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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 it ages and transitions into a gerontoplast. Plastoglobuli also exhibit a greater size variation under these conditions. They are also common in etioplasts, but decrease in number as the etioplasts mature into chloroplasts.
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 chloroplasts, the vast majority of plastoglobuli occur singularly, attached directly to their parent thylakoid. In old or stressed chloroplasts, plastoglobuli tend to occur in linked groups or chains, still always anchored to a thylakoid.
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 hemispherical starch plates. The starch is accumulated as the pyrenoids mature. In algae with carbon concentrating mechanisms, the enzyme rubisco is found in the pyrenoids. Starch can also accumulate around the pyrenoids when CO2 is scarce. Pyrenoids can divide to form new pyrenoids, or be produced "de novo".
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 lamellar thylakoids. The helices ascend at an angle of 20–25°, connecting to each granal thylakoid at a bridge-like slit junction. The helicoids may extend as large sheets that link multiple grana, or narrow to tube-like bridges between grana. While different parts of the thylakoid system contain different membrane proteins, the thylakoid membranes are continuous and the thylakoid space they enclose form a single continuous labyrinth.
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 use the energized electrons to pump hydrogen ions into the thylakoid space, decreasing the pH and turning it acidic. ATP synthase is a large protein complex that harnesses the concentration gradient of the hydrogen ions in the thylakoid space to generate ATP energy as the hydrogen ions flow back out into the stroma—much like a dam turbine.
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 bottoms of granal thylakoids contain only the relatively flat photosystem II protein complex. This allows them to stack tightly, forming grana with many layers of tightly appressed membrane, called granal membrane, increasing stability and surface area for light capture.
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 leaves of some land plants change color. β-carotene is a bright red-orange carotenoid found in nearly all chloroplasts, like chlorophyll a. Xanthophylls, especially the orange-red zeaxanthin, are also common. Many other forms of carotenoids exist that are only found in certain groups of chloroplasts.
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 across called phycobilisomes. Like photosystem I and ATP synthase, phycobilisomes jut into the stroma, preventing thylakoid stacking in red algal chloroplasts. Cryptophyte chloroplasts and some cyanobacteria don't have their phycobilin pigments organized into phycobilisomes, and keep them in their thylakoid space instead.
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 result of ATP energy being wasted and CO2 being released, all with no sugar being produced. This is a big problem, since O2 is produced by the initial light reactions of photosynthesis, causing issues down the line in the Calvin cycle which uses rubisco.
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 store CO2 in a four-carbon compound, which is why the process is called C4 photosynthesis. The four-carbon compound is then transported to the bundle sheath chloroplasts, where it drops off CO2 and returns to the mesophyll. Bundle sheath chloroplasts do not carry out the light reactions, preventing oxygen from building up in them and disrupting rubisco activity. Because of this, they lack thylakoids organized into grana stacks—though bundle sheath chloroplasts still have free-floating thylakoids in the stroma where they still carry out cyclic electron flow, a light-driven method of synthesizing ATP to power the Calvin cycle without generating oxygen. They lack photosystem II, and only have photosystem I—the only protein complex needed for cyclic electron flow. Because the job of bundle sheath chloroplasts is to carry out the Calvin cycle and make sugar, they often contain large starch grains.
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 chloroplasts can also be found in collenchyma tissue. A plant cell which contains chloroplasts is known as a chlorenchyma cell. A typical chlorenchyma cell of a land plant contains about 10 to 100 chloroplasts.
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 stomata. Palisade mesophyll cells can contain 30–70 chloroplasts per cell, while stomatal guard cells contain only around 8–15 per cell, as well as much less chlorophyll. Chloroplasts can also be found in the bundle sheath cells of a leaf, especially in C4 plants, which carry out the Calvin cycle in their bundle sheath cells. They are often absent from the epidermis of a leaf.
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 sideways so that light strikes them edge-on. This reduces exposure and protects them from photooxidative damage. This ability to distribute chloroplasts so that they can take shelter behind each other or spread out may be the reason why land plants evolved to have many small chloroplasts instead of a few big ones. Chloroplast movement is considered one of the most closely regulated stimulus-response systems that can be found in plants. Mitochondria have also been observed to follow chloroplasts as they move.
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 purposely damaging their photosynthetic system, producing reactive oxygen species. High levels of reactive oxygen species will cause the hypersensitive response. The reactive oxygen species also directly kill any pathogens within the cell. Lower levels of reactive oxygen species initiate systemic acquired resistance, triggering defense-molecule production in the rest of the plant.
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 unstable molecules, so they probably don't leave the chloroplast, but instead pass on their signal to an unknown second messenger molecule. All these molecules initiate retrograde signaling—signals from the chloroplast that regulate gene expression in the nucleus.
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. Photosynthesis is divided into two stages—the light reactions, where water is split to produce oxygen, and the dark reactions, or Calvin cycle, which builds sugar molecules from carbon dioxide. The two phases are linked by the energy carriers adenosine triphosphate (ATP) and nicotinamide adenine dinucleotide phosphate (NADP+).
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' energy to pump hydrogen ions into the thylakoid space, creating a concentration gradient, with more hydrogen ions (up to a thousand times as many) inside the thylakoid system than in the stroma. The hydrogen ions in the thylakoid space then diffuse back down their concentration gradient, flowing back out into the stroma through ATP synthase. ATP synthase uses the energy from the flowing hydrogen ions to phosphorylate adenosine diphosphate into adenosine triphosphate, or ATP. Because chloroplast ATP synthase projects out into the stroma, the ATP is synthesized there, in position to be used in the dark reactions.
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 transport more hydrogen ions into the thylakoid space to generate more ATP. This is termed cyclic photophosphorylation because the electrons are recycled. Cyclic photophosphorylation is common in C4 plants, which need more ATP than NADPH.
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 convert the 3-PGA into glyceraldehyde-3-phosphate, or G3P sugar molecules. Most of the G3P molecules are recycled back into RuBP using energy from more ATP, but one out of every six produced leaves the cycle—the end product of the dark reactions.
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 displace the thylakoids, but leave them intact. Waterlogged roots can also cause starch buildup in the chloroplasts, possibly due to less sucrose being exported out of the chloroplast (or more accurately, the plant cell). This depletes a plant's free phosphate supply, which indirectly stimulates chloroplast starch synthesis. While linked to low photosynthesis rates, the starch grains themselves may not necessarily interfere significantly with the efficiency of photosynthesis, and might simply be a side effect of another photosynthesis-depressing factor.
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 can waste up to half the carbon fixed by the Calvin cycle. Several mechanisms have evolved in different lineages that raise the carbon dioxide concentration relative to oxygen within the chloroplast, increasing the efficiency of photosynthesis. These mechanisms are called carbon dioxide concentrating mechanisms, or CCMs. These include Crassulacean acid metabolism, C4 carbon fixation, and pyrenoids. Chloroplasts in C4 plants are notable as they exhibit a distinct chloroplast dimorphism.
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 to where it is needed. The chloroplast is known to make the precursors to methionine but it is unclear whether the organelle carries out the last leg of the pathway or if it happens in the cytosol.
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 egg. Proplastids are commonly found in an adult plant's apical meristems. Chloroplasts do not normally develop from proplastids in root tip meristems—instead, the formation of starch-storing amyloplasts is more common.
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 body. While etioplasts lack chlorophyll, they have a yellow chlorophyll precursor stocked. Within a few minutes of light exposure, the prolamellar body begins to reorganize into stacks of thylakoids, and chlorophyll starts to be produced. This process, where the etioplast becomes a chloroplast, takes several hours. Gymnosperms do not require light to form chloroplasts.
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 for proplastids to develop straight into chromoplasts. Chromoplasts and amyloplasts can also become chloroplasts, like what happens when a carrot or a potato is illuminated. If a plant is injured, or something else causes a plant cell to revert to a meristematic state, chloroplasts and other plastids can turn back into proplastids. Chloroplast, amyloplast, chromoplast, proplast, etc., are not absolute states—intermediate forms are common.
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 prevents FtsZ from linking up and forming filaments. Another protein ARC3 may also be involved, but it is not very well understood. These proteins are active at the poles of the chloroplast, preventing Z-ring formation there, but near the center of the chloroplast, MinE inhibits them, allowing the Z-ring to form.
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, arranged in rows 6.4 nanometers apart, and shrinks to squeeze the chloroplast. This is when chloroplast constriction begins. In a few species like Cyanidioschyzon merolæ, chloroplasts have a third plastid-dividing ring located in the chloroplast's intermembrane space.
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 been observed to contain many large dumbbell-shaped chloroplasts. Exposure to white light can stimulate these chloroplasts to divide and reduce the population of dumbbell-shaped chloroplasts.
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 cultivation of genetically modified plants that are biologically contained, thus posing significantly lower environmental risks. This biological containment strategy is therefore suitable for establishing the coexistence of conventional and organic agriculture. While the reliability of this mechanism has not yet been studied for all relevant crop species, recent results in tobacco plants are promising, showing a failed containment rate of transplastomic plants at 3 in 1,000,000.
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 has the divisors 2 and 3 in addition to 1 and 6. The fundamental theorem of arithmetic establishes the central role of primes in number theory: any integer greater than 1 can be expressed as a product of primes that is unique up to ordering. The uniqueness in this theorem requires excluding 1 as a prime because one can include arbitrarily many instances of 1 in any factorization, e.g., 3, 1 · 3, 1 · 1 · 3, etc. are all valid factorizations of 3.
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 primality of large numbers. These include the Miller–Rabin primality test, which is fast but has a small probability of error, and the AKS primality test, which always produces the correct answer in polynomial time but is too slow to be practical. Particularly fast methods are available for numbers of special forms, such as Mersenne numbers. As of January 2016[update], the largest known prime number has 22,338,618 decimal digits.
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 the prime number theorem, proven at the end of the 19th century, which says that the probability that a given, randomly chosen number n is prime is inversely proportional to its number of digits, or to the logarithm of n.
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 branches of number theory, focusing on analytic or algebraic aspects of numbers. Primes are used in several routines in information technology, such as public-key cryptography, which makes use of properties such as the difficulty of factoring large numbers into their prime factors. Prime numbers give rise to various generalizations in other mathematical domains, mainly algebra, such as prime elements and prime ideals.
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 number greater than 2. Similarly, when written in the usual decimal system, all prime numbers larger than 5 end in 1, 3, 7, or 9, since even numbers are multiples of 2 and numbers ending in 0 or 5 are multiples of 5.
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 -- who did not agree. In the 19th century many mathematicians still considered the number 1 to be a prime. For example, Derrick Norman Lehmer's list of primes up to 10,006,721, reprinted as late as 1956, started with 1 as its first prime. Henri Lebesgue is said to be the last professional mathematician to call 1 prime. By the early 20th century, mathematicians began to accept that 1 is not a prime number, but rather forms its own special category as a "unit".
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 different factorizations of 15 into prime numbers, so the statement of that theorem would have to be modified. Similarly, the sieve of Eratosthenes would not work correctly if 1 were considered a prime: a modified version of the sieve that considers 1 as prime would eliminate all multiples of 1 (that is, all other numbers) and produce as output only the single number 1. Furthermore, the prime numbers have several properties that the number 1 lacks, such as the relationship of the number to its corresponding value of Euler's totient function or the sum of divisors function.
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 come from the Ancient Greeks. Euclid's Elements (circa 300 BC) contain important theorems about primes, including the infinitude of primes and the fundamental theorem of arithmetic. Euclid also showed how to construct a perfect number from a Mersenne prime. The Sieve of Eratosthenes, attributed to Eratosthenes, is a simple method to compute primes, although the large primes found today with computers are not generated this way.
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 this up to n = 4 (or 216 + 1). However, the very next Fermat number 232 + 1 is composite (one of its prime factors is 641), as Euler discovered later, and in fact no further Fermat numbers are known to be prime. The French monk Marin Mersenne looked at primes of the form 2p − 1, with p a prime. They are called Mersenne primes in his honor.
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 prime. Indeed, if is composite (with a and b ≠ 1) then one of the factors a or b is necessarily at most . For example, for , the trial divisions are by m = 2, 3, 4, 5, and 6. None of these numbers divides 37, so 37 is prime. This routine can be implemented more efficiently if a complete list of primes up to is known—then trial divisions need to be checked only for those m that are prime. For example, to check the primality of 37, only three divisions are necessary (m = 2, 3, and 5), given that 4 and 6 are composite.
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 correctly, it will always identify a prime number as prime and a composite number as composite. Probabilistic algorithms are normally faster, but do not completely prove that a number is prime. These tests rely on testing a given number in a partly random way. For example, a given test might pass all the time if applied to a prime number, but pass only with probability p if applied to a composite number. If we repeat the test n times and pass every time, then the probability that our number is composite is 1/(1-p)n, which decreases exponentially with the number of tests, so we can be as sure as we like (though never perfectly sure) that the number is prime. On the other hand, if the test ever fails, then we know that the number is composite.
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 with this test is that there are some composite numbers (the Carmichael numbers) that satisfy the Fermat identity even though they are not prime, so the test has no way of distinguishing between prime numbers and Carmichael numbers. Carmichael numbers are substantially rarer than prime numbers, though, so this test can be useful for practical purposes. More powerful extensions of the Fermat primality test, such as the Baillie-PSW, Miller-Rabin, and Solovay-Strassen tests, are guaranteed to fail at least some of the time when applied to a composite number.
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, where p is an arbitrary prime. The Lucas–Lehmer test is particularly fast for numbers of this form. This is why the largest known prime has almost always been a Mersenne prime since the dawn of electronic computers.
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 Foundation also offers $150,000 and $250,000 for primes with at least 100 million digits and 1 billion digits, respectively. Some of the largest primes not known to have any particular form (that is, no simple formula such as that of Mersenne primes) have been found by taking a piece of semi-random binary data, converting it to a number n, multiplying it by 256k for some positive integer k, and searching for possible primes within the interval [256kn + 1, 256k(n + 1) − 1].[citation needed]
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 natural number n > 3. However, computing A or μ requires the knowledge of infinitely many primes to begin with. Another formula is based on Wilson's theorem and generates the number 2 many times and all other primes exactly once.
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 "wrapped around" as soon as a multiple of 9 is passed. Primes are highlighted in red. The rows (=progressions) starting with a = 3, 6, or 9 contain at most one prime number. In all other rows (a = 1, 2, 4, 5, 7, and 8) there are infinitely many prime numbers. What is more, the primes are distributed equally among those rows in the long run—the density of all primes congruent a modulo 9 is 1/6.
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., exceeds any given number), so there must be infinitely many primes. Another example of the richness of the zeta function and a glimpse of modern algebraic number theory is the following identity (Basel problem), due to Euler,
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 roughly states that the irregularity in the distribution of primes only comes from random noise. From a mathematical viewpoint, it roughly states that the asymptotic distribution of primes (about x/log x of numbers less than x are primes, the prime number theorem) also holds for much shorter intervals of length about the square root of x (for intervals near x). This hypothesis is generally believed to be correct. In particular, the simplest assumption is that primes should have no significant irregularities without good reason.
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 every even integer n greater than 2 can be written as a sum of two primes. As of February 2011[update], this conjecture has been verified for all numbers up to n = 2 · 1017. Weaker statements than this have been proven, for example Vinogradov's theorem says that every sufficiently large odd integer can be written as a sum of three primes. Chen's theorem says that every sufficiently large even number can be expressed as the sum of a prime and a semiprime, the product of two primes. Also, any even integer can be written as the sum of six primes. The branch of number theory studying such questions is called additive number theory.
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 infinitely many pairs of consecutive primes that differ by 2n. It is conjectured there are infinitely many primes of the form n2 + 1. These conjectures are special cases of the broad Schinzel's hypothesis H. Brocard's conjecture says that there are always at least four primes between the squares of consecutive primes greater than 2. Legendre's conjecture states that there is a prime number between n2 and (n + 1)2 for every positive integer n. It is implied by the stronger Cramér's conjecture.
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 theorists such as British mathematician G. H. Hardy prided themselves on doing work that had absolutely no military significance. However, this vision was shattered in the 1970s, when it was publicly announced that prime numbers could be used as the basis for the creation of public key cryptography algorithms. Prime numbers are also used for hash tables and pseudorandom number generators.
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 base q (rather than base 10) has similar effect, provided that p is not a prime factor of q. Wilson's theorem says that an integer p > 1 is prime if and only if the factorial (p − 1)! + 1 is divisible by p. Moreover, an integer n > 4 is composite if and only if (n − 1)! is divisible by n.
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 perform the multiplication of two (large) numbers x and y than to calculate x and y (assumed coprime) if only the product xy is known. The Diffie–Hellman key exchange relies on the fact that there are efficient algorithms for modular exponentiation, while the reverse operation the discrete logarithm is thought to be a hard problem.
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 for this is believed to be that the prime number intervals between emergences make it very difficult for predators to evolve that could specialize as predators on Magicicadas. If Magicicadas appeared at a non-prime number intervals, say every 12 years, then predators appearing every 2, 3, 4, 6, or 12 years would be sure to meet them. Over a 200-year period, average predator populations during hypothetical outbreaks of 14- and 15-year cicadas would be up to 2% higher than during outbreaks of 13- and 17-year cicadas. Though small, this advantage appears to have been enough to drive natural selection in favour of a prime-numbered life-cycle for these insects.
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 or the finite field with p elements, whence the name. Often a second, additional meaning is intended by using the word prime, namely that any object can be, essentially uniquely, decomposed into its prime components. For example, in knot theory, a prime knot is a knot that is indecomposable in the sense that it cannot be written as the knot sum of two nontrivial knots. Any knot can be uniquely expressed as a connected sum of prime knots. Prime models and prime 3-manifolds are other examples of this type.
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 have a multiplicative inverse) and satisfies the following requirement: given x and y in R such that p divides the product xy, then p divides x or y. An element is irreducible if it is not a unit and cannot be written as a product of two ring elements that are not units. In the ring Z of integers, the set of prime elements equals the set of irreducible elements, which is
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 primes. Not every prime (in Z) is a Gaussian prime: in the bigger ring Z[i], 2 factors into the product of the two Gaussian primes (1 + i) and (1 − i). Rational primes (i.e. prime elements in Z) of the form 4k + 3 are Gaussian primes, whereas rational primes of the form 4k + 1 are not.
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 geometry. The prime ideals of the ring of integers are the ideals (0), (2), (3), (5), (7), (11), … The fundamental theorem of arithmetic generalizes to the Lasker–Noether theorem, which expresses every ideal in a Noetherian commutative ring as an intersection of primary ideals, which are the appropriate generalizations of prime powers.
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 problem of algebraic number theory, bears some resemblance with ramification in geometry. Such ramification questions occur even in number-theoretic questions solely concerned with integers. For example, prime ideals in the ring of integers of quadratic number fields can be used in proving quadratic reciprocity, a statement that concerns the solvability of quadratic equations
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 \|−\|p yields the field of p-adic numbers. These are essentially all possible ways to complete Q, by Ostrowski's theorem. Certain arithmetic questions related to Q or more general global fields may be transferred back and forth to the completed (or local) fields. This local-global principle again underlines the importance of primes to number theory.
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 prime numbers to create unpredictable rhythms: the primes 41, 43, 47 and 53 appear in the third étude, "Neumes rythmiques". According to Messiaen this way of composing was "inspired by the movements of nature, movements of free and unequal durations".
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 eventually empties into the North Sea in the Netherlands. The biggest city on the river Rhine is Cologne, Germany with a population of more than 1,050,000 people. It is the second-longest river in Central and Western Europe (after the Danube), at about 1,230 km (760 mi),[note 2][note 1] with an average discharge of about 2,900 m3/s (100,000 cu ft/s).
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 of Greek orthography, while the vocalisation -i- is due to the Proto-Germanic adoption of the Gaulish name as *Rīnaz, via Old Frankish giving Old English Rín, Old High German Rīn, Dutch Rijn (formerly also spelled Rhijn)). The diphthong in modern German Rhein (also adopted in Romansh Rein, Rain) is a Central German development of the early modern period, the Alemannic name Rī(n) retaining the older vocalism,[note 4] as does Ripuarian Rhing, while Palatine has diphthongized Rhei, Rhoi. Spanish is with French in adopting the Germanic vocalism Rin-, while Italian, Occitan and Portuguese retain the Latin Ren-.
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 projects completed in the 19th and 20th century.[note 7] The "total length of the Rhine", to the inclusion of Lake Constance and the Alpine Rhine is more difficult to measure objectively; it was cited as 1,232 kilometres (766 miles) by the Dutch Rijkswaterstaat in 2010.[note 1]
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). Near Sargans a natural dam, only a few metres high, prevents it from flowing into the open Seeztal valley and then through Lake Walen and Lake Zurich into the river Aare. The Alpine Rhine begins in the most western part of the Swiss canton of Graubünden, and later forms the border between Switzerland to the West and Liechtenstein and later Austria to the East.
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 originally branched into at least two arms and formed small islands by precipitating sediments. In the local Alemannic dialect, the singular is pronounced "Isel" and this is also the local pronunciation of Esel ("Donkey"). Many local fields have an official name containing this element.
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 water has a darker color than the Rhine; the latter's lighter suspended load comes from higher up the mountains. It is expected that the continuous input of sediment into the lake will silt up the lake. This has already happened to the former Lake Tuggenersee.
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 and Baden-Württemberg, the Austrian state of Vorarlberg, and the Swiss cantons of Thurgau and St. Gallen. The Rhine flows into it from the south following the Swiss-Austrian border. It is located at approximately 47°39′N 9°19′E / 47.650°N 9.317°E / 47.650; 9.317.
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 flow reappears on the surface at the northern (German) shore of the lake, off the island of Lindau. The water then follows the northern shore until Hagnau am Bodensee. A small fraction of the flow is diverted off the island of Mainau into Lake Überlingen. Most of the water flows via the Constance hopper into the Rheinrinne ("Rhine Gutter") and Seerhein. Depending on the water level, this flow of the Rhine water is clearly visible along the entire length of the lake.
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 the Dutch border. The Aare also contains the waters from the 4,274 m (14,022 ft) summit of Finsteraarhorn, the highest point of the Rhine basin. The Rhine roughly forms the German-Swiss border from Lake Constance with the exceptions of the canton of Schaffhausen and parts of the cantons of Zürich and Basel-Stadt, until it turns north at the so-called Rhine knee at Basel, leaving Switzerland.
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 as Upper Rhine through the Upper Rhine Plain, which is about 300 km long and up to 40 km wide. The most important tributaries in this area are the Ill below of Strasbourg, the Neckar in Mannheim and the Main across from Mainz. In Mainz, the Rhine leaves the Upper Rhine Valley and flows through the Mainz Basin.
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 dug, which carries a significant part of the river water, and all of the traffic. In some places, there are large compensation pools, for example the huge Bassin de compensation de Plobsheim in Alsace.
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 drain the Vosges and Jura Mountains uplands. Most of Luxembourg and a very small part of Belgium also drain to the Rhine via the Moselle. As it approaches the Dutch border, the Rhine has an annual mean discharge of 2,290 m3/s (81,000 cu ft/s) and an average width of 400 m (1,300 ft).
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 which is known for its many castles and vineyards. It is a UNESCO World Heritage Site (2002) and known as "the Romantic Rhine", with more than 40 castles and fortresses from the Middle Ages and many quaint and lovely country villages.
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 of Europe's largest inland port and functions as a hub to the sea ports of Rotterdam, Antwerp and Amsterdam. The Ruhr, which joins the Rhine in Duisburg, is nowadays a clean river, thanks to a combination of stricter environmental controls, a transition from heavy industry to light industry and cleanup measures, such as the reforestation of Slag and brownfields. The Ruhr currently provides the region with drinking water. It contributes 70 m3/s (2,500 cu ft/s) to the Rhine. Other rivers in the Ruhr Area, above all, the Emscher, still carry a considerable degree of pollution.
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 vines, settlements crowded on the narrow river banks and scores of castles lined up along the top of the steep slopes, the Middle Rhine Valley can be considered the epitome of the Rhine romanticism.
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 Duisburg with the largest river port in Europe (Duisport). The region downstream of Duisburg is more agricultural. In Wesel, 30 km downstream of Duisburg, is located the western end of the second east-west shipping route, the Wesel-Datteln Canal, which runs parallel to the Lippe. Between Emmerich and Cleves the Emmerich Rhine Bridge, the longest suspension bridge in Germany, crosses the 400 m wide river. Near Krefeld, the river crosses the Uerdingen line, the line which separates the areas where Low German and High German are spoken.
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 and Haringvliet estuaries, into the North Sea. The Beneden Merwede branches off, near Hardinxveld-Giessendam and continues as the Noord, to join the Lek, near the village of Kinderdijk, to form the Nieuwe Maas; then flows past Rotterdam and continues via Het Scheur and the Nieuwe Waterweg, to the North Sea. The Oude Maas branches off, near Dordrecht, farther down rejoining the Nieuwe Maas to form Het Scheur.
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 Waal. However, at Wijk bij Duurstede, the Nederrijn changes its name and becomes the Lek. It flows farther west, to rejoin the Noord River into the Nieuwe Maas and to the North Sea.
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 north branch of the Rhine is called Kromme Rijn ("Bent Rhine") past Utrecht, first Leidse Rijn ("Rhine of Leiden") and then, Oude Rijn ("Old Rhine"). The latter flows west into a sluice at Katwijk, where its waters can be discharged into the North Sea. This branch once formed the line along which the Limes Germanicus were built. During periods of lower sea levels within the various ice ages, the Rhine took a left turn, creating the Channel River, the course of which now lies below the English Channel.
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 used for the river delta where the Rhine flows into Lake Constance, so it is clearer to call the larger one Rhine-Meuse delta, or even Rhine–Meuse–Scheldt delta, as the Scheldt ends in the same delta.
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 rather often. The largest and southern main branch begins as Waal and continues as Boven Merwede ("Upper Merwede"), Beneden Merwede ("Lower Merwede"), Noord River ("North River"), Nieuwe Maas ("New Meuse"), Het Scheur ("the Rip") and Nieuwe Waterweg ("New Waterway"). The middle flow begins as Nederrijn, then changes into Lek, then joins the Noord, thereby forming Nieuwe Maas. The northern flow keeps the name IJssel until it flows into Lake IJsselmeer. Three more flows carry significant amounts of water: the Nieuwe Merwede ("New Merwede"), which branches off from the southern branch where it changes from Boven to Beneden Merwede; the Oude Maas ("Old Meuse"), which branches off from the southern branch where it changes from Beneden Merwede into Noord, and Dordtse Kil, which branches off from Oude Maas.
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 1421 to 1904, the Meuse and Waal merged further upstream at Gorinchem to form Merwede. For flood protection reasons, the Meuse was separated from the Waal through a lock and diverted into a new outlet called "Bergse Maas", then Amer and then flows into the former bay Hollands Diep.
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 in the second half of the 20th Century fundamentally. Currently Rhine water runs into the sea, or into former marine bays now separated from the sea, in five places, namely at the mouths of the Nieuwe Merwede, Nieuwe Waterway (Nieuwe Maas), Dordtse Kil, Spui and IJssel.
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 Nijmegen, and even today, after the regulatory action of the Delta Works, the tide acts far inland. At the Waal, for example, the most landward tidal influence can be detected between Brakel and Zaltbommel.
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 about 180 MBP, in the Jurassic Period, the two plates reversed direction and began to compress the Tethys floor, causing it to be subducted under Eurasia and pushing up the edge of the latter plate in the Alpine Orogeny of the Oligocene and Miocene Periods. Several microplates were caught in the squeeze and rotated or were pushed laterally, generating the individual features of Mediterranean geography: Iberia pushed up the Pyrenees; Italy, the Alps, and Anatolia, moving west, the mountains of Greece and the islands. The compression and orogeny continue today, as shown by the ongoing raising of the mountains a small amount each year and the active volcanoes.
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 river system had developed in the Upper Rhine Graben, that continued northward and is considered the first Rhine river. At that time, it did not yet carry discharge from the Alps; instead, the watersheds of the Rhone and Danube drained the northern flanks of the Alps.
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 current Alpine watershed from the Rhône, including the Aar. Since that time, the Rhine has added the watershed above Lake Constance (Vorderrhein, Hinterrhein, Alpenrhein; captured from the Rhône), the upper reaches of the Main, beyond Schweinfurt and the Vosges Mountains, captured from the Meuse, to its watershed.
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 to the northwest, through the present North Sea. During the so-called Anglian glaciation (~450,000 yr BP, marine oxygen isotope stage 12), the northern part of the present North Sea was blocked by the ice and a large lake developed, that overflowed through the English Channel. This caused the Rhine's course to be diverted through the English Channel. Since then, during glacial times, the river mouth was located offshore of Brest, France and rivers, like the Thames and the Seine, became tributaries to the Rhine. During interglacials, when sea level rose to approximately the present level, the Rhine built deltas, in what is now the Netherlands.
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 lower Rhine flowed roughly west through the Netherlands and extended to the southwest, through the English Channel and finally, to the Atlantic Ocean. The English Channel, the Irish Channel and most of the North Sea were dry land, mainly because sea level was approximately 120 m (390 ft) lower than today.
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 ice-sheets covered Scandinavia, the Baltics, Scotland and the Alps, but left the space between as open tundra. The loess or wind-blown dust over that tundra, settled in and around the Rhine Valley, contributing to its current agricultural usefulness.
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 about 13,000 BP. By 9000 BP, Europe was fully forested. With globally shrinking ice-cover, ocean water levels rose and the English Channel and North Sea re-inundated. Meltwater, adding to the ocean and land subsidence, drowned the former coasts of Europe transgressionally.
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 location. In the southern North Sea, due to ongoing tectonic subsidence, the sea level is still rising, at the rate of about 1–3 cm (0.39–1.18 in) per century (1 metre or 39 inches in last 3000 years).
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 tectonic subsidence have strongly influenced delta evolution. Other factors of importance to the shape of the delta are the local tectonic activities of the Peel Boundary Fault, the substrate and geomorphology, as inherited from the Last Glacial and the coastal-marine dynamics, such as barrier and tidal inlet formations.
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, ending peat formation in the delta. The shifting of river channels to new locations, on the floodplain (termed avulsion), was the main process distributing sediment across the subrecent delta. Over the past 6000 years, approximately 80 avulsions have occurred. Direct human impact in the delta started with peat mining, for salt and fuel, from Roman times onward. This was followed by embankment, of the major distributaries and damming of minor distributaries, which took place in the 11–13th century AD. Thereafter, canals were dug, bends were short cut and groynes were built, to prevent the river's channels from migrating or silting up.
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 three branches: the River Waal (6/9 of total discharge), the River Nederrijn – Lek (2/9 of total discharge) and the River IJssel (1/9 of total discharge). This discharge distribution has been maintained since 1709, by river engineering works, including the digging of the Pannerdens canal and since the 20th century, with the help of weirs in the Nederrijn river.
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 areal of the La Tène culture covered almost its entire length, forming a contact zone with the Jastorf culture, i.e. the locus of early Celtic-Germanic cultural contact. In Roman geography, the Rhine formed the boundary between Gallia and Germania by definition; e.g. Maurus Servius Honoratus, Commentary on the Aeneid of Vergil (8.727) (Rhenus) fluvius Galliae, qui Germanos a Gallia dividit "(The Rhine is a) river of Gaul, which divides the Germanic people from Gaul."
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 few scattered bridge-heads. The northern section of this frontier, where the Rhine is deep and broad, remained the Roman boundary until the empire fell. The southern part was different. The upper Rhine and upper Danube are easily crossed. The frontier which they form is inconveniently long, enclosing an acute-angled wedge of foreign territory between the modern Baden and Württemberg. The Germanic populations of these lands seem in Roman times to have been scanty, and Roman subjects from the modern Alsace-Lorraine had drifted across the river eastwards.
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 Germanica and XX Valeria (Pannonian troops); two legions at oppidum Ubiorum ("town of the Ubii"), which was renamed to Colonia Agrippina, descending to Cologne, V Alaudae, a Celtic legion recruited from Gallia Narbonensis and XXI, possibly a Galatian legion from the other side of the empire.
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 killing a dragon on the Drachenfels (Siebengebirge) ("dragons rock"), near Bonn at the Rhine and of the Burgundians and their court at Worms, at the Rhine and Kriemhild's golden treasure, which was thrown into the Rhine by Hagen.
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 to the Burgundian Netherlands in the 15th century; Holland remained contentious territory throughout the European wars of religion and the eventual collapse of the Holy Roman Empire, when the length of the Rhine fell to the First French Empire and its client states. The Alsace on the left banks of the Upper Rhine was sold to Burgundy by Archduke Sigismund of Austria in 1469 and eventually fell to France in the Thirty Years' War. The numerous historic castles in Rhineland-Palatinate attest to the importance of the river as a commercial route.

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