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Plastoglobuli (singular plastoglobulus, sometimes spelled plastoglobule(s)), are spherical bubbles of lipids and proteins about 45–60 nanometers across. They are surrounded by a lipid monolayer. Plastoglobuli are found in all chloroplasts, but become more common when the chloroplast is under oxidative stress, or when i...
Plastoglobuli were once thought to be free-floating in the stroma, but it is now thought that they are permanently attached either to a thylakoid or to another plastoglobulus attached to a thylakoid, a configuration that allows a plastoglobulus to exchange its contents with the thylakoid network. In normal green chloro...
The chloroplasts of some hornworts and algae contain structures called pyrenoids. They are not found in higher plants. Pyrenoids are roughly spherical and highly refractive bodies which are a site of starch accumulation in plants that contain them. They consist of a matrix opaque to electrons, surrounded by two hemisph...
In the helical thylakoid model, grana consist of a stack of flattened circular granal thylakoids that resemble pancakes. Each granum can contain anywhere from two to a hundred thylakoids, though grana with 10–20 thylakoids are most common. Wrapped around the grana are helicoid stromal thylakoids, also known as frets or...
Embedded in the thylakoid membranes are important protein complexes which carry out the light reactions of photosynthesis. Photosystem II and photosystem I contain light-harvesting complexes with chlorophyll and carotenoids that absorb light energy and use it to energize electrons. Molecules in the thylakoid membrane u...
There are two types of thylakoids—granal thylakoids, which are arranged in grana, and stromal thylakoids, which are in contact with the stroma. Granal thylakoids are pancake-shaped circular disks about 300–600 nanometers in diameter. Stromal thylakoids are helicoid sheets that spiral around grana. The flat tops and bot...
In addition to chlorophylls, another group of yellow–orange pigments called carotenoids are also found in the photosystems. There are about thirty photosynthetic carotenoids. They help transfer and dissipate excess energy, and their bright colors sometimes override the chlorophyll green, like during the fall, when the ...
Phycobilins are a third group of pigments found in cyanobacteria, and glaucophyte, red algal, and cryptophyte chloroplasts. Phycobilins come in all colors, though phycoerytherin is one of the pigments that makes many red algae red. Phycobilins often organize into relatively large protein complexes about 40 nanometers a...
To fix carbon dioxide into sugar molecules in the process of photosynthesis, chloroplasts use an enzyme called rubisco. Rubisco has a problem—it has trouble distinguishing between carbon dioxide and oxygen, so at high oxygen concentrations, rubisco starts accidentally adding oxygen to sugar precursors. This has the end...
As a result, chloroplasts in C4 mesophyll cells and bundle sheath cells are specialized for each stage of photosynthesis. In mesophyll cells, chloroplasts are specialized for the light reactions, so they lack rubisco, and have normal grana and thylakoids, which they use to make ATP and NADPH, as well as oxygen. They st...
Not all cells in a multicellular plant contain chloroplasts. All green parts of a plant contain chloroplasts—the chloroplasts, or more specifically, the chlorophyll in them are what make the photosynthetic parts of a plant green. The plant cells which contain chloroplasts are usually parenchyma cells, though chloroplas...
In some plants such as cacti, chloroplasts are found in the stems, though in most plants, chloroplasts are concentrated in the leaves. One square millimeter of leaf tissue can contain half a million chloroplasts. Within a leaf, chloroplasts are mainly found in the mesophyll layers of a leaf, and the guard cells of stom...
The chloroplasts of plant and algal cells can orient themselves to best suit the available light. In low-light conditions, they will spread out in a sheet—maximizing the surface area to absorb light. Under intense light, they will seek shelter by aligning in vertical columns along the plant cell's cell wall or turning ...
Plants have two main immune responses—the hypersensitive response, in which infected cells seal themselves off and undergo programmed cell death, and systemic acquired resistance, where infected cells release signals warning the rest of the plant of a pathogen's presence. Chloroplasts stimulate both responses by purpos...
Chloroplasts can serve as cellular sensors. After detecting stress in a cell, which might be due to a pathogen, chloroplasts begin producing molecules like salicylic acid, jasmonic acid, nitric oxide and reactive oxygen species which can serve as defense-signals. As cellular signals, reactive oxygen species are unstabl...
One of the main functions of the chloroplast is its role in photosynthesis, the process by which light is transformed into chemical energy, to subsequently produce food in the form of sugars. Water (H2O) and carbon dioxide (CO2) are used in photosynthesis, and sugar and oxygen (O2) is made, using light energy. Photosyn...
Like mitochondria, chloroplasts use the potential energy stored in an H+, or hydrogen ion gradient to generate ATP energy. The two photosystems capture light energy to energize electrons taken from water, and release them down an electron transport chain. The molecules between the photosystems harness the electrons' en...
While photosystem II photolyzes water to obtain and energize new electrons, photosystem I simply reenergizes depleted electrons at the end of an electron transport chain. Normally, the reenergized electrons are taken by NADP+, though sometimes they can flow back down more H+-pumping electron transport chains to transpo...
The Calvin cycle starts by using the enzyme Rubisco to fix CO2 into five-carbon Ribulose bisphosphate (RuBP) molecules. The result is unstable six-carbon molecules that immediately break down into three-carbon molecules called 3-phosphoglyceric acid, or 3-PGA. The ATP and NADPH made in the light reactions is used to co...
Alternatively, glucose monomers in the chloroplast can be linked together to make starch, which accumulates into the starch grains found in the chloroplast. Under conditions such as high atmospheric CO2 concentrations, these starch grains may grow very large, distorting the grana and thylakoids. The starch granules dis...
Photorespiration can occur when the oxygen concentration is too high. Rubisco cannot distinguish between oxygen and carbon dioxide very well, so it can accidentally add O2 instead of CO2 to RuBP. This process reduces the efficiency of photosynthesis—it consumes ATP and oxygen, releases CO2, and produces no sugar. It ca...
Chloroplasts alone make almost all of a plant cell's amino acids in their stroma except the sulfur-containing ones like cysteine and methionine. Cysteine is made in the chloroplast (the proplastid too) but it is also synthesized in the cytosol and mitochondria, probably because it has trouble crossing membranes to get ...
Chloroplasts are a special type of a plant cell organelle called a plastid, though the two terms are sometimes used interchangeably. There are many other types of plastids, which carry out various functions. All chloroplasts in a plant are descended from undifferentiated proplastids found in the zygote, or fertilized e...
If angiosperm shoots are not exposed to the required light for chloroplast formation, proplastids may develop into an etioplast stage before becoming chloroplasts. An etioplast is a plastid that lacks chlorophyll, and has inner membrane invaginations that form a lattice of tubes in their stroma, called a prolamellar bo...
Plastid differentiation is not permanent, in fact many interconversions are possible. Chloroplasts may be converted to chromoplasts, which are pigment-filled plastids responsible for the bright colors seen in flowers and ripe fruit. Starch storing amyloplasts can also be converted to chromoplasts, and it is possible fo...
The division process starts when the proteins FtsZ1 and FtsZ2 assemble into filaments, and with the help of a protein ARC6, form a structure called a Z-ring within the chloroplast's stroma. The Min system manages the placement of the Z-ring, ensuring that the chloroplast is cleaved more or less evenly. The protein MinD...
Next, the two plastid-dividing rings, or PD rings form. The inner plastid-dividing ring is located in the inner side of the chloroplast's inner membrane, and is formed first. The outer plastid-dividing ring is found wrapped around the outer chloroplast membrane. It consists of filaments about 5 nanometers across, arran...
Light has been shown to be a requirement for chloroplast division. Chloroplasts can grow and progress through some of the constriction stages under poor quality green light, but are slow to complete division—they require exposure to bright white light to complete division. Spinach leaves grown under green light have be...
Recently, chloroplasts have caught attention by developers of genetically modified crops. Since, in most flowering plants, chloroplasts are not inherited from the male parent, transgenes in these plastids cannot be disseminated by pollen. This makes plastid transformation a valuable tool for the creation and cultivatio...
A prime number (or a prime) is a natural number greater than 1 that has no positive divisors other than 1 and itself. A natural number greater than 1 that is not a prime number is called a composite number. For example, 5 is prime because 1 and 5 are its only positive integer factors, whereas 6 is composite because it ...
The property of being prime (or not) is called primality. A simple but slow method of verifying the primality of a given number n is known as trial division. It consists of testing whether n is a multiple of any integer between 2 and . Algorithms much more efficient than trial division have been devised to test the pri...
There are infinitely many primes, as demonstrated by Euclid around 300 BC. There is no known simple formula that separates prime numbers from composite numbers. However, the distribution of primes, that is to say, the statistical behaviour of primes in the large, can be modelled. The first result in that direction is t...
Many questions regarding prime numbers remain open, such as Goldbach's conjecture (that every even integer greater than 2 can be expressed as the sum of two primes), and the twin prime conjecture (that there are infinitely many pairs of primes whose difference is 2). Such questions spurred the development of various br...
Hence, 6 is not prime. The image at the right illustrates that 12 is not prime: 12 = 3 · 4. No even number greater than 2 is prime because by definition, any such number n has at least three distinct divisors, namely 1, 2, and n. This implies that n is not prime. Accordingly, the term odd prime refers to any prime numb...
Most early Greeks did not even consider 1 to be a number, so they could not consider it to be a prime. By the Middle Ages and Renaissance many mathematicians included 1 as the first prime number. In the mid-18th century Christian Goldbach listed 1 as the first prime in his famous correspondence with Leonhard Euler -- w...
A large body of mathematical work would still be valid when calling 1 a prime, but Euclid's fundamental theorem of arithmetic (mentioned above) would not hold as stated. For example, the number 15 can be factored as 3 · 5 and 1 · 3 · 5; if 1 were admitted as a prime, these two presentations would be considered differen...
There are hints in the surviving records of the ancient Egyptians that they had some knowledge of prime numbers: the Egyptian fraction expansions in the Rhind papyrus, for instance, have quite different forms for primes and for composites. However, the earliest surviving records of the explicit study of prime numbers c...
After the Greeks, little happened with the study of prime numbers until the 17th century. In 1640 Pierre de Fermat stated (without proof) Fermat's little theorem (later proved by Leibniz and Euler). Fermat also conjectured that all numbers of the form 22n + 1 are prime (they are called Fermat numbers) and he verified t...
The most basic method of checking the primality of a given integer n is called trial division. This routine consists of dividing n by each integer m that is greater than 1 and less than or equal to the square root of n. If the result of any of these divisions is an integer, then n is not a prime, otherwise it is a prim...
Modern primality tests for general numbers n can be divided into two main classes, probabilistic (or "Monte Carlo") and deterministic algorithms. Deterministic algorithms provide a way to tell for sure whether a given number is prime or not. For example, trial division is a deterministic algorithm because, if performed...
A particularly simple example of a probabilistic test is the Fermat primality test, which relies on the fact (Fermat's little theorem) that np≡n (mod p) for any n if p is a prime number. If we have a number b that we want to test for primality, then we work out nb (mod b) for a random value of n as our test. A flaw wit...
are prime. Prime numbers of this form are known as factorial primes. Other primes where either p + 1 or p − 1 is of a particular shape include the Sophie Germain primes (primes of the form 2p + 1 with p prime), primorial primes, Fermat primes and Mersenne primes, that is, prime numbers that are of the form 2p − 1, wher...
The following table gives the largest known primes of the mentioned types. Some of these primes have been found using distributed computing. In 2009, the Great Internet Mersenne Prime Search project was awarded a US$100,000 prize for first discovering a prime with at least 10 million digits. The Electronic Frontier Fou...
are prime for any natural number n. Here represents the floor function, i.e., largest integer not greater than the number in question. The latter formula can be shown using Bertrand's postulate (proven first by Chebyshev), which states that there always exists at least one prime number p with n < p < 2n − 2, for any na...
can have infinitely many primes only when a and q are coprime, i.e., their greatest common divisor is one. If this necessary condition is satisfied, Dirichlet's theorem on arithmetic progressions asserts that the progression contains infinitely many primes. The picture below illustrates this with q = 9: the numbers are...
The zeta function is closely related to prime numbers. For example, the aforementioned fact that there are infinitely many primes can also be seen using the zeta function: if there were only finitely many primes then ζ(1) would have a finite value. However, the harmonic series 1 + 1/2 + 1/3 + 1/4 + ... diverges (i.e., ...
The unproven Riemann hypothesis, dating from 1859, states that except for s = −2, −4, ..., all zeroes of the ζ-function have real part equal to 1/2. The connection to prime numbers is that it essentially says that the primes are as regularly distributed as possible.[clarification needed] From a physical viewpoint, it r...
In addition to the Riemann hypothesis, many more conjectures revolving about primes have been posed. Often having an elementary formulation, many of these conjectures have withstood a proof for decades: all four of Landau's problems from 1912 are still unsolved. One of them is Goldbach's conjecture, which asserts that ...
A third type of conjectures concerns aspects of the distribution of primes. It is conjectured that there are infinitely many twin primes, pairs of primes with difference 2 (twin prime conjecture). Polignac's conjecture is a strengthening of that conjecture, it states that for every positive integer n, there are infinit...
For a long time, number theory in general, and the study of prime numbers in particular, was seen as the canonical example of pure mathematics, with no applications outside of the self-interest of studying the topic with the exception of use of prime numbered gear teeth to distribute wear evenly. In particular, number ...
Giuga's conjecture says that this equation is also a sufficient condition for p to be prime. Another consequence of Fermat's little theorem is the following: if p is a prime number other than 2 and 5, 1/p is always a recurring decimal, whose period is p − 1 or a divisor of p − 1. The fraction 1/p expressed likewise in ...
Several public-key cryptography algorithms, such as RSA and the Diffie–Hellman key exchange, are based on large prime numbers (for example, 512-bit primes are frequently used for RSA and 1024-bit primes are typical for Diffie–Hellman.). RSA relies on the assumption that it is much easier (i.e., more efficient) to perfo...
The evolutionary strategy used by cicadas of the genus Magicicada make use of prime numbers. These insects spend most of their lives as grubs underground. They only pupate and then emerge from their burrows after 7, 13 or 17 years, at which point they fly about, breed, and then die after a few weeks at most. The logic ...
The concept of prime number is so important that it has been generalized in different ways in various branches of mathematics. Generally, "prime" indicates minimality or indecomposability, in an appropriate sense. For example, the prime field is the smallest subfield of a field F containing both 0 and 1. It is either Q...
Prime numbers give rise to two more general concepts that apply to elements of any commutative ring R, an algebraic structure where addition, subtraction and multiplication are defined: prime elements and irreducible elements. An element p of R is called prime element if it is neither zero nor a unit (i.e., does not ha...
The fundamental theorem of arithmetic continues to hold in unique factorization domains. An example of such a domain is the Gaussian integers Z[i], that is, the set of complex numbers of the form a + bi where i denotes the imaginary unit and a and b are arbitrary integers. Its prime elements are known as Gaussian prime...
In ring theory, the notion of number is generally replaced with that of ideal. Prime ideals, which generalize prime elements in the sense that the principal ideal generated by a prime element is a prime ideal, are an important tool and object of study in commutative algebra, algebraic number theory and algebraic geomet...
Prime ideals are the points of algebro-geometric objects, via the notion of the spectrum of a ring. Arithmetic geometry also benefits from this notion, and many concepts exist in both geometry and number theory. For example, factorization or ramification of prime ideals when lifted to an extension field, a basic proble...
In particular, this norm gets smaller when a number is multiplied by p, in sharp contrast to the usual absolute value (also referred to as the infinite prime). While completing Q (roughly, filling the gaps) with respect to the absolute value yields the field of real numbers, completing with respect to the p-adic norm \|...
Prime numbers have influenced many artists and writers. The French composer Olivier Messiaen used prime numbers to create ametrical music through "natural phenomena". In works such as La Nativité du Seigneur (1935) and Quatre études de rythme (1949–50), he simultaneously employs motifs with lengths given by different p...
The Rhine (Romansh: Rein, German: Rhein, French: le Rhin, Dutch: Rijn) is a European river that begins in the Swiss canton of Graubünden in the southeastern Swiss Alps, forms part of the Swiss-Austrian, Swiss-Liechtenstein border, Swiss-German and then the Franco-German border, then flows through the Rhineland and even...
The variant forms of the name of the Rhine in modern languages are all derived from the Gaulish name Rēnos, which was adapted in Roman-era geography (1st century BC) as Greek Ῥῆνος (Rhēnos), Latin Rhenus.[note 3] The spelling with Rh- in English Rhine as well as in German Rhein and French Rhin is due to the influence o...
The length of the Rhine is conventionally measured in "Rhine-kilometers" (Rheinkilometer), a scale introduced in 1939 which runs from the Old Rhine Bridge at Constance (0 km) to Hoek van Holland (1036.20 km). The river length is significantly shortened from the river's natural course due to number of canalisation proje...
Near Tamins-Reichenau the Anterior Rhine and the Posterior Rhine join and form the Rhine. The river makes a distinctive turn to the north near Chur. This section is nearly 86 km long, and descends from a height of 599 m to 396 m. It flows through a wide glacial alpine valley known as the Rhine Valley (German: Rheintal)...
The mouth of the Rhine into Lake Constance forms an inland delta. The delta is delimited in the West by the Alter Rhein ("Old Rhine") and in the East by a modern canalized section. Most of the delta is a nature reserve and bird sanctuary. It includes the Austrian towns of Gaißau, Höchst and Fußach. The natural Rhine or...
A regulation of the Rhine was called for, with an upper canal near Diepoldsau and a lower canal at Fußach, in order to counteract the constant flooding and strong sedimentation in the western Rhine Delta. The Dornbirner Ach had to be diverted, too, and it now flows parallel to the canalized Rhine into the lake. Its wat...
Lake Constance consists of three bodies of water: the Obersee ("upper lake"), the Untersee ("lower lake"), and a connecting stretch of the Rhine, called the Seerhein ("Lake Rhine"). The lake is situated in Germany, Switzerland and Austria near the Alps. Specifically, its shorelines lie in the German states of Bavaria a...
The flow of cold, gray mountain water continues for some distance into the lake. The cold water flows near the surface and at first doesn't mix with the warmer, green waters of Upper Lake. But then, at the so-called Rheinbrech, the Rhine water abruptly falls into the depths because of the greater density of cold water....
The Rhine emerges from Lake Constance, flows generally westward, as the Hochrhein, passes the Rhine Falls, and is joined by its major tributary, the river Aare. The Aare more than doubles the Rhine's water discharge, to an average of nearly 1,000 m3/s (35,000 cu ft/s), and provides more than a fifth of the discharge at...
In the centre of Basel, the first major city in the course of the stream, is located the "Rhine knee"; this is a major bend, where the overall direction of the Rhine changes from West to North. Here the High Rhine ends. Legally, the Central Bridge is the boundary between High and Upper Rhine. The river now flows North ...
The Upper Rhine region was changed significantly by a Rhine straightening program in the 19th Century. The rate of flow was increased and the ground water level fell significantly. Dead branches dried up and the amount of forests on the flood plains decreased sharply. On the French side, the Grand Canal d'Alsace was du...
The Rhine is the longest river in Germany. It is here that the Rhine encounters some more of its main tributaries, such as the Neckar, the Main and, later, the Moselle, which contributes an average discharge of more than 300 m3/s (11,000 cu ft/s). Northeastern France drains to the Rhine via the Moselle; smaller rivers ...
Between Bingen and Bonn, the Middle Rhine flows through the Rhine Gorge, a formation which was created by erosion. The rate of erosion equaled the uplift in the region, such that the river was left at about its original level while the surrounding lands raised. The gorge is quite deep and is the stretch of the river wh...
Until the early 1980s, industry was a major source of water pollution. Although many plants and factories can be found along the Rhine up into Switzerland, it is along the Lower Rhine that the bulk of them are concentrated, as the river passes the major cities of Cologne, Düsseldorf and Duisburg. Duisburg is the home o...
The dominant economic sectors in the Middle Rhine area are viniculture and tourism. The Rhine Gorge between Rüdesheim am Rhein and Koblenz is listed as a UNESCO World Heritage Site. Near Sankt Goarshausen, the Rhine flows around the famous rock Lorelei. With its outstanding architectural monuments, the slopes full of v...
The Lower Rhine flows through North Rhine-Westphalia. Its banks are usually heavily populated and industrialized, in particular the agglomerations Cologne, Düsseldorf and Ruhr area. Here the Rhine flows through the largest conurbation in Germany, the Rhine-Ruhr region. One of the most important cities in this region is...
From here, the situation becomes more complicated, as the Dutch name Rijn no longer coincides with the main flow of water. Two thirds of the water flow volume of the Rhine flows farther west, through the Waal and then, via the Merwede and Nieuwe Merwede (De Biesbosch), merging with the Meuse, through the Hollands Diep ...
The other third of the water flows through the Pannerdens Kanaal and redistributes in the IJssel and Nederrijn. The IJssel branch carries one ninth of the water flow of the Rhine north into the IJsselmeer (a former bay), while the Nederrijn carries approximately two ninths of the flow west along a route parallel to the...
The name Rijn, from here on, is used only for smaller streams farther to the north, which together formed the main river Rhine in Roman times. Though they retained the name, these streams no longer carry water from the Rhine, but are used for draining the surrounding land and polders. From Wijk bij Duurstede, the old n...
The Rhine-Meuse Delta, the most important natural region of the Netherlands begins near Millingen aan de Rijn, close to the Dutch-German border with the division of the Rhine into Waal and Nederrijn. Since the Rhine contributes most of the water, the shorter term Rhine Delta is commonly used. However, this name is also...
The shape of the Rhine delta is determined by two bifurcations: first, at Millingen aan de Rijn, the Rhine splits into Waal and Pannerdens Kanaal, which changes its name to Nederrijn at Angeren, and second near Arnhem, the IJssel branches off from the Nederrijn. This creates three main flows, two of which change names ...
Before the St. Elizabeth's flood (1421), the Meuse flowed just south of today's line Merwede-Oude Maas to the North Sea and formed an archipelago-like estuary with Waal and Lek. This system of numerous bays, estuary-like extended rivers, many islands and constant changes of the coastline, is hard to imagine today. From...
The hydrography of the current delta is characterized by the delta's main arms, disconnected arms (Hollandse IJssel, Linge, Vecht, etc.) and smaller rivers and streams. Many rivers have been closed ("dammed") and now serve as drainage channels for the numerous polders. The construction of Delta Works changed the Delta ...
The Rhine-Meuse Delta is a tidal delta, shaped not only by the sedimentation of the rivers, but also by tidal currents. This meant that high tide formed a serious risk because strong tidal currents could tear huge areas of land into the sea. Before the construction of the Delta Works, tidal influence was palpable up to...
In southern Europe, the stage was set in the Triassic Period of the Mesozoic Era, with the opening of the Tethys Ocean, between the Eurasian and African tectonic plates, between about 240 MBP and 220 MBP (million years before present). The present Mediterranean Sea descends from this somewhat larger Tethys sea. At abou...
From the Eocene onwards, the ongoing Alpine orogeny caused a N–S rift system to develop in this zone. The main elements of this rift are the Upper Rhine Graben, in southwest Germany and eastern France and the Lower Rhine Embayment, in northwest Germany and the southeastern Netherlands. By the time of the Miocene, a riv...
Through stream capture, the Rhine extended its watershed southward. By the Pliocene period, the Rhine had captured streams down to the Vosges Mountains, including the Mosel, the Main and the Neckar. The northern Alps were then drained by the Rhone. By the early Pleistocene period, the Rhine had captured most of its cur...
Around 2.5 million years ago (ending 11,600 years ago) was the geological period of the Ice Ages. Since approximately 600,000 years ago, six major Ice Ages have occurred, in which sea level dropped 120 m (390 ft) and much of the continental margins became exposed. In the Early Pleistocene, the Rhine followed a course t...
The last glacial ran from ~74,000 (BP = Before Present), until the end of the Pleistocene (~11,600 BP). In northwest Europe, it saw two very cold phases, peaking around 70,000 BP and around 29,000–24,000 BP. The last phase slightly predates the global last ice age maximum (Last Glacial Maximum). During this time, the l...
Most of the Rhine's current course was not under the ice during the last Ice Age; although, its source must still have been a glacier. A tundra, with Ice Age flora and fauna, stretched across middle Europe, from Asia to the Atlantic Ocean. Such was the case during the Last Glacial Maximum, ca. 22,000–14,000 yr BP, when...
As northwest Europe slowly began to warm up from 22,000 years ago onward, frozen subsoil and expanded alpine glaciers began to thaw and fall-winter snow covers melted in spring. Much of the discharge was routed to the Rhine and its downstream extension. Rapid warming and changes of vegetation, to open forest, began abo...
Since 7500 yr ago, a situation with tides and currents, very similar to present has existed. Rates of sea-level rise had dropped so far, that natural sedimentation by the Rhine and coastal processes together, could compensate the transgression by the sea; in the last 7000 years, the coast line was roughly at the same l...
At the begin of the Holocene (~11,700 years ago), the Rhine occupied its Late-Glacial valley. As a meandering river, it reworked its ice-age braidplain. As sea-level continued to rise in the Netherlands, the formation of the Holocene Rhine-Meuse delta began (~8,000 years ago). Coeval absolute sea-level rise and tectoni...
Since ~3000 yr BP (= years Before Present), human impact is seen in the delta. As a result of increasing land clearance (Bronze Age agriculture), in the upland areas (central Germany), the sediment load of the Rhine has strongly increased and delta growth has sped up. This caused increased flooding and sedimentation, e...
At present, the branches Waal and Nederrijn-Lek discharge to the North Sea, through the former Meuse estuary, near Rotterdam. The river IJssel branch flows to the north and enters the IJsselmeer, formerly the Zuider Zee brackish lagoon; however, since 1932, a freshwater lake. The discharge of the Rhine is divided among...
The Rhine was not known to Herodotus and first enters the historical period in the 1st century BC in Roman-era geography. At that time, it formed the boundary between Gaul and Germania. The Upper Rhine had been part of the areal of the late Hallstatt culture since the 6th century BC, and by the 1st century BC, the area...
From the death of Augustus in AD 14 until after AD 70, Rome accepted as her Germanic frontier the water-boundary of the Rhine and upper Danube. Beyond these rivers she held only the fertile plain of Frankfurt, opposite the Roman border fortress of Moguntiacum (Mainz), the southernmost slopes of the Black Forest and a f...
The Romans kept eight legions in five bases along the Rhine. The actual number of legions present at any base or in all, depended on whether a state or threat of war existed. Between about AD 14 and 180, the assignment of legions was as follows: for the army of Germania Inferior, two legions at Vetera (Xanten), I Germa...
Germanic tribes crossed the Rhine in the Migration period, by the 5th century establishing the kingdoms of Francia on the Lower Rhine, Burgundy on the Upper Rhine and Alemannia on the High Rhine. This "Germanic Heroic Age" is reflected in medieval legend, such as the Nibelungenlied which tells of the hero Siegfried kil...
By the 6th century, the Rhine was within the borders of Francia. In the 9th, it formed part of the border between Middle and Western Francia, but in the 10th century, it was fully within the Holy Roman Empire, flowing through Swabia, Franconia and Lower Lorraine. The mouths of the Rhine, in the county of Holland, fell ...

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