow, built in the 16th and 17th centuries in the so-called Moscow Baroque style, was part of a chain of monastic ensembles that were integrated into the defence system of the city. The convent was directly associated with the political, cultural and religious history of Russia, and closely linked to the Moscow Kremlin. It was used by women of the Tsar’s family and the aristocracy. Members of the Tsar’s family and entourage were also buried in its cemetery. The convent provides an example of the highest accomplishments of Russian architecture with rich interiors and an important collection of paintings and artefacts.
Historic Centre of the City of Yaroslavl Yaroslavl,
Yaroslavl Oblast
57°39′10″N 39°52′34″E / 57.65278°N 39.87611°E / 57.65278; 39.87611 (Yaroslavl) 2005 ii, iv 1170 16th century on Situated at the confluence of the Volga and Kotorosl Rivers some 250 km north-east of Moscow, the historic city of Yaroslavl developed into a major commercial centre from the 11th century. It is renowned for its numerous 17th-century churches and is an outstanding example of the urban planning reform Empress Catherine the Great ordered for the whole of Russia in 1763. While keeping some of its significant historic structures, the town was renovated in the neoclassical style on a radial urban master plan. It has also kept elements from the 16th century in the Spassky Monastery, one of the oldest in the Upper Volga region, built on the site of a pagan temple in the late 12th century but reconstructed over time.
Struve Geodetic Arc [n 3] Gogland,
Leningrad Oblast
60°5′7″N 26°57′40″E / 60.08528°N 26.96111°E / 60.08528; 26.96111 (Struve Geodetic Arc) 2005 ii, iii, vi 1187 19th century The Struve Arc is a chain of survey triangulations stretching from Hammerfest in Norway to the Black Sea, through 10 countries and over 2,820 km. These are points of a survey, carried out between 1816 and 1855 by the astronomer Friedrich Georg Wilhelm Struve, which represented the first accurate measuring of a long segment of a meridian. This helped to establish the exact size and shape of the planet and marked an important step in the development of earth sciences and topographic mapping. It is an extraordinary example of scientific collaboration among scientists from different countries, and of collaboration between monarchs for a scientific cause. The original arc consisted of 258 main triangles with 265 main station points. The listed site includes 34 of the original station points, with different markings, i.e. a drilled hole in rock, iron cross, cairns, or built obelisks.
Bolgar Historical and Archaeological Complex Tatarstan
54°58′44″N 49°03′23″E / 54.97889°N 49.05639°E / 54.97889; 49.05639 (Bolghar) 2014 ii, iv 981 13th century on This property lies on the shores of the Volga River, south of its confluence with the River Kama, and south of the capital of Tatarstan, Kazan. It contains evidence of the medieval city of Bolgar, an early settlement of the civilization of Volga-Bolgars, which existed between the 7th and the 15th centuries, and was the first capital of the Golden Horde in the 13th century. Bolgar represents the historical cultural exchanges and transformations of Eurasia over several centuries that played a pivotal role in the formation of civilizations, customs and cultural traditions. The property provides remarkable evidence of historic continuity and cultural diversity. It is a symbolic reminder of the acceptance of Islam by the Volga-Bolgars in 922 AD and remains a sacred, pilgrimage destination to the Tatar Muslims.
Assumption Cathedral and Monastery of the town-island of Sviyazhsk Tatarstan
55°46′13″N 48°39′10″E / 55.77028°N 48.65278°E / 55.77028; 48.65278 (Assumption Cathedral of Sviyazhsk) 2017 ii, iv 1525 16th century on The Assumption Cathedral is located in the town-island of Sviyazhsk and is part of the monastery of the same name. Situated at the confluence of the Volga, the Sviyaga and the Shchuka rivers, at the crossroads of the Silk and Volga routes, Sviyazhsk was founded by Ivan the Terrible in 1551. It was from this outpost that he initiated the conquest of the Kazan Khanate. The cathedral’s frescoes are among the rarest examples of Eastern Orthodox mural paintings.
Jump up ^ shared with Mongolia
Jump up ^ shared with Lithuania
Jump up ^ shared with Belarus, Estonia, Finland, Latvia, Lithuania, Moldova, Norway, Sweden, Ukraine
Tentative list[edit]
Daursky Reserve
Commander Islands
Magadansky Reserve
Krasnoyarsk Stolby
Vasyugan Swamp
Ilmensky Mountains
Bikin River
Valaam
Bashkir Ural
Kenozero Lake
Jeyrakh–Assa Reservation
Cathedral of Christ the Saviour, Sheremetev's Shelter, Izmaylovo Estate
Sviyazhsk
Irkutsk
Rostov Kremlin
Yeniseysk
Uglich Kremlin,
Church of Dimitri
Pskov Kremlin
Sikachi-Alyan
Astrakhan Kremlin
Tanais
Location of UNESCO Tentative list World Heritage Sites within Russia
Legend: World Cultural Heritage Site; World Natural Heritage Site; World Cultural & Natural Heritage Site
There are currently twenty-four properties in the tentative list.[2]
The sites are sorted in the chronological order of their addition to the tentative list.
Name
Image
Location
Period
Year
submit.
Criteria
Ref.
num.
Natural
Daurian Steppes (Daursky State Biosphere Reserve) Zabaykalsky Krai
50°03′37″N 115°39′07″E / 50.06028°N 115.65194°E / 50.06028; 115.65194 (Daursky Reserve) – 2005 x 1996
The Commander Islands (Komandorsky Nature Reserve) Kamchatka Krai
55°N 167°E / 55°N 167°E / 55; 167 (Commander Islands) – 2005 vii, viii, ix, x 1997
Magadan Nature Reserve (ru) Magadan Oblast
61°25′N 141°0′E / 61.417°N 141.000°E / 61.417; 141.000 (Magadansky Reserve) – 2005 vii, viii, ix, x 1998
Krasnoyarsk Stolby (ru) Krasnoyarsk Krai
55°57′40″N 92°48′25″E / 55.96111°N 92.80694°E / 55.96111; 92.80694 (Krasnoyarsk Stolby) – 2007 vii, viii, ix, x 5113
The Great Vasyugan Mire Tomsk Oblast,
Novosibirsk Oblast,
Omsk Oblast
56°58′51″N 79°41′24″E / 56.98083°N 79.69000°E / 56.98083; 79.69000 (Vasyugan Swamp) – 2007 vii, viii, ix, x 5114
The Ilmensky Mountains (ru) Chelyabinsk Oblast
55°09′N 60°15′E / 55.150°N 60.250°E / 55.150; 60.250 (Ilmensky Mountains) – 2008 vi, vii, viii 5388
Bikin River Valley (Extension of the "Central Sikhote-Alin") Primorsky Krai
46°39′N 135°27′E / 46.650°N 135.450°E / 46.650; 135.450 (Bikin River) – 2010 x 5571
Mixed
The Valamo archipelago Karelia
61°23′20″N 30°56′49″E / 61.38889°N 30.94694°E / 61.38889; 30.94694 (Valaam) 19th century on 1996 (mixed) 572
Bashkir Ural (ru) Bashkortostan
53°08′30″N 57°00′30″E / 53.14167°N 57.00833°E / 53.14167; 57.00833 (Bashkir Ural) – 2012 i, iii, v, vi, viii, x 5666
Testament of Kenozero Lake Arkhangelsk Oblast
62°N 38°E / 62°N 38°E / 62; 38 (Kenozero Lake) – 2014 iii, iv, v, vi, vii 5922
Cultural
Historical and Cultural Jeyrakh–Assa Reservation Ingushetia
42°49′N 44°40′E / 42.817°N 44.667°E / 42.817; 44.667 (Jeyrakh–Assa Reservation) 5th century B. C. – 17th century 1996 (cultural) 569
The architectural and historical complex "Shelter of count N. P. Cheremetev" (ru) Moscow
55°45′58″N 37°37′58″E / 55.76611°N 37.63278°E / 55.76611; 37.63278 (Sheremetev's Shelter) 17th–18th century 1996 i, iii, vi 571
The ensemble of former city building of Sviyazhsk Tatarstan
55°46′N 48°39′E / 55.767°N 48.650°E / 55.767; 48.650 (Sviyazhsk) 16th century on 1998 (cultural) 1111
Historic Centre of Irkutsk Irkutsk Oblast
52°18′N 104°17′E / 52.300°N 104.283°E / 52.300; 104.283 (Irkutsk) 19th century on 1998 (cultural) 1166
Cathedral of Christ the Saviour Moscow
55°45′N 37°37′E / 55.750°N 37.617°E / 55.750; 37.617 (Cathedral of Christ the Saviour) 1994–1997 1998 ii, vi 1168
Rostov Kremlin (ru) Yaroslavl Oblast
57°11′N 39°25′E / 57.183°N 39.417°E / 57.183; 39.417 (Rostov Kremlin) 17th century 1998 ii, iii, iv, vi 1185
Historic Centre of Yeniseysk Krasnoyarsk Krai
58°37′23″N 92°09′16″E / 58.62306°N 92.15444°E / 58.62306; 92.15444 (Yeniseysk) 17th century on 2000 ii, iii, iv 1460
Church of Prince Dimitri "On Blood" (ru) [t 1] Uglich, Yaroslavl Oblast
57°31′43″N 38°19′01″E / 57.52861°N 38.31694°E / 57.52861; 38.31694 (Church of Dimitri in Uglich Kremlin) 17th century 2001 i, ii, iv 1518
Great Pskov [t 2] Pskov Oblast
57°49′34″N 28°19′36″E / 57.82611°N 28.32667°E / 57.82611; 28.32667 (Pskov Kremlin) 10th century on 2002 i, ii, iv 1638
Petroglyphs of Sikachi-Alyan (ru) Khabarovsk Krai
48°45′N 135°39′E / 48.750°N 135.650°E / 48.750; 135.650 (Sikachi-Alyan) 13,000 B. C. – 3,000 B. C. 2003 (cultural) 1787
Ensemble of Astrakhan Kremlin [t 3] Astrakhan, Astrakhan Oblast
46°20′50″N 48°02′00″E / 46.34722°N 48.03333°E / 46.34722; 48.03333 (Astrakhan Kremlin) 16th–19th century 2008 ii, iii, iv 5368
The archeological site of Tanais Rostov Oblast
47°16′14″N 39°20′39″E / 47.27056°N 39.34417°E / 47.27056; 39.34417 (Tanais) 3rd century B. C. – 5th century A. D. 2009 ii, iii, v 5422
Russian Kremlins:
Ensemble of the Astrakhan Kremlin,[t 3]
Ensemble of the Uglich Kremlin (ru),[t 1]
Ensemble of the Pskov Kremlin[t 2] Astrakhan Oblast 46°20′50″N 48°02′00″E / 46.34722°N 48.03333°E / 46.34722; 48.03333 (Astrakhan Kremlin),
Yaroslavl Oblast 57°31′43″N 38°19′01″E / 57.52861°N 38.31694°E / 57.52861; 38.31694 (Church of Dimitri in Uglich Kremlin),
Pskov Oblast 57°49′34″N 28°19′36″E / 57.82611°N 28.32667°E / 57.82611; 28.32667 (Pskov Kremlin) 10th century on 2010 ii, iii, iv 5517
Architectural and Park Ensemble "Tsar's Country Estate Izmaylovo" Moscow
55°47′31.88″N 37°45′40.31″E / 55.7921889°N 37.7611972°E / 55.7921889; 37.7611972 (Izmaylovo Estate) 17th–19th century 2011 ii, iv 5622
^ Jump up to: a b Church of Prince Dimitri "On Blood" is submitted both separately and as a part of Uglich Kremlin
^ Jump up to: a b Pskov Kremlin is submitted both as a part of Great Pskov and jointly with Russian Kremlins
^ Jump up to: a b Astrakhan Kremlin is submitted both separately and jointly with Russian Kremlins
See also[edit]
List of World Heritage Sites in Europe
List of World Heritage Sites in Asia
References[edit]
^ Jump up to: a b "Russian Federation". UNESCO World Heritage Centre. Retrieved March 27, 2018.
^ Jump up to: a b "Tentative list of the Russian Federation". UNESCO World Heritage Centre. Retrieved 2014-06-25.
[hide]
v
t
e
World Heritage Sites in Russia by federal district

Central
Church of the Ascension in Kolomenskoye
Moscow Kremlin and Red Square
Novodevichy Convent
Trinity Sergius Lavra
White Monuments of Vladimir and Suzdal
Historic Centre of Yaroslavl

Southern
Western Caucasus
Northwestern
Curonian Spit1
Ferapontov Monastery
Kizhi Pogost
Virgin Komi Forests
Historic Monuments of Novgorod and Surroundings
Historic Centre of Saint Petersburg and Surroundings
Solovetsky Islands
Struve Geodetic Arc2
Far Eastern
Lena Pillars
Volcanoes of Kamchatka
Central Sikhote-Alin
Wrangel Island
Siberian
Golden Mountains of Altai
Lake Baikal
Landscapes of Dauria3
Putorana Plateau
Uvs Nuur Basin3
Volga
Assumption Cathedral of Sviyazhsk
Bolghar
Kazan Kremlin
North Caucasian
Citadel, Ancient City and Fortress Buildings of Derbent
1 Shared with Lithuania
2 Shared with nine other countries
3 Shared with Mongolia
[show]
v
t
e
Lists of World Heritage Sites in Asia
Northern and Central Asia
Kazakhstan
Kyrgyzstan
Tajikistan
Turkmenistan
Uzbekistan
Western Asia
(including Arab States)
Armenia
Azerbaijan
Bahrain
Georgia
Iran
Iraq
Israel
Jordan
Lebanon
Oman
Palestine
Saudi Arabia
Syria
Turkey
United Arab Emirates
Yemen
Eastern Asia
China
Japan
Mongolia
North Korea
South Korea
Southern Asia
Afghanistan
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India
Nepal
Pakistan
Sri Lanka
Southeast Asia
Cambodia
Indonesia
Laos
Malaysia
Myanmar
Philippines
Singapore
Thailand
Vietnam
[show]
v
t
e
Lists of World Heritage Sites in Europe
Northern Europe
Denmark
Estonia
Finland
Iceland
Latvia
Lithuania
Norway
Sweden
Western Europe
Austria
Belgium
France
Germany
Ireland
Luxembourg
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Armenia
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Georgia
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Romania
Russia
Slovakia
Ukraine
Southern Europe
Albania
Andorra
Bosnia and Herzegovina
Croatia
Cyprus
Greece
Gibraltar (UK)
Holy See
Italy
Macedonia
Malta
Montenegro
Portugal
San Marino
Serbia
Slovenia
Spain
Turkey
[show]
v
t
e
Lists of World Heritage Sites
Africa
Americas
North America
Central America
Caribbean
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Asia
Eastern
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Southern
Western
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World Heritage in Danger
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By country
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Endoplasmic reticulum
From Wikipedia, the free encyclopedia
Jump to: navigation, search
Cell biology
The animal cell
Components of a typical animal cell:
Nucleolus
Nucleus
Ribosome (little dots)
Vesicle
Rough endoplasmic reticulum
Golgi apparatus (or "Golgi body")
Cytoskeleton
Smooth endoplasmic reticulum
Mitochondrion
Vacuole
Cytosol (fluid that contains organelles, comprising the cytoplasm)
Lysosome
Centrosome
Cell membrane
Micrograph of rough endoplasmic reticulum network around the nucleus (shown in lower right-hand side of the picture). Dark small circles in the network are mitochondria.
The endoplasmic reticulum (ER) is a type of organelle found in eukaryotic cells that forms an interconnected network of flattened, membrane-enclosed sacs or tube-like structures known as cisternae. The membranes of the ER are continuous with the outer nuclear membrane. The endoplasmic reticulum occurs in most types of eukaryotic cells, but is absent from red blood cells and spermatozoa. There are two types of endoplasmic reticulum: rough and smooth. The outer (cytosolic) face of the rough endoplasmic reticulum is studded with ribosomes that are the sites of protein synthesis. The rough endoplasmic reticulum is especially prominent in cells such as hepatocytes. The smooth endoplasmic reticulum lacks ribosomes and functions in lipid manufacture and metabolism, the production of steroid hormones, and detoxification.[1] The smooth ER is especially abundant in mammalian liver and gonad cells. The lacy membranes of the endoplasmic reticulum were first seen in 1945 using electron microscopy.
Contents
[hide]
1 History
2 Structure
2.1 Rough endoplasmic reticulum
2.2 Smooth endoplasmic reticulum
2.2.1 Sarcoplasmic reticulum
3 Functions
3.1 Protein transport
4 Clinical significance
5 References
6 External links
History[edit]
The lacy membranes of the endoplasmic reticulum were first seen in 1945 by Keith R. Porter, Albert Claude, Brody Meskers and Ernest F. Fullam, using electron microscopy.[2] The word reticulum, which means "network", was applied to describe this fabric of membranes.
Structure[edit]
1 Nucleus 2 Nuclear pore 3 Rough endoplasmic reticulum (RER) 4 Smooth endoplasmic reticulum (SER) 5 Ribosome on the rough ER 6 Proteins that are transported 7 Transport vesicle 8 Golgi apparatus 9 Cis face of the Golgi apparatus 10 Trans face of the Golgi apparatus 11 Cisternae of the Golgi apparatus
3D rendering of endoplasmic reticulum
The general structure of the endoplasmic reticulum is a network of membranes called cisternae. These sac-like structures are held together by the cytoskeleton. The phospholipid membrane encloses the cisternal space (or lumen), which is continuous with the perinuclear space but separate from the cytosol. The functions of the endoplasmic reticulum can be summarized as the synthesis and export of proteins and membrane lipids, but varies between ER and cell type and cell function. The quantity of both rough and smooth endoplasmic reticulum in a cell can slowly interchange from one type to the other, depending on the changing metabolic activities of the cell. Transformation can include embedding of new proteins in membrane as well as structural changes. Changes in protein content may occur without noticeable structural changes.[3][4][citation needed]
Rough endoplasmic reticulum[edit]
An animation showing how a protein destined for the secretory pathway is synthesized into the rough endoplasmic reticulum (which appears at upper right in animation when approximately half of animation is done). The total animation time is about 2 minutes.
The surface of the rough endoplasmic reticulum (often abbreviated RER or Rough ER) (also called granular endoplasmic reticulum) is studded with protein-manufacturing ribosomes giving it a "rough" appearance (hence its name).[5] The binding site of the ribosome on the rough endoplasmic reticulum is the translocon.[6] However, the ribosomes are not a stable part of this organelle's structure as they are constantly being bound and released from the membrane. A ribosome only binds to the RER once a specific protein-nucleic acid complex forms in the cytosol. This special complex forms when a free ribosome begins translating the mRNA of a protein destined for the secretory pathway.[7] The first 5–30 amino acids polymerized encode a signal peptide, a molecular message that is recognized and bound by a signal recognition particle (SRP). Translation pauses and the ribosome complex binds to the RER translocon where translation continues with the nascent (new) protein forming into the RER lumen and/or membrane. The protein is processed in the ER lumen by an enzyme (a signal peptidase), which removes the signal peptide. Ribosomes at this point may be released back into the cytosol; however, non-translating ribosomes are also known to stay associated with translocons.[8]
The membrane of the rough endoplasmic reticulum forms large double membrane sheets that are located near, and continuous with, the outer layer of the nuclear envelope.[9] The double membrane sheets are stacked and connected through several right or left-handed helical ramps, the so-called Terasaki ramps, giving rise to a structure resembling a multi-storey car park.[10][11] Although there is no continuous membrane between the endoplasmic reticulum and the Golgi apparatus, membrane-bound transport vesicles shuttle proteins between these two compartments.[12] Vesicles are surrounded by coating proteins called COPI and COPII. COPII targets vesicles to the Golgi apparatus and COPI marks them to be brought back to the rough endoplasmic reticulum. The rough endoplasmic reticulum works in concert with the Golgi complex to target new proteins to their proper destinations. A second method of transport out of the endoplasmic reticulum involves areas called membrane contact sites, where the membranes of the endoplasmic reticulum and other organelles are held closely together, allowing the transfer of lipids and other small molecules.[13][14]
The rough endoplasmic reticulum is key in multiple functions:
Manufacture of lysosomal enzymes with a mannose-6-phosphate marker added in the cis-Golgi network.[citation needed]
Manufacture of secreted proteins, either secreted constitutively with no tag or secreted in a regulatory manner involving clathrin and paired basic amino acids in the signal peptide.
Integral membrane proteins that stay embedded in the membrane as vesicles exit and bind to new membranes. Rab proteins are key in targeting the membrane; SNAP and SNARE proteins are key in the fusion event.
Initial glycosylation as assembly continues. This is N-linked (O-linking occurs in the Golgi).
N-linked glycosylation: If the protein is properly folded, Oligosaccharyltransferase recognizes the AA sequence NXS or NXT (with the S/T residue phosphorylated) and adds a 14-sugar backbone (2-N-acetylglucosamine, 9-branching mannose, and 3-glucose at the end) to the side-chain nitrogen of Asn.
Smooth endoplasmic reticulum[edit]
Electromicrograph showing smooth ER (arrow) in mouse tissue, at 110,510 x magnification.
In most cells the smooth endoplasmic reticulum (abbreviated SER) is scarce. Instead there are areas where the ER is partly smooth and partly rough, this area is called the transitional ER. The transitional ER gets its name because it contains ER exit sites. These are areas where the transport vesicles that contain lipids and proteins made in the ER, detach from the ER and start moving to the Golgi apparatus. Specialized cells can have a lot of smooth endoplasmic reticulum and in these cells the smooth ER has many functions.[15] It synthesizes lipids, phospholipids, and steroids. Cells which secrete these products, such as those in the testes, ovaries, and sebaceous glands have an abundance of smooth endoplasmic reticulum.[16] It also carries out the metabolism of carbohydrates, detoxification of natural metabolism products and of alcohol and drugs, attachment of receptors on cell membrane proteins, and steroid metabolism.[17] In muscle cells, it regulates calcium ion concentration. Smooth endoplasmic reticulum is found in a variety of cell types (both animal and plant), and it serves different functions in each. The smooth endoplasmic reticulum also contains the enzyme glucose-6-phosphatase, which converts glucose-6-phosphate to glucose, a step in gluconeogenesis. It is connected to the nuclear envelope and consists of tubules that are located near the cell periphery. These tubes sometimes branch forming a network that is reticular in appearance.[9] In some cells, there are dilated areas like the sacs of rough endoplasmic reticulum. The network of smooth endoplasmic reticulum allows for an increased surface area to be devoted to the action or storage of key enzymes and the products of these enzymes.
Sarcoplasmic reticulum[edit]
Main article: Sarcoplasmic reticulum
See also: Calcium-induced calcium release
Skeletal muscle fiber, with sarcoplasmic reticulum colored in blue.
The sarcoplasmic reticulum (SR), from the Greek σάρξ sarx ("flesh"), is smooth ER found in myocytes. The only structural difference between this organelle and the smooth endoplasmic reticulum is the medley of proteins they have, both bound to their membranes and drifting within the confines of their lumens. This fundamental difference is indicative of their functions: The endoplasmic reticulum synthesizes molecules, while the sarcoplasmic reticulum stores calcium ions and pumps them out into the sarcoplasm when the muscle fiber is stimulated.[18][19] After their release from the sarcoplasmic reticulum, calcium ions interact with contractile proteins that utilize ATP to shorten the muscle fiber. The sarcoplasmic reticulum plays a major role in excitation-contraction coupling.[20]
Functions[edit]
The endoplasmic reticulum serves many general functions, including the folding of protein molecules in sacs called cisternae and the transport of synthesized proteins in vesicles to the Golgi apparatus. Correct folding of newly made proteins is made possible by several endoplasmic reticulum chaperone proteins, including protein disulfide isomerase (PDI), ERp29, the Hsp70 family member BiP/Grp78, calnexin, calreticulin, and the peptidylpropyl isomerase family. Only properly folded proteins are transported from the rough ER to the Golgi apparatus – unfolded proteins cause an unfolded protein response as a stress response in the ER. Disturbances in redox regulation, calcium regulation, glucose deprivation, and viral infection[21] or the over-expression of proteins[22] can lead to endoplasmic reticulum stress response (ER stress), a state in which the folding of proteins slows, leading to an increase in unfolded proteins. This stress is emerging as a potential cause of damage in hypoxia/ischemia, insulin resistance, and other disorders.[23]
Protein transport[edit]
Secretory proteins, mostly glycoproteins, are moved across the endoplasmic reticulum membrane. Proteins that are transported by the endoplasmic reticulum throughout the cell are marked with an address tag called a signal sequence. The N-terminus (one end) of a polypeptide chain (i.e., a protein) contains a few amino acids that work as an address tag, which are removed when the polypeptide reaches its destination. Nascent peptides reach the ER via the translocon, a membrane-embedded multiprotein complex. Proteins that are destined for places outside the endoplasmic reticulum are packed into transport vesicles and moved along the cytoskeleton toward their destination. In human fibroblasts, the ER is always co-distributed with microtubules and the depolymerisation of the latter cause its co-aggregation with mitochondria, which are also associated with the ER.[24]
The endoplasmic reticulum is also part of a protein sorting pathway. It is, in essence, the transportation system of the eukaryotic cell. The majority of its resident proteins are retained within it through a retention motif. This motif is composed of four amino acids at the end of the protein sequence. The most common retention sequences are KDEL for lumen located proteins and KKXX for transmembrane protein.[25] However, variations of KDEL and KKXX do occur, and other sequences can also give rise to endoplasmic reticulum retention. It is not known whether such variation can lead to sub-ER localizations. There are three KDEL (1, 2 and 3) receptors in mammalian cells, and they have a very high degree of sequence identity. The functional differences between these receptors remain to be established.[26]
Clinical significance[edit]
Abnormalities in XBP1 lead to a heightened endoplasmic reticulum stress response and subsequently causes a higher susceptibility for inflammatory processes that may even contribute to Alzheimer's disease.[27] In the colon, XBP1 anomalies have been linked to the inflammatory bowel diseases including Crohn's disease.[28]
The unfolded protein response (UPR) is a cellular stress response related to the endoplasmic reticulum.[29] The UPR is activated in response to an accumulation of unfolded or misfolded proteins in the lumen of the endoplasmic reticulum. The UPR functions to restore normal function of the cell by halting protein translation, degrading misfolded proteins, and activating the signaling pathways that lead to increasing the production of molecular chaperones involved in protein folding. Sustained overactivation of the UPR has been implicated in prion diseases as well as several other neurodegenerative diseases and the inhibition of the UPR could become a treatment for those diseases.[30]
References[edit]
Jump up ^ "Endoplasmic Reticulum (Rough and Smooth)". Retrieved 21 November 2015.
Jump up ^ Porter KR, Claude A, Fullam EF (March 1945). "A study of tissue culture cells by electron microscopy". J Exp Med. 81 (3): 233–246. doi:10.1084/jem.81.3.233. PMC 2135493  . PMID 19871454.
Jump up ^ Alberts, Bruce; Johnson, Alexander; Lewis, Julian; Raff, Martin; Roberts, Keith; Walter, Peter (2002). Molecular biology of the cell (4th ed.). New York: Garland Science. ISBN 0-8153-3218-1.
Jump up ^ Cooper, Geoffrey M. (2000). The cell : a molecular approach (2nd ed.). Washington (DC): ASM Press. ISBN 0-87893-106-6.
Jump up ^ "reticulum". The Free Dictionary.
Jump up ^ Görlich D, Prehn S, Hartmann E, Kalies KU, Rapoport TA (Oct 1992). "A mammalian homolog of SEC61p and SECYp is associated with ribosomes and nascent polypeptides during translocation". Cell. 71 (3): 489–503. doi:10.1016/0092-8674(92)90517-G. PMID 1423609.
Jump up ^ Lodish, Harvey; et al. (2003). Molecular Cell Biology (5th ed.). W. H. Freeman. pp. 659–666. ISBN 0-7167-4366-3.
Jump up ^ Seiser, R. M. (2000). "The Fate of Membrane-bound Ribosomes Following the Termination of Protein Synthesis". Journal of Biological Chemistry. 275 (43): 33820–33827. doi:10.1074/jbc.M004462200. ISSN 0021-9258. PMID 10931837.
^ Jump up to: a b Shibata, Yoko; Voeltz, Gia K.; Rapoport, Tom A. (2006). "Rough Sheets and Smooth Tubules". Cell. 126 (3): 435–439. doi:10.1016/j.cell.2006.07.019. ISSN 0092-8674.
Jump up ^ Terasaki, Mark; Shemesh, Tom; Kasthuri, Narayanan; Klemm, Robin W.; Schalek, Richard; Hayworth, Kenneth J.; Hand, Arthur R.; Yankova, Maya; Huber, Greg; Lichtman, Jeff W.; Rapoport, Tom A.; Kozlov, Michael M. (2013). "Stacked Endoplasmic Reticulum Sheets Are Connected by Helicoidal Membrane Motifs". Cell. 154 (2): 285–296. doi:10.1016/j.cell.2013.06.031. PMC 3767119  . PMID 23870120.
Jump up ^ Guven, J; Huber, G; Valencia, DM (31 October 2014). "Terasaki spiral ramps in the rough endoplasmic reticulum". Physical review letters. 113 (18): 188101. doi:10.1103/PhysRevLett.113.188101. PMID 25396396.
Jump up ^ Endoplasmic reticulum. (n.d.). McGraw-Hill Encyclopedia of Science and Technology. Retrieved September 13, 2006, from Answers.com Web site: http://www.answers.com/topic/endoplasmic-reticulum
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Lipid and protein composition of Endoplasmic reticulum in OPM database
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