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Arabic numerals
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Arabic numerals, also called Hindu–Arabic numerals,[1][2] are the ten digits: 0, 1, 2, 3, 4, 5, 6, 7, 8, 9, based on the Hindu–Arabic numeral system,[3] the most common system for the symbolic representation of numbers in the world today. In this numeral system, a sequence of digits such as "975" is read as a single number, using the position of the digit in the sequence to interpret its value. They are descended from the Hindu-Arabic numeral system developed by Indian mathematicians around AD 500.[3]
The system was adopted by Arabic mathematicians in Baghdad and passed on to the Arabs farther west. There is some evidence to suggest that the numerals in their current form developed from Arabic letters in the Maghreb, the western region of the Arab world.[4] The current form of the numerals developed in North Africa, distinct in form from the Indian and Eastern Arabic numerals. It was in the North African city of Bejaia that the Italian scholar Fibonacci first encountered the numerals; his work was crucial in making them known throughout Europe. The use of Arabic numerals spread around the world through European trade, books and colonialism.
The term Arabic numerals is ambiguous. It most commonly refers to the numerals widely used in Europe and the Americas; to avoid confusion, Unicode calls these European digits. Arabic numerals is also the European name for the entire family of related numerals of Arabic and Indian numerals. It may also be intended to mean the numerals used by Arabs, in which case it generally refers to the Eastern Arabic numerals. It would be more appropriate to refer to the Arabic numeral system, where the value of a digit in a number depends on its position.
Although the phrase "Arabic numeral" is frequently capitalized, it is sometimes written in lower case: for instance, in its entry in the Oxford English Dictionary,[5] which helps to distinguish it from "Arabic numerals" as the East Arabic numerals specific to the Arabs.
Contents
[hide]
1 History
1.1 Origins
1.1.1 Popular myths
1.2 Adoption in Europe
1.3 Adoption in Russia
1.4 Adoption in China
2 Evolution of symbols
3 See also
4 References
5 Sources
6 Further reading
7 External links
History[edit]
Origins[edit]
Main article: History of the Hindu–Arabic numeral system
The decimal Hindu–Arabic numeral system was developed in India by around AD 700.[6] The development was gradual, spanning several centuries, but the decisive step was probably provided by Brahmagupta's formulation of zero as a number in AD 628. The system was revolutionary by including zero in positional notation, thereby limiting the number of individual digits to ten. It is considered an important milestone in the development of mathematics. One may distinguish between this positional system, which is identical throughout the family, and the precise glyphs used to write the numerals, which varied regionally.
The glyphs most commonly used in conjunction with the Latin script since early modern times are 0 1 2 3 4 5 6 7 8 9. The first universally accepted inscription containing the use of the 0 glyph in India is first recorded in the 9th century, in an inscription at Gwalior in Central India dated to 870. Numerous Indian documents on copper plates exist, with the same symbol for zero in them, dated back as far as the 6th century AD, but their dates are uncertain. Inscriptions in Indonesia and Cambodia dating to AD 683 have also been found.[7]
Brahmi numerals (lower row) in India in the 1st century AD
The numerals used in the Bakhshali manuscript, dated to sometime between the 3rd and 7th century AD.
Modern-day Arab telephone keypad with two forms of Arabic numerals: Western Arabic/European numerals on the left and Eastern Arabic numerals on the right
The numeral system came to be known to the court of Baghdad, where mathematicians such as the Persian Al-Khwarizmi, whose book On the Calculation with Hindu Numerals was written about 825 in Arabic, and the Arab mathematician Al-Kindi, who wrote four volumes, On the Use of the Indian Numerals (Ketab fi Isti'mal al-'Adad al-Hindi) about 830, propagated it in the Arab world. Their work was principally responsible for the diffusion of the Indian system of numeration in the Middle East and the West.[8]
In the 10th century, Middle-Eastern mathematicians extended the decimal numeral system to include fractions, as recorded in a treatise by Syrian mathematician Abu'l-Hasan al-Uqlidisi in 952–953. The decimal point notation was introduced by Sind ibn Ali, who also wrote the earliest treatise on Arabic numerals.
A distinctive West Arabic variant of the symbols begins to emerge around the 10th century in the Maghreb and Al-Andalus (sometimes called ghubar numerals, though the term is not always accepted), which are the direct ancestor of the modern "Arabic numerals" used throughout the world.[9] Woepecke has proposed that the Western Arabic numerals were already in use in Spain before the arrival of the Moors, purportedly received via Alexandria, but this theory is not accepted by scholars.[10][11][12]
Popular myths[edit]
Some popular myths have argued that the original forms of these symbols indicated their numeric value through the number of angles they contained, but no evidence exists of any such origin.[13]
Adoption in Europe[edit]
Adoption of the Hindu numerals through the Arabs by Europe
Woodcut showing the 16th century astronomical clock of Uppsala Cathedral, with two clockfaces, one with Arabic and one with Roman numerals.
A German manuscript page teaching use of Arabic numerals (Talhoffer Thott, 1459). At this time, knowledge of the numerals was still widely seen as esoteric, and Talhoffer presents them with the Hebrew alphabet and astrology.
Late 18th-century French revolutionary "decimal" clockface.
In 825 Al-Khwārizmī wrote a treatise in Arabic, On the Calculation with Hindu Numerals,[14] which survives only as the 12th-century Latin translation, Algoritmi de numero Indorum.[15][16] Algoritmi, the translator's rendition of the author's name, gave rise to the word algorithm.[17]
The first mentions of the numerals in the West are found in the Codex Vigilanus of 976.[18]
From the 980s, Gerbert of Aurillac (later, Pope Sylvester II) used his position to spread knowledge of the numerals in Europe. Gerbert studied in Barcelona in his youth. He was known to have requested mathematical treatises concerning the astrolabe from Lupitus of Barcelona after he had returned to France.
Leonardo Fibonacci (Leonardo of Pisa), a mathematician born in the Republic of Pisa who had studied in Béjaïa (Bougie), Algeria, promoted the Indian numeral system in Europe with his 1202 book Liber Abaci:
When my father, who had been appointed by his country as public notary in the customs at Bugia acting for the Pisan merchants going there, was in charge, he summoned me to him while I was still a child, and having an eye to usefulness and future convenience, desired me to stay there and receive instruction in the school of accounting. There, when I had been introduced to the art of the Indians' nine symbols through remarkable teaching, knowledge of the art very soon pleased me above all else and I came to understand it.
The numerals are arranged with their lowest value digit to the right, with higher value positions added to the left. This arrangement was adopted identically into the numerals as used in Europe. Languages written in the Latin alphabet run from left-to-right, unlike languages written in the Arabic alphabet. Hence, from the point of view of the reader, numerals in Western texts are written with the highest power of the base first whereas numerals in Arabic texts are written with the lowest power of the base first.
The reason the digits are more commonly known as "Arabic numerals" in Europe and the Americas is that they were introduced to Europe in the 10th century by Arabic-speakers of North Africa, who were then using the digits from Libya to Morocco. Arabs, on the other hand, call the system "Hindu numerals",[19][20] referring to their origin in India. This is not to be confused with what the Arabs call the "Hindi numerals", namely the Eastern Arabic numerals (٠ - ١ - ٢ - ٣ -٤ - ٥ - ٦ - ٧ - ٨ - ٩) used in the Middle East, or any of the numerals currently used in Indian languages (e.g. Devanagari: ०.१.२.३.४.५.६.७.८.९).[13]
The European acceptance of the numerals was accelerated by the invention of the printing press, and they became widely known during the 15th century. Early evidence of their use in Britain includes: an equal hour horary quadrant from 1396,[21] in England, a 1445 inscription on the tower of Heathfield Church, Sussex; a 1448 inscription on a wooden lych-gate of Bray Church, Berkshire; and a 1487 inscription on the belfry door at Piddletrenthide church, Dorset; and in Scotland a 1470 inscription on the tomb of the first Earl of Huntly in Elgin Cathedral. (See G.F. Hill, The Development of Arabic Numerals in Europe for more examples.) In central Europe, the King of Hungary Ladislaus the Posthumous, started the use of Arabic numerals, which appear for the first time in a royal document of 1456.[22] By the mid-16th century, they were in common use in most of Europe.[23] Roman numerals remained in use mostly for the notation of Anno Domini years, and for numbers on clockfaces.
Today, Roman numerals are still used for enumeration of lists (as an alternative to alphabetical enumeration), for sequential volumes, to differentiate monarchs or family members with the same first names, and (in lower case) to number pages in prefatory material in books.
Adoption in Russia[edit]
Cyrillic numerals were a numbering system derived from the Cyrillic alphabet, used by South and East Slavic peoples. The system was used in Russia as late as the early 18th century when Peter the Great replaced it with Arabic numerals.
Adoption in China[edit]
Iron plate with an order 6 magic square in Persian/ Arabic numbers from China, dating to the Yuan Dynasty (1271–1368).
Arabic numerals were introduced to China during the Yuan Dynasty (1271–1368) by the Muslim Hui people. In the early 17th century, European-style Arabic numerals were introduced by Spanish and Portuguese Jesuits.[24][25][26]
Evolution of symbols[edit]
Main article: Algorism
The numeral system employed, known as algorism, is positional decimal notation. Various symbol sets are used to represent numbers in the Hindu–Arabic numeral system, potentially including both symbols that evolved from the Brahmi numerals, and symbols that developed independently. The symbols used to represent the system have split into various typographical variants since the Middle Ages:
The widespread Western Arabic numerals used with the Latin script, in the table below labelled European, descended from the West Arabic numerals developed in al-Andalus (Andalucía, Spain) and the Maghreb. Spanish scholars, because of the geographic proximity, trade, and constant warfare with the Muslim kingdoms of Southern Spain, saw a potential in the simplicity of Arabic numbers, and decided to adopt those symbols, and later other Europeans followed suit. There are two typographic styles for rendering European numerals, known as lining figures and text figures.
The Arabic–Indic or Eastern Arabic numerals, used with the Arabic script, developed primarily in what is now Iraq. A variant of the Eastern Arabic numerals used in the Persian and Urdu languages is shown below as East Arabic-Indic.
The Devanagari numerals used with Devanagari and related variants are grouped as Indian numerals.
The evolution of the numerals in early Europe is shown here in a table created by the French scholar Jean-Étienne Montucla in his Histoire de la Mathematique, which was published in 1757:
The Arabic numeral glyphs 0–9 are encoded in ASCII and Unicode at positions 0x30 to 0x39, matching up with the second hexadecimal digit for convenience:
Binary
Octal
Decimal
Hexadecimal
Glyph
0011 0000 060 48 30 0
0011 0001 061 49 31 1
0011 0010 062 50 32 2
0011 0011 063 51 33 3
0011 0100 064 52 34 4
0011 0101 065 53 35 5
0011 0110 066 54 36 6
0011 0111 067 55 37 7
0011 1000 070 56 38 8
0011 1001 071 57 39 9
See also[edit]
Abjad numerals
Chinese numerals
Counting rods – decimal positional numeral system with zero
Decimal
Greek numerals
Japanese numerals
Maya numerals
Regional variations in modern handwritten Arabic numerals
References[edit]
Jump up ^ Schipp, Bernhard; Krämer, Walter (2008), Statistical Inference, Econometric Analysis and Matrix Algebra: Festschrift in Honour of Götz Trenkler, Springer, p. 387, ISBN 9783790821208
Jump up ^ Lumpkin, Beatrice; Strong, Dorothy (1995), Multicultural science and math connections: middle school projects and activities, Walch Publishing, p. 118, ISBN 9780825126598
^ Jump up to: a b Bulliet, Richard; Crossley, Pamela; Headrick,, Daniel; Hirsch, Steven; Johnson, Lyman (2010). The Earth and Its Peoples: A Global History, Volume 1. Cengage Learning. p. 192. ISBN 1439084742. Indian mathematicians invented the concept of zero and developed the "Arabic" numerals and system of place-value notation used in most parts of the world today [better source needed]
Jump up ^ On the Origin of Arabic Numerals - A. Boucenna - Université Ferhat Abbas Setif (in French)
Jump up ^ "Arabic", Oxford English Dictionary, 2nd edition
Jump up ^ O'Connor, J. J. and E. F. Robertson. 2000. Indian Numerals, MacTutor History of Mathematics Archive, School of Mathematics and Statistics, University of St. Andrews, Scotland.
Jump up ^ Plofker 2009, p. 45.
Jump up ^ The MacTutor History of Mathematics archive
Jump up ^ Kunitzsch, The Transmission of Hindu-Arabic Numerals Reconsidered 2003, pp. 12–13: "While specimens of Western Arabic numerals from the early period—the tenth to thirteenth centuries—are still not available, we know at least that Hindu reckoning (called ḥisāb al-ghubār) was known in the West from the tenth century onward..."
Jump up ^ Kunitzsch, The Transmission of Hindu-Arabic Numerals Reconsidered 2003, pp. 12–13: "Since edition of and research on the Pseudo-Boethius[41] we now know that the texts running under his name and carrying Arabic numerals date from the eleventh century. Thus the assumed way of transmission from Alexandria to Spain is impossible and this theory can no longer be taken as serious."
Jump up ^ Smith, D. E.; Karpinski, L. C. (2013) [first published in Boston, 1911], The Hindu-Arabic Numerals, Dover, Chapter V, ISBN 0486155110
Jump up ^ Gandz, Solomon (November 1931), "The Origin of the Ghubār Numerals, or the Arabian Abacus and the Articuli", Isis, 16 (2): 393–424, doi:10.1086/346615, JSTOR 224714
^ Jump up to: a b Ifrah, Georges (1998). The universal history of numbers: from prehistory to the invention of the computer; translated from the French by David Bellos. London: Harvill Press. pp. 356–357. ISBN 9781860463242.
Jump up ^ Philosophy Of Mathematics Francis, John – 2008 – Page 38
Jump up ^ The Ellipse: A Historical and Mathematical Journey Arthur Mazer – 2011
Jump up ^ "al-Khwarizmi - Muslim mathematician".
Jump up ^ Models of Computation: An Introduction to Computability Theory – Page 1 Maribel Fernández – 2009
Jump up ^ "MATHORIGINS.COM_V". www.mathorigins.com.
Jump up ^ Rowlett, Russ (4 July 2004), Roman and "Arabic" Numerals, University of North Carolina at Chapel Hill, retrieved 22 June 2009
Jump up ^ Achenbach, Joel (16 September 1994), Article: Take a Number, Please., The Washington Post, retrieved 22 June 2009
Jump up ^ "14th century timepiece unearthed in Qld farm shed". ABC News.
Jump up ^ Erdélyi: Magyar művelődéstörténet 1-2. kötet. Kolozsvár, 1913, 1918
Jump up ^ Mathforum.org
Jump up ^ Helaine Selin, ed. (31 July 1997). Encyclopaedia of the history of science, technology, and medicine in non-western cultures. Springer. pp. 198–. ISBN 978-0-7923-4066-9. Retrieved 3 March 2012.
Jump up ^ Meuleman, Johan H. (23 August 2002). Islam in the era of globalization: Muslim attitudes towards modernity and identity. Psychology Press. p. 272. ISBN 978-0-7007-1691-3. Retrieved 3 March 2012.
Jump up ^ Peng Yoke Ho (16 October 2000). Li, Qi and Shu: An Introduction to Science and Civilization in China. Courier Dover Publications. p. 106. ISBN 978-0-486-41445-4. Retrieved 3 March 2012.
Sources[edit]
Kunitzsch, Paul (2003), "The Transmission of Hindu-Arabic Numerals Reconsidered", in J. P. Hogendijk; A. I. Sabra, The Enterprise of Science in Islam: New Perspectives, MIT Press, pp. 3–22, ISBN 978-0-262-19482-2
Plofker, Kim (2009), Mathematics in India, Princeton University Pres, ISBN 978-0-691-12067-6
Further reading[edit]
Ore, Oystein (1988), "Hindu-Arabic numerals", Number Theory and Its History, Dover, pp. 19–24, ISBN 0486656209 .
Burnett, Charles (2006), "The Semantics of Indian Numerals in Arabic, Greek and Latin", Journal of Indian Philosophy, Springer-Netherlands, 34 (1–2): 15–30, doi:10.1007/s10781-005-8153-z .
Encyclopædia Britannica (Kim Plofker) (2007), "mathematics, South Asian", Encyclopædia Britannica Online, 189 (4761): 1–12, Bibcode:1961Natur.189S.273., doi:10.1038/189273c0, retrieved 18 May 2007 .
Hayashi, Takao (1995), The Bakhshali Manuscript, An ancient Indian mathematical treatise, Groningen: Egbert Forsten, ISBN 906980087X .
Ifrah, Georges (2000), A Universal History of Numbers: From Prehistory to Computers, New York: Wiley, ASIN 0471393401, ISBN 0471393401 CS1 maint: ASIN uses ISBN (link) .
Katz, Victor J. (ed.) (20 July 2007), The Mathematics of Egypt, Mesopotamia, China, India, and Islam: A Sourcebook, Princeton, New Jersey: Princeton University Press, ISBN 0691114854 CS1 maint: Extra text: authors list (link) .
External links[edit]
Wikimedia Commons has media related to:
Arabic numerals (category)
Development of Hindu Arabic and Traditional Chinese Arithmetic
History of Counting Systems and Numerals. Retrieved 11 December 2005.
The Evolution of Numbers. 16 April 2005.
O'Connor, J. J. and Robertson, E. F. Indian numerals. November 2000.
History of the numerals
Arabic numerals
Hindu-Arabic numerals
Numeral & Numbers' history and curiosities
Gerbert d'Aurillac's early use of Hindu-Arabic numerals at Convergence
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Digestion
From Wikipedia, the free encyclopedia
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For the industrial process, see anaerobic digestion. For the treatment of precipitates in analytical chemistry, see Precipitation (chemistry) § Digestion.
Digestive system
Details
Latin
systema digestorium
Anatomical terminology
[edit on Wikidata]
Digestion is the breakdown of large insoluble food molecules into small water-soluble food molecules so that they can be absorbed into the watery blood plasma. In certain organisms, these smaller substances are absorbed through the small intestine into the blood stream. Digestion is a form of catabolism that is often divided into two processes based on how food is broken down: mechanical and chemical digestion. The term mechanical digestion refers to the physical breakdown of large pieces of food into smaller pieces which can subsequently be accessed by digestive enzymes. In chemical digestion, enzymes break down food into the small molecules the body can use.
In the human digestive system, food enters the mouth and mechanical digestion of the food starts by the action of mastication (chewing), a form of mechanical digestion, and the wetting contact of saliva. Saliva, a liquid secreted by the salivary glands, contains salivary amylase, an enzyme which starts the digestion of starch in the food; the saliva also contains mucus, which lubricates the food, and hydrogen carbonate, which provides the ideal conditions of pH (alkaline) for amylase to work. After undergoing mastication and starch digestion, the food will be in the form of a small, round slurry mass called a bolus. It will then travel down the esophagus and into the stomach by the action of peristalsis. Gastric juice in the stomach starts protein digestion. Gastric juice mainly contains hydrochloric acid and pepsin. As these two chemicals may damage the stomach wall, mucus is secreted by the stomach, providing a slimy layer that acts as a shield against the damaging effects of the chemicals. At the same time protein digestion is occurring, mechanical mixing occurs by peristalsis, which is waves of muscular contractions that move along the stomach wall. This allows the mass of food to further mix with the digestive enzymes.
After some time (typically 1–2 hours in humans, 4–6 hours in dogs, 3–4 hours in house cats),[citation needed] the resulting thick liquid is called chyme. When the pyloric sphincter valve opens, chyme enters the duodenum where it mixes with digestive enzymes from the pancreas and bile juice from the liver and then passes through the small intestine, in which digestion continues. When the chyme is fully digested, it is absorbed into the blood. 95% of absorption of nutrients occurs in the small intestine. Water and minerals are reabsorbed back into the blood in the colon (large intestine) where the pH is slightly acidic about 5.6 ~ 6.9. Some vitamins, such as biotin and vitamin K (K2MK7) produced by bacteria in the colon are also absorbed into the blood in the colon. Waste material is eliminated from the rectum during defecation.[1]
Contents
[hide]
1 Digestive system
1.1 Secretion systems
1.1.1 Channel transport system
1.1.2 Molecular syringe
1.1.3 Conjugation machinery
1.1.4 Release of outer membrane vesicles
1.2 Gastrovascular cavity
1.3 Phagosome
1.4 Specialised organs and behaviours
1.4.1 Beaks
1.4.2 Tongue
1.4.3 Teeth
1.4.4 Crop
1.4.5 Abomasum
1.4.6 Specialised behaviours
1.5 In earthworms
2 Overview of vertebrate digestion
3 Human digestion process
3.1 Neural and biochemical control mechanisms
4 Breakdown into nutrients
4.1 Protein digestion
4.2 Fat digestion
4.3 Carbohydrate digestion
4.4 DNA and RNA digestion
5 Non-destructive digestion
6 Digestive hormones
7 Significance of pH
8 See also
9 References
10 External links
Digestive system
Digestive systems take many forms. There is a fundamental distinction between internal and external digestion. External digestion developed earlier in evolutionary history, and most fungi still rely on it.[2] In this process, enzymes are secreted into the environment surrounding the organism, where they break down an organic material, and some of the products diffuse back to the organism. Animals have a tube (gastrointestinal tract) in which internal digestion occurs, which is more efficient because more of the broken down products can be captured, and the internal chemical environment can be more efficiently controlled.[3]
Some organisms, including nearly all spiders, simply secrete biotoxins and digestive chemicals (e.g., enzymes) into the extracellular environment prior to ingestion of the consequent "soup". In others, once potential nutrients or food is inside the organism, digestion can be conducted to a vesicle or a sac-like structure, through a tube, or through several specialized organs aimed at making the absorption of nutrients more efficient.
Schematic drawing of bacterial conjugation. 1- Donor cell produces pilus. 2- Pilus attaches to recipient cell, bringing the two cells together. 3- The mobile plasmid is nicked and a single strand of DNA is transferred to the recipient cell. 4- Both cells recircularize their plasmids, synthesize second strands, and reproduce pili; both cells are now viable donors.
Secretion systems
Main article: Secretion § Secretion in Gram negative bacteria
Bacteria use several systems to obtain nutrients from other organisms in the environments.
Channel transport system
In a channel transupport system, several proteins form a contiguous channel traversing the inner and outer membranes of the bacteria. It is a simple system, which consists of only three protein subunits: the ABC protein, membrane fusion protein (MFP), and outer membrane protein (OMP)[specify]. This secretion system transports various molecules, from ions, drugs, to proteins of various sizes (20 – 900 kDa). The molecules secreted vary in size from the small Escherichia coli peptide colicin V, (10 kDa) to the Pseudomonas fluorescens cell adhesion protein LapA of 900 kDa.[4]
Molecular syringe
A type III secretion system means that a molecular syringe is used through which a bacterium (e.g. certain types of Salmonella, Shigella, Yersinia) can inject nutrients into protist cells. One such mechanism was first discovered in Y. pestis and showed that toxins could be injected directly from the bacterial cytoplasm into the cytoplasm of its host's cells rather than simply be secreted into the extracellular medium.[5]
Conjugation machinery
The conjugation machinery of some bacteria (and archaeal flagella) is capable of transporting both DNA and proteins. It was discovered in Agrobacterium tumefaciens, which uses this system to introduce the Ti plasmid and proteins into the host, which develops the crown gall (tumor).[6] The VirB complex of Agrobacterium tumefaciens is the prototypic system.[7]
The nitrogen fixing Rhizobia are an interesting case, wherein conjugative elements naturally engage in inter-kingdom conjugation. Such elements as the Agrobacterium Ti or Ri plasmids contain elements that can transfer to plant cells. Transferred genes enter the plant cell nucleus and effectively transform the plant cells into factories for the production of opines, which the bacteria use as carbon and energy sources. Infected plant cells form crown gall or root tumors. The Ti and Ri plasmids are thus endosymbionts of the bacteria, which are in turn endosymbionts (or parasites) of the infected plant.
The Ti and Ri plasmids are themselves conjugative. Ti and Ri transfer between bacteria uses an independent system (the tra, or transfer, operon) from that for inter-kingdom transfer (the vir, or virulence, operon). Such transfer creates virulent strains from previously avirulent Agrobacteria.
Release of outer membrane vesicles
In addition to the use of the multiprotein complexes listed above, Gram-negative bacteria possess another method for release of material: the formation of outer membrane vesicles.[8][9] Portions of the outer membrane pinch off, forming spherical structures made of a lipid bilayer enclosing periplasmic materials. Vesicles from a number of bacterial species have been found to contain virulence factors, some have immunomodulatory effects, and some can directly adhere to and intoxicate host cells. While release of vesicles has been demonstrated as a general response to stress conditions, the process of loading cargo proteins seems to be selective.[10]
Venus Flytrap (Dionaea muscipula) leaf
Gastrovascular cavity
The gastrovascular cavity functions as a stomach in both digestion and the distribution of nutrients to all parts of the body. Extracellular digestion takes place within this central cavity, which is lined with the gastrodermis, the internal layer of epithelium. This cavity has only one opening to the outside that functions as both a mouth and an anus: waste and undigested matter is excreted through the mouth/anus, which can be described as an incomplete gut.
In a plant such as the Venus Flytrap that can make its own food through photosynthesis, it does not eat and digest its prey for the traditional objectives of harvesting energy and carbon, but mines prey primarily for essential nutrients (nitrogen and phosphorus in particular) that are in short supply in its boggy, acidic habitat.[11]
Trophozoites of Entamoeba histolytica with ingested erythrocytes
Phagosome
A phagosome is a vacuole formed around a particle absorbed by phagocytosis. The vacuole is formed by the fusion of the cell membrane around the particle. A phagosome is a cellular compartment in which pathogenic microorganisms can be killed and digested. Phagosomes fuse with lysosomes in their maturation process, forming phagolysosomes. In humans, Entamoeba histolytica can phagocytose red blood cells.[12]
Specialised organs and behaviours
To aid in the digestion of their food animals evolved organs such as beaks, tongues, teeth, a crop, gizzard, and others.
A Catalina Macaw's seed-shearing beak
Squid beak with ruler for size comparison
Beaks
Birds have bony beaks that are specialised according to the bird's ecological niche. For example, macaws primarily eat seeds, nuts, and fruit, using their impressive beaks to open even the toughest seed. First they scratch a thin line with the sharp point of the beak, then they shear the seed open with the sides of the beak.
The mouth of the squid is equipped with a sharp horny beak mainly made of cross-linked proteins. It is used to kill and tear prey into manageable pieces. The beak is very robust, but does not contain any minerals, unlike the teeth and jaws of many other organisms, including marine species.[13] The beak is the only indigestible part of the squid.
Tongue
Main article: Tongue
The tongue is skeletal muscle on the floor of the mouth that manipulates food for chewing (mastication) and swallowing (deglutition). It is sensitive and kept moist by saliva. The underside of the tongue is covered with a smooth mucous membrane. The tongue also has a touch sense for locating and positioning food particles that require further chewing. The tongue is utilized to roll food particles into a bolus before being transported down the esophagus through peristalsis.
The sublingual region underneath the front of the tongue is a location where the oral mucosa is very thin, and underlain by a plexus of veins. This is an ideal location for introducing certain medications to the body. The sublingual route takes advantage of the highly vascular quality of the oral cavity, and allows for the speedy application of medication into the cardiovascular system, bypassing the gastrointestinal tract.
Teeth
Main article: Teeth
Teeth (singular tooth) are small whitish structures found in the jaws (or mouths) of many vertebrates that are used to tear, scrape, milk and chew food. Teeth are not made of bone, but rather of tissues of varying density and hardness, such as enamel, dentine and cementum. Human teeth have a blood and nerve supply which enables proprioception. This is the ability of sensation when chewing, for example if we were to bite into something too hard for our teeth, such as a chipped plate mixed in food, our teeth send a message to our brain and we realise that it cannot be chewed, so we stop trying.
The shapes, sizes and numbers of types of animals' teeth are related to their diets. For example, herbivores have a number of molars which are used to grind plant matter, which is difficult to digest. Carnivores have canine teeth which are used to kill and tear meat.
Crop
A crop, or croup, is a thin-walled expanded portion of the alimentary tract used for the storage of food prior to digestion. In some birds it is an expanded, muscular pouch near the gullet or throat. In adult doves and pigeons, the crop can produce crop milk to feed newly hatched birds.[14]
Certain insects may have a crop or enlarged esophagus.
Rough illustration of a ruminant digestive system
Abomasum
Main article: Abomasum
Herbivores have evolved cecums (or an abomasum in the case of ruminants). Ruminants have a fore-stomach with four chambers. These are the rumen, reticulum, omasum, and abomasum. In the first two chambers, the rumen and the reticulum, the food is mixed with saliva and separates into layers of solid and liquid material. Solids clump together to form the cud (or bolus). The cud is then regurgitated, chewed slowly to completely mix it with saliva and to break down the particle size.
Fibre, especially cellulose and hemi-cellulose, is primarily broken down into the volatile fatty acids, acetic acid, propionic acid and butyric acid in these chambers (the reticulo-rumen) by microbes: (bacteria, protozoa, and fungi). In the omasum, water and many of the inorganic mineral elements are absorbed into the blood stream.
The abomasum is the fourth and final stomach compartment in ruminants. It is a close equivalent of a monogastric stomach (e.g., those in humans or pigs), and digesta is processed here in much the same way. It serves primarily as a site for acid hydrolysis of microbial and dietary protein, preparing these protein sources for further digestion and absorption in the small intestine. Digesta is finally moved into the small intestine, where the digestion and absorption of nutrients occurs. Microbes produced in the reticulo-rumen are also digested in the small intestine.
A flesh fly "blowing a bubble", possibly to concentrate its food by evaporating water
Specialised behaviours
Regurgitation has been mentioned above under abomasum and crop, referring to crop milk, a secretion from the lining of the crop of pigeons and doves with which the parents feed their young by regurgitation.[15]
Many sharks have the ability to turn their stomachs inside out and evert it out of their mouths in order to get rid of unwanted contents (perhaps developed as a way to reduce exposure to toxins).
Other animals, such as rabbits and rodents, practise coprophagia behaviours – eating specialised faeces in order to re-digest food, especially in the case of roughage. Capybara, rabbits, hamsters and other related species do not have a complex digestive system as do, for example, ruminants. Instead they extract more nutrition from grass by giving their food a second pass through the gut. Soft faecal pellets of partially digested food are excreted and generally consumed immediately. They also produce normal droppings, which are not eaten.
Young elephants, pandas, koalas, and hippos eat the faeces of their mother, probably to obtain the bacteria required to properly digest vegetation. When they are born, their intestines do not contain these bacteria (they are completely sterile). Without them, they would be unable to get any nutritional value from many plant components.
In earthworms
An earthworm's digestive system consists of a mouth, pharynx, esophagus, crop, gizzard, and intestine. The mouth is surrounded by strong lips, which act like a hand to grab pieces of dead grass, leaves, and weeds, with bits of soil to help chew. The lips break the food down into smaller pieces. In the pharynx, the food is lubricated by mucus secretions for easier passage. The esophagus adds calcium carbonate to neutralize the acids formed by food matter decay. Temporary storage occurs in the crop where food and calcium carbonate are mixed. The powerful muscles of the gizzard churn and mix the mass of food and dirt. When the churning is complete, the glands in the walls of the gizzard add enzymes to the thick paste, which helps chemically breakdown the organic matter. By peristalsis, the mixture is sent to the intestine where friendly bacteria continue chemical breakdown. This releases carbohydrates, protein, fat, and various vitamins and minerals for absorption into the body.
Overview of vertebrate digestion
In most vertebrates, digestion is a multistage process in the digestive system, starting from ingestion of raw materials, most often other organisms. Ingestion usually involves some type of mechanical and chemical processing. Digestion is separated into four steps:
Ingestion: placing food into the mouth (entry of food in the digestive system),
Mechanical and chemical breakdown: mastication and the mixing of the resulting bolus with water, acids, bile and enzymes in the stomach and intestine to break down complex molecules into simple structures,
Absorption: of nutrients from the digestive system to the circulatory and lymphatic capillaries through osmosis, active transport, and diffusion, and
Egestion (Excretion): Removal of undigested materials from the digestive tract through defecation.
Underlying the process is muscle movement throughout the system through swallowing and peristalsis. Each step in digestion requires energy, and thus imposes an "overhead charge" on the energy made available from absorbed substances. Differences in that overhead cost are important influences on lifestyle, behavior, and even physical structures. Examples may be seen in humans, who differ considerably from other hominids (lack of hair, smaller jaws and musculature, different dentition, length of intestines, cooking, etc.).
The major part of digestion takes place in the small intestine. The large intestine primarily serves as a site for fermentation of indigestible matter by gut bacteria and for resorption of water from digests before excretion.
In mammals, preparation for digestion begins with the cephalic phase in which saliva is produced in the mouth and digestive enzymes are produced in the stomach. Mechanical and chemical digestion begin in the mouth where food is chewed, and mixed with saliva to begin enzymatic processing of starches. The stomach continues to break food down mechanically and chemically through churning and mixing with both acids and enzymes. Absorption occurs in the stomach and gastrointestinal tract, and the process finishes with defecation.[1]
Human digestion process
Main article: Human digestive system
Upper and lower human gastrointestinal tract
The human gastrointestinal tract is around 9 meters long. Food digestion physiology varies between individuals and upon other factors such as the characteristics of the food and size of the meal, and the process of digestion normally takes between 24 and 72 hours.[16]
Digestion begins in the mouth with the secretion of saliva and its digestive enzymes. Food is formed into a bolus by the mechanical mastication and swallowed into the esophagus from where it enters the stomach through the action of peristalsis. Gastric juice contains hydrochloric acid and pepsin which would damage the walls of the stomach and mucus is secreted for protection. In the stomach further release of enzymes break down the food further and this is combined with the churning action of the stomach. The partially digested food enters the duodenum as a thick semi-liquid chyme. In the small intestine, the larger part of digestion takes place and this is helped by the secretions of bile, pancreatic juice and intestinal juice. The intestinal walls are lined with villi, and their epithelial cells is covered with numerous microvilli to improve the absorption of nutrients by increasing the surface area of the intestine.
In the large intestine the passage of food is slower to enable fermentation by the gut flora to take place. Here water is absorbed and waste material stored as feces to be removed by defecation via the anal canal and anus.
Neural and biochemical control mechanisms
Different phases of digestion take place including: the cephalic phase, gastric phase, and intestinal phase.
The cephalic phase occurs at the sight, thought and smell of food, which stimulate the cerebral cortex. Taste and smell stimuli are sent to the hypothalamus and medulla oblongata. After this it is routed through the vagus nerve and release of acetylcholine. Gastric secretion at this phase rises to 40% of maximum rate. Acidity in the stomach is not buffered by food at this point and thus acts to inhibit parietal (secretes acid) and G cell (secretes gastrin) activity via D cell secretion of somatostatin.
The gastric phase takes 3 to 4 hours. It is stimulated by distension of the stomach, presence of food in stomach and decrease in pH. Distention activates long and myenteric reflexes. This activates the release of acetylcholine, which stimulates the release of more gastric juices. As protein enters the stomach, it binds to hydrogen ions, which raises the pH of the stomach. Inhibition of gastrin and gastric acid secretion is lifted. This triggers G cells to release gastrin, which in turn stimulates parietal cells to secrete gastric acid. Gastric acid is about 0.5% hydrochloric acid (HCl), which lowers the pH to the desired pH of 1-3. Acid release is also triggered by acetylcholine and histamine.
The intestinal phase has two parts, the excitatory and the inhibitory. Partially digested food fills the duodenum. This triggers intestinal gastrin to be released. Enterogastric reflex inhibits vagal nuclei, activating sympathetic fibers causing the pyloric sphincter to tighten to prevent more food from entering, and inhibits local reflexes.
Breakdown into nutrients
This section needs expansion with: digestion of other substances. You can help by adding to it. (August 2011)
Protein digestion
Protein digestion occurs in the stomach and duodenum in which 3 main enzymes, pepsin secreted by the stomach and trypsin and chymotrypsin secreted by the pancreas, break down food proteins into polypeptides that are then broken down by various exopeptidases and dipeptidases into amino acids. The digestive enzymes however are mostly secreted as their inactive precursors, the zymogens. For example, trypsin is secreted by pancreas in the form of trypsinogen, which is activated in the duodenum by enterokinase to form trypsin. Trypsin then cleaves proteins to smaller polypeptides.
Fat digestion
Main article: Fatty acid metabolism § Dietary sources of fatty acids, their digestion, absorption, transport in the blood and storage
Digestion of some fats can begin in the mouth where lingual lipase breaks down some short chain lipids into diglycerides. However fats are mainly digested in the small intestine.[17] The presence of fat in the small intestine produces hormones that stimulate the release of pancreatic lipase from the pancreas and bile from the liver which helps in the emulsification of fats for absorption of fatty acids.[17] Complete digestion of one molecule of fat (a triglyceride) results a mixture of fatty acids, mono- and di-glycerides, as well as some undigested triglycerides, but no free glycerol molecules.[17]
Carbohydrate digestion
In humans, dietary starches are composed of glucose units arranged in long chains called amylose, a polysaccharide. During digestion, bonds between glucose molecules are broken by salivary and pancreatic amylase, resulting in progressively smaller chains of glucose. This results in simple sugars glucose and maltose (2 glucose molecules) that can be absorbed by the small intestine.
Lactase is an enzyme that breaks down the disaccharide lactose to its component parts, glucose and galactose. Glucose and galactose can be absorbed by the small intestine. Approximately 65 percent of the adult population produce only small amounts of lactase and are unable to eat unfermented milk-based foods. This is commonly known as lactose intolerance. Lactose intolerance varies widely by ethnic heritage; more than 90 percent of peoples of east Asian descent are lactose intolerant, in contrast to about 5 percent of people of northern European descent.[18]
Sucrase is an enzyme that breaks down the disaccharide sucrose, commonly known as table sugar, cane sugar, or beet sugar. Sucrose digestion yields the sugars fructose and glucose which are readily absorbed by the small intestine.
DNA and RNA digestion
DNA and RNA are broken down into mononucleotides by the nucleases deoxyribonuclease and ribonuclease (DNase and RNase) from the pancreas.
Non-destructive digestion
Some nutrients are complex molecules (for example vitamin B12) which would be destroyed if they were broken down into their functional groups. To digest vitamin B12 non-destructively, haptocorrin in saliva strongly binds and protects the B12 molecules from stomach acid as they enter the stomach and are cleaved from their protein complexes.[19]
After the B12-haptocorrin complexes pass from the stomach via the pylorus to the duodenum, pancreatic proteases cleave haptocorrin from the B12 molecules which rebind to intrinsic factor (IF). These B12-IF complexes travel to the ileum portion of the small intestine where cubilin receptors enable assimilation and circulation of B12-IF complexes in the blood.[20]
Digestive hormones
Action of the major digestive hormones
There are at least five hormones that aid and regulate the digestive system in mammals. There are variations across the vertebrates, as for instance in birds. Arrangements are complex and additional details are regularly discovered. For instance, more connections to metabolic control (largely the glucose-insulin system) have been uncovered in recent years.
Gastrin – is in the stomach and stimulates the gastric glands to secrete pepsinogen (an inactive form of the enzyme pepsin) and hydrochloric acid. Secretion of gastrin is stimulated by food arriving in stomach. The secretion is inhibited by low pH.
Secretin – is in the duodenum and signals the secretion of sodium bicarbonate in the pancreas and it stimulates the bile secretion in the liver. This hormone responds to the acidity of the chyme.
Cholecystokinin (CCK) – is in the duodenum and stimulates the release of digestive enzymes in the pancreas and stimulates the emptying of bile in the gall bladder. This hormone is secreted in response to fat in chyme.
Gastric inhibitory peptide (GIP) – is in the duodenum and decreases the stomach churning in turn slowing the emptying in the stomach. Another function is to induce insulin secretion.
Motilin – is in the duodenum and increases the migrating myoelectric complex component of gastrointestinal motility and stimulates the production of pepsin.
Significance of pH
Digestion is a complex process controlled by several factors. pH plays a crucial role in a normally functioning digestive tract. In the mouth, pharynx and esophagus, pH is typically about 6.8, very weakly acidic. Saliva controls pH in this region of the digestive tract. Salivary amylase is contained in saliva and starts the breakdown of carbohydrates into monosaccharides. Most digestive enzymes are sensitive to pH and will denature in a high or low pH environment.
The stomach's high acidity inhibits the breakdown of carbohydrates within it. This acidity confers two benefits: it denatures proteins for further digestion in the small intestines, and provides non-specific immunity, damaging or eliminating various pathogens.[citation needed]
In the small intestines, the duodenum provides critical pH balancing to activate digestive enzymes. The liver secretes bile into the duodenum to neutralize the acidic conditions from the stomach, and the pancreatic duct empties into the duodenum, adding bicarbonate to neutralize the acidic chyme, thus creating a neutral environment. The mucosal tissue of the small intestines is alkaline with a pH of about 8.5.[citation needed]
See also
Digestive system of gastropods
Digestive system of humpback whales
Erepsin
Gastroesophageal reflux disease
Discovery and development of proton pump inhibitors
References
^ Jump up to: a b Maton, Anthea; Jean Hopkins; Charles William McLaughlin; Susan Johnson; Maryanna Quon Warner; David LaHart; Jill D. Wright (1993). Human Biology and Health. Englewood Cliffs, New Jersey, USA: Prentice Hall. ISBN 0-13-981176-1. OCLC 32308337.
Jump up ^ Dusenbery, David B. (1996). “Life at Small Scale”, pp. 113-115. Scientific American Library, New York. ISBN 0-7167-5060-0.
Jump up ^ Dusenbery, David B. (2009). Living at Micro Scale, p. 280. Harvard University Press, Cambridge, Mass. ISBN 978-0-674-03116-6.
Jump up ^ Wooldridge K (editor) (2009). Bacterial Secreted Proteins: Secretory Mechanisms and Role in Pathogenesis. Caister Academic Press. ISBN 978-1-904455-42-4. CS1 maint: Extra text: authors list (link)
Jump up ^ Salyers, A. A. & Whitt, D. D. (2002). Bacterial Pathogenesis: A Molecular Approach, 2nd ed., Washington, D.C.: ASM Press. ISBN 1-55581-171-X
Jump up ^ Cascales E, Christie PJ (2003). "The versatile Type IV secretion systems". Nature Reviews Microbiology. 1 (2): 137–149. doi:10.1038/nrmicro753. PMC 3873781  . PMID 15035043.
Jump up ^ Christie PJ; Atmakuri K; Jabubowski S; Krishnamoorthy V; Cascales E. (2005). "Biogenesis, architecture, and function of bacterial Type IV secretion systems". Annu Rev Microbiol. 59: 451–485. doi:10.1146/annurev.micro.58.030603.123630. PMC 3872966  . PMID 16153176.
Jump up ^ Chatterjee, S. N.; Das, J (1967). "Electron microscopic observations on the excretion of cell-wall material by Vibrio cholerae". Journal of General Microbiology. 49 (1): 1–11. doi:10.1099/00221287-49-1-1. PMID 4168882.
Jump up ^ Kuehn, M. J.; Kesty, N. C. (2005). "Bacterial outer membrane vesicles and the host-pathogen interaction". Genes & Development. 19 (22): 2645–55. doi:10.1101/gad.1299905. PMID 16291643.
Jump up ^ McBroom, A. J.; Kuehn, M. J. (2007). "Release of outer membrane vesicles by Gram-negative bacteria is a novel envelope stress response". Molecular Microbiology. 63 (2): 545–558. doi:10.1111/j.1365-2958.2006.05522.x. PMC 1868505  . PMID 17163978.
Jump up ^ Leege, Lissa. "How does the Venus flytrap digest flies?". Scientific American. Retrieved 2008-08-20.
Jump up ^ Boettner, D. R.; Huston, C. D.; Linford, A. S.; Buss, S. N.; Houpt, E.; Sherman, N. E.; Petri, W. A. (2008). "Entamoeba histolytica Phagocytosis of Human Erythrocytes Involves PATMK, a Member of the Transmembrane Kinase Family". PLoS Pathogens. 4 (1): e8. doi:10.1371/journal.ppat.0040008. PMC 2211552  . PMID 18208324.
Jump up ^ Miserez, A; Li, Y; Waite, H; Zok, F (2007). "Jumbo squid beaks: Inspiration for design of robust organic composites". Acta Biomaterialia. 3 (1): 139–149. doi:10.1016/j.actbio.2006.09.004. PMID 17113369.
Jump up ^ Gordon John Larkman Ramel (2008-09-29). "The Alimentary Canal in Birds". Retrieved 2008-12-16.
Jump up ^ Levi, Wendell (1977). The Pigeon. Sumter, S.C.: Levi Publishing Co, Inc. ISBN 0-85390-013-2.
Jump up ^ Kong F, Singh RP (June 2008). "Disintegration of solid foods in human stomach". J. Food Sci. 73 (5): R67–80. doi:10.1111/j.1750-3841.2008.00766.x. PMID 18577009.
^ Jump up to: a b c Digestion of fats (triacylglycerols)
Jump up ^ "Genetics Home Reference". US National Library of Medicine. US National Institutes of Health. Retrieved 27 June 2015.
Jump up ^ Nexo E, Hoffmann-Lücke E (July 2011). "Holotranscobalamin, a marker of vitamin B-12 status: analytical aspects and clinical utility". Am. J. Clin. Nutr. 94 (1): 359S–365S. doi:10.3945/ajcn.111.013458. PMC 3127504  . PMID 21593496.
Jump up ^ Viola-Villegas N, Rabideau AE, Bartholomä M, Zubieta J, Doyle RP (August 2009). "Targeting the cubilin receptor through the vitamin B(12) uptake pathway: cytotoxicity and mechanistic insight through fluorescent Re(I) delivery". J. Med. Chem. 52 (16): 5253–61. doi:10.1021/jm900777v. PMID 19627091.
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