title: English phonology

text: A phoneme of a language or dialect is an abstraction of a speech sound or of a group of different sounds which are all perceived to have the same function by speakers of that particular language or dialect. For example, the English word "through" consists of three phonemes: the initial "th" sound, the "r" sound, and a vowel sound. The phonemes in this and many other English words do not always correspond directly to the letters used to spell them (English orthography is not as strongly phonemic as that of many other languages).


title: English phonology

text: The number and distribution of phonemes in English vary from dialect to dialect, and also depend on the interpretation of the individual researcher. The number of consonant phonemes is generally put at 24 (or slightly more depending on the dialect). The number of vowels is subject to greater variation; in the system presented on this page there are 20–25 vowel phonemes in Received Pronunciation, 14–16 in General American and 19–21 in Australian English. The pronunciation keys used in dictionaries generally contain a slightly greater number of symbols than this, to take account of certain sounds used in foreign words and certain noticeable distinctions that may not be—strictly speaking—phonemic.


title: English phonology

text: The following table shows the 24 consonant phonemes found in most dialects of English, plus , whose distribution is more limited. Fortis consonants are always voiceless, aspirated in syllable onset (except in clusters beginning with ), and sometimes also glottalized to an extent in syllable coda (most likely to occur with , see T-glottalization), while lenis consonants are always unaspirated and un-glottalized, and generally partially or fully voiced. The alveolars are usually apical, i.e. pronounced with the tip of the tongue touching or approaching the roof of the mouth, though some speakers produce them laminally, i.e. with the blade of the tongue.


title: English phonology

text: The following table shows typical examples of the occurrence of the above consonant phonemes in words, using minimal pairs where possible.


title: English phonology

text: In most dialects, the fortis stops and affricate have various different allophones, and are distinguished from the lenis stops and affricate by several phonetic features.


title: English phonology

text: English, much like other Germanic languages, has a particularly large number of vowel phonemes, and in addition the vowels of English differ considerably between dialects. Consequently, corresponding vowels may be transcribed with various symbols depending on the dialect under consideration. When considering English as a whole, lexical sets are often used, each named by a word containing the vowel or vowels in question. For example, the set consists of words which, like "lot", have in Received Pronunciation and in General American. The " vowel" then refers to the vowel that appears in those words in whichever dialect is being considered, or (at a greater level of abstraction) to a diaphoneme, which represents this interdialectal correspondence. A commonly used system of lexical sets, devised by John C. Wells, is presented below; for each set, the corresponding phonemes are given for RP and General American, using the notation that will be used on this page.


title: English phonology

text: For a table that shows the pronunciations of these vowels in a wider range of English dialects, see IPA chart for English dialects.


title: English phonology

text: The following tables show the vowel phonemes of three standard varieties of English. The notation system used here for Received Pronunciation (RP) is fairly standard; the others less so. The feature descriptions given here (front, close, etc.) are abstracted somewhat; the actual pronunciations of these vowels are somewhat more accurately conveyed by the IPA symbols used (see Vowel for a chart indicating the meanings of these symbols; though note also the points listed below the following tables). The symbols given in the table are traditional but redirect to their modern implementation.


title: English phonology

text: Listed here are some of the significant cases of allophony of vowels found within standard English dialects.


title: English phonology

text: Unstressed syllables in English may contain almost any vowel, but in practice vowels in stressed and unstressed syllables tend to use different inventories of phonemes. In particular, long vowels are used less often in unstressed syllables than stressed syllables. Additionally there are certain sounds—characterized by central position and weakness—that are particularly often found as the nuclei of unstressed syllables. These include:


title: English phonology

text: Vowel reduction in unstressed syllables is a significant feature of English. Syllables of the types listed above often correspond to a syllable containing a different vowel ("full vowel") used in other forms of the same morpheme where that syllable is stressed. For example, the first "o" in "photograph", being stressed, is pronounced with the vowel, but in "photography", where it is unstressed, it is reduced to schwa. Also, certain common words ("a", "an", "of", "for", etc.) are pronounced with a schwa when they are unstressed, although they have different vowels when they are in a stressed position (see Weak and strong forms in English).


title: English phonology

text: Some unstressed syllables, however, retain full (unreduced) vowels, i.e. vowels other than those listed above. Examples are the in ambition" and the in "finite". Some phonologists regard such syllables as not being fully unstressed (they may describe them as having "tertiary stress"); some dictionaries have marked such syllables as having secondary stress. However linguists such as Ladefoged and regard this as a difference purely of vowel quality and not of stress, and thus argue that vowel reduction itself is phonemic in English. Examples of words where vowel reduction seems to be distinctive for some speakers include "chickaree vs. "chicory (the latter has the reduced vowel of , whereas the former has the vowel without reduction), and "Pharaoh vs. "farrow" (both have the vowel, but in the latter word it may reduce to ).


title: English phonology

text: Lexical stress is phonemic in English. For example, the noun increase" and the verb "increase are distinguished by the positioning of the stress on the first syllable in the former, and on the second syllable in the latter. (See initial-stress-derived noun.) Stressed syllables in English are louder than non-stressed syllables, as well as being longer and having a higher pitch.


title: English phonology

text: In traditional approaches, in any English word consisting of more than one syllable, each syllable is ascribed one of three degrees of stress: "primary", "secondary" or "unstressed". Ordinarily, in each such word there will be exactly one syllable with primary stress, possibly one syllable having secondary stress, and the remainder are unstressed (unusually-long words may have multiple syllables with secondary stress). For example, the word "amazing" has primary stress on the second syllable, while the first and third syllables are unstressed, whereas the word "organization" has primary stress on the fourth syllable, secondary stress on the first, and the second, third, and fifth unstressed. This is often shown in pronunciation keys using the IPA symbols for primary and secondary stress (which are ˈ and ˌ respectively), placed before the syllables to which they apply. The two words just given may therefore be represented (in RP) as and .


title: English phonology

text: Some analysts identify an additional level of stress ("tertiary" stress). This is generally ascribed to syllables that are pronounced with less force than those with secondary stress, but nonetheless contain a "full" or "unreduced" vowel (vowels that are considered to be reduced are listed under above). Hence the third syllable of "organization", if pronounced with as shown above (rather than being reduced to or ), might be said to have tertiary stress. (The precise identification of secondary and tertiary stress differs between analyses; dictionaries do not generally show tertiary stress, although some have taken the approach of marking all syllables with unreduced vowels as having at least secondary stress.)


title: English phonology

text: In some analyses, then, the concept of lexical stress may become conflated with that of vowel reduction. An approach which attempts to separate these two is provided by Peter Ladefoged, who states that it is possible to describe English with only one degree of stress, as long as unstressed syllables are phonemically distinguished for vowel reduction. In this approach, the distinction between primary and secondary stress is regarded as a phonetic or prosodic detail rather than a phonemic feature – primary stress is seen as an example of the predictable "tonic" stress that falls on the "final" stressed syllable of a prosodic unit. For more details of this analysis, see Stress and vowel reduction in English.


title: English phonology

text: Phonotactics is the study of the sequences of phonemes that occur in languages and the sound structures that they form. In this study it is usual to represent consonants in general with the letter C and vowels with the letter V, so that a syllable such as 'be' is described as having CV structure. The IPA symbol used to show a division between syllables is the dot . Syllabification is the process of dividing continuous speech into discrete syllables, a process in which the position of a syllable division is not always easy to decide upon.


title: English phonology

text: Most languages of the world syllabify and sequences as and or , with consonants preferentially acting as the onset of a syllable containing the following vowel. According to one view, English is unusual in this regard, in that stressed syllables attract following consonants, so that and syllabify as and , as long as the consonant cluster is a possible syllable coda; in addition, preferentially syllabifies with the preceding vowel even when both syllables are unstressed, so that occurs as . This is the analysis used in the "Longman Pronunciation Dictionary". However, this view is not widely accepted, as explained in the following section.


title: English phonology

text: The syllable structure in English is (C)V(C), with near-maximal examples being "strengths" (, a (C)V(C) syllable, although it can also be pronounced or ) and "twelfths" (, a (C)V(C) syllable, although it can also be pronounced , , or ). From the phonetic point of view, the analysis of syllable structures is a complex task: because of widespread occurrences of articulatory overlap, English speakers rarely produce an audible release of individual consonants in consonant clusters. This coarticulation can lead to articulatory gestures that seem very much like deletions or complete assimilations. For example, "hundred pounds" may sound like and "jumped back" (in slow speech, ) may sound like , but X-ray and electropalatographic studies demonstrate that inaudible and possibly weakened contacts or lingual gestures may still be made. Thus the second in "hundred pounds" does not entirely assimilate to a labial place of articulation, rather the labial gesture co-occurs with the alveolar one; the "missing" in "jumped back" may still be articulated, though not heard.


title: English phonology

text: Division into syllables is a difficult area, and different theories have been proposed. A widely accepted approach is the maximal onset principle: this states that, subject to certain constraints, any consonants in between vowels should be assigned to the following syllable. Thus the word "leaving" should be divided rather than *, and "hasty" is rather than * or *. However, when such a division results in an onset cluster which is not allowed in English, the division must respect this. Thus if the word "extra" were divided * the resulting onset of the second syllable would be , a cluster which does not occur initially in English. The division is therefore preferred. If assigning a consonant or consonants to the following syllable would result in the preceding syllable ending in an unreduced short vowel, this is avoided. Thus the word "comma" (in RP) should be divided and not *, even though the latter division gives the maximal onset to the following syllable.


title: English phonology

text: In some cases, no solution is completely satisfactory: for example, in British English (RP) the word "hurry" could be divided or , but the former would result in an analysis with a syllable-final (which is held to be non-occurring) while the latter would result in a syllable final (which is said not to occur in this accent). Some phonologists have suggested a compromise analysis where the consonant in the middle belongs to both syllables, and is described as ambisyllabic. In this way, it is possible to suggest an analysis of "hurry" which comprises the syllables and , the medial being ambisyllabic. Where the division coincides with a word boundary, or the boundary between elements of a compound word, it is not usual in the case of dictionaries to insist on the maximal onset principle in a way that divides words in a counter-intuitive way; thus the word "hardware" would be divided by the M.O.P., but dictionaries prefer the division .


title: English phonology

text: In the approach used by the "Longman Pronunciation Dictionary", Wells claims that consonants syllabify with the preceding rather than following vowel when the preceding vowel is the nucleus of a more salient syllable, with stressed syllables being the most salient, reduced syllables the least, and full unstressed vowels ("secondary stress") intermediate. But there are lexical differences as well, frequently but not exclusively with compound words. For example, in "dolphin" and "selfish," Wells argues that the stressed syllable ends in , but in "shellfish," the belongs with the following syllable: → , but → , where the is a little longer and the is not reduced. Similarly, in "toe-strap" Wells argues that the second is a full plosive, as usual in syllable onset, whereas in "toast-rack" the second is in many dialects reduced to the unreleased allophone it takes in syllable codas, or even elided: → ; likewise "nitrate" → with a voiceless (and for some people an affricated "tr" as in "tree"), vs "night-rate" → with a voiced . Cues of syllable boundaries include aspiration of syllable onsets and (in the US) flapping of coda "(a tease" → vs. "at ease" → ), epenthetic stops like in syllable codas ("fence" → but "inside" → ), and r-colored vowels when the is in the coda vs. labialization when it is in the onset "(key-ring" → but "fearing" → ).


title: English phonology

text: Certain English onsets appear only in contractions: e.g. ("<nowiki>'</nowiki>sblood"), and or ("<nowiki>'</nowiki>swounds" or "<nowiki>'</nowiki>dswounds"). Some, such as ("pshaw"), ("fwoosh"), or ("vroom"), can occur in interjections. An archaic voiceless fricative plus nasal exists, ("fnese"), as does an archaic ("snew").


title: English phonology

text: Several additional onsets occur in loan words (with varying degrees of anglicization) such as ("bwana"), ("moiré"), ("noire"), ("zwitterion"), ("zwieback"), ("Dvorak"), ("kvetch"), ("schvartze"), ("Tver"), ("Zwickau"), ("Kjell"), ("Kshatriya"), ("Tlaloc"), ("Vladimir"), ("zloty"), ("Tskhinvali"), ("Hmong"), ("Khmer"), and ("Nganasan").


title: English phonology

text: Some clusters of this type can be converted to regular English phonotactics by simplifying the cluster: e.g. ("dziggetai"), ("Hrolf"), ("croissant"), ("Nguyen"), ("pfennig"), ("phthalic"), ("tsunami"), ("!kung"), and ("Xhosa").


title: English phonology

text: Most (in theory, all) of the following except those that end with , , , , or can be extended with or representing the morpheme -s/-z. Similarly, most (in theory, all) of the following except those that end with or can be extended with or representing the morpheme -t/-d.


title: English phonology

text: For some speakers, a fricative before is elided so that these never appear phonetically: becomes , becomes , or becomes .


title: English phonology

text: Prosodic stress is extra stress given to words or syllables when they appear in certain positions in an utterance, or when they receive special emphasis.


title: English phonology

text: According to Ladefoged's analysis (as referred to under above), English normally has prosodic stress on the final stressed syllable in an intonation unit. This is said to be the origin of the distinction traditionally made at the lexical level between primary and secondary stress: when a word like "admiration" (traditionally transcribed as something like ) is spoken in isolation, or at the end of a sentence, the syllable "ra" (the final stressed syllable) is pronounced with greater force than the syllable "ad", although when the word is not pronounced with this final intonation there may be no difference between the levels of stress of these two syllables.


title: English phonology

text: Prosodic stress can shift for various pragmatic functions, such as focus or contrast. For instance, in the dialogue "Is it brunch tomorrow? No, it's dinner tomorrow", the extra stress shifts from the last stressed syllable of the sentence, "tomorrow", to the last stressed syllable of the emphasized word, "dinner."


title: English phonology

text: Grammatical function words are usually prosodically unstressed, although they can acquire stress when emphasized (as in "Did you find the cat? Well, I found a cat"). Many English function words have distinct strong and weak pronunciations; for example, the word "a" in the last example is pronounced , while the more common unstressed "a" is pronounced . See Weak and strong forms in English.


title: English phonology

text: English is claimed to be a "stress-timed" language. That is, stressed syllables tend to appear with a more or less regular rhythm, while non-stressed syllables are shortened to accommodate this. For example, in the sentence "One make of car is better than another", the syllables "one", "make", "car", "bett-" and ' will be stressed and relatively long, while the other syllables will be considerably shorter. The theory of stress-timing predicts that each of the three unstressed syllables in between "bett-" and ' will be shorter than the syllable "of" between "make" and "car", because three syllables must fit into the same amount of time as that available for "of". However, it should not be assumed that all varieties of English are stress-timed in this way. The English spoken in the West Indies, in Africa and in India are probably better characterized as syllable-timed, though the lack of an agreed scientific test for categorizing an accent or language as stress-timed or syllable-timed may lead one to doubt the value of such a characterization.


title: English phonology

text: Phonological contrasts in intonation can be said to be found in three different and independent domains. In the work of Halliday the following names are proposed:


title: English phonology

text: These terms ("the Three Ts") have been used in more recent work, though they have been criticized for being difficult to remember. American systems such as ToBI also identify contrasts involving boundaries between intonation phrases (Halliday's "tonality"), placement of pitch accent ("tonicity"), and choice of tone or tones associated with the pitch accent ("tone").


title: English phonology

text: Example of phonological contrast involving placement of intonation unit boundaries (boundary marked by comma):


title: English phonology

text: Example of phonological contrast (British English) involving choice of tone (\ = falling tone, \/ = fall-rise tone)


title: English phonology

text: There is typically a contrast involving tone between wh-questions and yes/no questions, the former having a falling tone (e.g. "Where did you \PUT it?") and the latter a rising tone (e.g. "Are you going /OUT?"), though studies of spontaneous speech have shown frequent exceptions to this rule. Tag questions asking for information are said to carry rising tones (e.g. "They are coming on Tuesday, /AREN'T they?") while those asking for confirmation have falling tone (e.g. "Your name's John, \ISN'T it.").


title: English phonology

text: The pronunciation system of English has undergone many changes throughout the history of the language, from the phonological system of Old English, to that of Middle English, through to that of the present day. Variation between dialects has always been significant. Former pronunciations of many words are reflected in their spellings, as English orthography has generally not kept pace with phonological changes since the Middle English period.


title: English phonology

text: The English consonant system has been relatively stable over time, although a number of significant changes have occurred. Examples include the loss (in most dialects) of the and sounds still reflected by the in words like "night" and "taught", and the splitting of voiced and voiceless allophones of fricatives into separate phonemes (such as the two different phonemes represented by). There have also been many changes in consonant clusters, mostly reductions, for instance those that produced the usual modern pronunciations of such letter combinations as , and .


title: English phonology

text: The development of vowels has been much more complex. One of the most notable series of changes is that known as the Great Vowel Shift, which began around the late 14th century. Here the and in words like "price" and "mouth" became diphthongized, and other long vowels became higher: became (as in "meet"), became and later (as in "name"), became (as in "goose"), and became and later (in RP now ; as in "bone"). These shifts are responsible for the modern pronunciations of many written vowel combinations, including those involving a silent final.


title: English phonology

text: Many other changes in vowels have taken place over the centuries (see the separate articles on the low back, high back and high front vowels, short A, and diphthongs). These various changes mean that many words that formerly rhymed (and may be expected to rhyme based on their spelling) no longer do. For example, in Shakespeare's time, following the Great Vowel Shift, "food", "good" and "blood" all had the vowel , but in modern pronunciation "good" has been shortened to , while "blood" has been shortened and lowered to in most accents. In other cases, words that were formerly distinct have come to be pronounced the same – examples of such mergers include "meet–meat", "pane–pain" and "toe–tow".


title: English phonology

text: The phonemic status of the velar nasal consonant is disputed; one analysis claims that the only nasal phonemes in English are and , while is an allophone of found before velar consonants. Evidence in support of this analysis is found in accents of the north-west Midlands of England where is only found before or , with "sung" being pronounced as . However, in most other accents of English "sung" is pronounced , producing a three-way phonemic contrast "sum" – "sun" – "sung" and supporting the analysis of the phonemic status of . In support of treating the velar nasal as an allophone of , claims on psychological grounds that did not form part of a series of three nasal consonants: "no naïve English-speaking person can be made to feel in his bones that it belongs to a single series with "m" and "n". ... It still "feels" like "ƞg"." More recent writers have indicated that analyses of as an allophone of may still have merit, even though may appear both with and without a following velar consonant; in such analyses, an underlying that is deleted by a phonological rule would account for occurrences of not followed by a velar consonant. Thus the phonemic representation of "sing" would be and that of "singer" is ; in order to reach the phonetic form and , it is necessary to apply a rule that changes to before or , then a second rule that deletes when it follows .


title: English phonology

text: In the above cases, the is not deleted. The words are all single morphemes, unlike "singer" and "singing" which are composed of two morphemes, "sing" plus "-er" or "-ing". Rule 2 can be amended to include a symbol # for a morpheme boundary (including word boundary):


title: English phonology

text: According to this rule, the words "hangar" ('shed for aircraft'), which contains no internal morpheme boundary, and "hanger" ('object for hanging clothes'), which comprises two morphemes, are expected to constitute a minimal pair as "hangar" versus "hanger" ; in actuality, their pronunciations are not consistently distinguished in this manner, as "hangar" is frequently pronounced .


title: English phonology

text: Additionally, there are exceptions in the form of comparative and superlative forms of adjectives, where Rule 2 must be prevented from applying. The ending "-ish" is another possible exception.


title: English phonology

text: As a result, there is, in theory, a minimal pair consisting of "longer" ( 'more long') and "longer" ( 'person who longs'), though it is doubtful that native speakers make this distinction regularly. Names of persons and places, and loanwords, are less predictable. "Singapore" may be pronounced with or without ; "bungalow" usually has ; and "Inge" may or may not have .


title: English phonology

text: It is often stated that English has a particularly large number of vowel phonemes and that there are 20 vowel phonemes in Received Pronunciation, 14–16 in General American, and 20–21 in Australian English. These numbers, however, reflect just one of many possible phonological analyses. A number of "biphonemic" analyses have proposed that English has a basic set of short (sometimes called "simple" or "checked") vowels, each of which can be shown to be a phoneme and which can be combined with another phoneme to form long vowels and diphthongs. One of these biphonemic analyses asserts that diphthongs and long vowels may be interpreted as comprising a short vowel linked to a consonant. The fullest exposition of this approach is found in , where all long vowels and diphthongs ("complex nuclei") are made up of a short vowel combined with either (for which the authors use the symbol ), or (plus for rhotic accents), each thus comprising two phonemes. Using this system, the word "bite" would be transcribed , "bout" as , "bar" as and "bra" as . One attraction that the authors claim for this analysis is that it regularizes the distribution of the consonants , , and (as well as in non-rhotic accents), which would otherwise not be found in syllable-final position. suggest nine simple vowel phonemes to allow them to represent all the accents of American and British English they surveyed, symbolized (front vowels); (central vowels); and (back vowels).


title: English phonology

text: The analysis from came out of a desire to build an "overall system" to accommodate all English dialects, with dialectal distinctions arising from differences in the ordering of phonological rules, as well as in the presence or absence of such rules. Another category of biphonemic analyses of English treats long vowels and diphthongs as conjunctions of two vowels. Such analyses, as found in or for example, are less concerned with dialectal variation. In , for example, there are seven basic vowels and these may be doubled (geminated) to represent long vowels, as shown in the table below:


title: English phonology

text: Some of the short vowels may also be combined with ( "bay", "buy", "boy"), with ( , "beau") or with ( "peer", "pair", "poor"). The vowel inventory of English RP in MacCarthy's system therefore totals only seven phonemes. Analyses such as these could also posit six vowel phonemes, if the vowel of the final syllable in "comma" is considered to be an unstressed allophone of that of "strut". These seven vowels might be symbolized , , , , , and . Six or seven vowels is a figure that would put English much closer to the average number of vowel phonemes in other languages.


title: English phonology

text: A radically different approach to the English vowel system was proposed by Chomsky and Halle. Their "Sound Pattern of English" proposed that English has lax and tense vowel phonemes which are operated on by a complex set of phonological rules to transform underlying phonological forms into surface phonetic representations. This generative analysis is not easily comparable with conventional analyses, but the total number of vowel phonemes proposed falls well short of the figure of 20 often claimed as the number of English vowel phonemes.


title: Wildlife

text: Wildlife refers to undomesticated animal species, but has come to include all organisms that grow or live wild in an area without being introduced by humans. Wildlife was also synonymous to game: those birds and mammals that were hunted for sport. Wildlife can be found in all ecosystems. Deserts, plains, grasslands, woodlands, forests, and other areas, including the most developed urban areas, all have distinct forms of wildlife. While the term in popular culture usually refers to animals that are untouched by human factors, most scientists agree that much wildlife is affected by human activities. Some wildlife threaten human safety, health, property, and quality of life. However, many wild animals, even the dangerous ones, have value to human beings. This value might be economic, educational, or emotional in nature.


title: Wildlife

text: Humans have historically tended to separate civilization from wildlife in a number of ways, including the legal, social, and moral senses. Some animals, however, have adapted to suburban environments. This includes such animals as feral cats, dogs, mice, and rats. Some religions declare certain animals to be sacred, and in modern times, concern for the natural environment has provoked activists to protest against the exploitation of wildlife for human benefit or entertainment.


title: Wildlife

text: Global wildlife populations have decreased by 68% since 1970 as a result of human activity, particularly overconsumption, population growth, and intensive farming, according to a 2020 World Wildlife Fund's "Living Planet Report" and the Zoological Society of London's Living Planet Index measure, which is further evidence that humans have unleashed a sixth mass extinction event. According to CITES, it has been estimated that annually the international wildlife trade amounts to billions of dollars and it affects hundreds of millions of animal and plant specimen.


title: Wildlife

text: Stone Age people and hunter-gatherers relied on wildlife, both plants and animals, for their food. In fact, some species may have been hunted to extinction by early human hunters. Today, hunting, fishing, and gathering wildlife is still a significant food source in some parts of the world. In other areas, hunting and non-commercial fishing are mainly seen as a sport or recreation. Meat sourced from wildlife that is not traditionally regarded as game is known as bushmeat. The increasing demand for wildlife as a source of traditional food in East Asia is decimating populations of sharks, primates, pangolins and other animals, which they believe have aphrodisiac properties.


title: Wildlife

text: A November 2008 report from biologist and author Sally Kneidel, PhD, documented numerous wildlife species for sale in informal markets along the Amazon River, including wild-caught marmosets sold for as little as $1.60 (5 Peruvian soles). Many Amazon species, including peccaries, agoutis, turtles, turtle eggs, anacondas, armadillos are sold primarily as food.


title: Wildlife

text: Wildlife has long been a common subject for educational television shows. National Geographic Society specials appeared on CBS since 1965, later moving to American Broadcasting Company and then Public Broadcasting Service. In 1963, NBC debuted "Wild Kingdom," a popular program featuring zoologist Marlin Perkins as host. The BBC natural history unit in the United Kingdom was a similar pioneer, the first wildlife series LOOK presented by Sir Peter Scott, was a studio-based show, with filmed inserts. David Attenborough first made his appearance in this series, which was followed by the series Zoo Quest during which he and cameraman Charles Lagus went to many exotic places looking for and filming elusive wildlife—notably the Komodo dragon in Indonesia and lemurs in Madagascar.


title: Wildlife

text: Since 1984, the Discovery Channel and its spinoff Animal Planet in the US have dominated the market for shows about wildlife on cable television, while on Public Broadcasting Service the NATURE strand made by WNET-13 in New York and NOVA by WGBH in Boston are notable. Wildlife television is now a multimillion-dollar industry with specialist documentary film-makers in many countries including UK, US, New Zealand, Australia, Austria, Germany, Japan, and Canada.


title: Wildlife

text: There are many magazines and websites which cover wildlife including National Wildlife Magazine, Birds & Blooms, Birding (magazine), wildlife.net and Ranger Rick for children.


title: Wildlife

text: Many animal species have spiritual significance in different cultures around the world, and they and their products may be used as sacred objects in religious rituals. For example, eagles, hawks and their feathers have great cultural and spiritual value to Native Americans as religious objects. In Hinduism the cow is regarded as sacred.


title: Wildlife

text: Muslims conduct sacrifices on Eid al-Adha, to commemorate the sacrificial spirit of Ibrāhīm in Islam ( Arabic-Abraham) in love of God. Camels, sheep, goats may be offered as sacrifice during the three days of Eid.


title: Wildlife

text: In Christianity the Bible has a variety of animal symbols, the Lamb is a famous title of Jesus. In the New Testament the Gospels Mark, Luke and John have animal symbols: "Mark is a lion, Luke is a bull and John is an eagle."


title: Wildlife

text: This subsection focuses on anthropogenic forms of wildlife destruction. The loss of animals from ecological communities is also known as "defaunation."


title: Wildlife

text: Exploitation of wild populations has been a characteristic of modern man since our exodus from Africa 130,000 – 70,000 years ago. The rate of extinctions of entire species of plants and animals across the planet has been so high in the last few hundred years it is widely believed that we are in the sixth great extinction event on earth; the Holocene Mass Extinction. The 2019 "Global Assessment Report on Biodiversity and Ecosystem Services", published by the United Nations' Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services, says that roughly one million species of plants and animals face extinction within decades as the result of human actions.


title: Wildlife

text: Destruction of wildlife does not always lead to an extinction of the species in question, however, the dramatic loss of entire species across Earth dominates any review of wildlife destruction as extinction is the level of damage to a wild population from which there is no return.


title: Wildlife

text: The four most general reasons that lead to destruction of wildlife include overkill, habitat destruction and fragmentation, impact of introduced species and chains of extinction.


title: Wildlife

text: Overkill happens whenever hunting occurs at rates greater than the reproductive capacity of the population is being exploited. The effects of this are often noticed much more dramatically in slow-growing populations such as many larger species of fish. Initially when a portion of a wild population is hunted, an increased availability of resources (food, etc.) is experienced increasing growth and reproduction as density dependent inhibition is lowered. Hunting, fishing and so on, has lowered the competition between members of a population. However, if this hunting continues at rate greater than the rate at which new members of the population can reach breeding age and produce more young, the population will begin to decrease in numbers.


title: Wildlife

text: Populations that are confined to islands, whether literal islands or just areas of habitat that are effectively an "island" for the species concerned, have also been observed to be at greater risk of dramatic population rise of deaths declines following unsustainable hunting.


title: Wildlife

text: The habitat of any given species is considered its preferred area or territory. Many processes associated with human habitation of an area cause loss of this area and decrease the carrying capacity of the land for that species. In many cases these changes in land use cause a patchy break-up of the wild landscape. Agricultural land frequently displays this type of extremely fragmented, or relictual, habitat. Farms sprawl across the landscape with patches of uncleared woodland or forest dotted in-between occasional paddocks.


title: Wildlife

text: Examples of habitat destruction include grazing of bushland by farmed animals, changes to natural fire regimes, forest clearing for timber production and wetland draining for city expansion.


title: Wildlife

text: Mice, cats, rabbits, dandelions and poison ivy are all examples of species that have become invasive threats to wild species in various parts of the world. Frequently species that are uncommon in their home range become out-of-control invasions in distant but similar climates. The reasons for this have not always been clear and Charles Darwin felt it was unlikely that exotic species would ever be able to grow abundantly in a place in which they had not evolved. The reality is that the vast majority of species exposed to a new habitat do not reproduce successfully. Occasionally, however, some populations do take hold and after a period of acclimation can increase in numbers significantly, having destructive effects on many elements of the native environment of which they have become part.


title: Wildlife

text: This final group is one of secondary effects. All wild populations of living things have many complex intertwining links with other living things around them. Large herbivorous animals such as the hippopotamus have populations of insectivorous birds that feed off the many parasitic insects that grow on the hippo. Should the hippo die out, so too will these groups of birds, leading to further destruction as other species dependent on the birds are affected. Also referred to as a domino effect, this series of chain reactions is by far the most destructive process that can occur in any ecological community.


title: Wildlife

text: Another example is the black drongos and the cattle egrets found in India. These birds feed on insects on the back of cattle, which helps to keep them disease-free. Destroying the nesting habitats of these birds would cause a decrease in the cattle population because of the spread of insect-borne diseases.


title: Botany

text: Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek word ("botanē") meaning "pasture", "herbs" "grass", or "fodder"; is in turn derived from (), "to feed" or "to graze". Traditionally, botany has also included the study of fungi and algae by mycologists and phycologists respectively, with the study of these three groups of organisms remaining within the sphere of interest of the International Botanical Congress. Nowadays, botanists (in the strict sense) study approximately 410,000 species of land plants of which some 391,000 species are vascular plants (including approximately 369,000 species of flowering plants), and approximately 20,000 are bryophytes.


title: Botany

text: Botany originated in prehistory as herbalism with the efforts of early humans to identify – and later cultivate – edible, medicinal and poisonous plants, making it one of the oldest branches of science. Medieval physic gardens, often attached to monasteries, contained plants of medical importance. They were forerunners of the first botanical gardens attached to universities, founded from the 1540s onwards. One of the earliest was the Padua botanical garden. These gardens facilitated the academic study of plants. Efforts to catalogue and describe their collections were the beginnings of plant taxonomy, and led in 1753 to the binomial system of nomenclature of Carl Linnaeus that remains in use to this day for the naming of all biological species.


title: Botany

text: In the 19th and 20th centuries, new techniques were developed for the study of plants, including methods of optical microscopy and live cell imaging, electron microscopy, analysis of chromosome number, plant chemistry and the structure and function of enzymes and other proteins. In the last two decades of the 20th century, botanists exploited the techniques of molecular genetic analysis, including genomics and proteomics and DNA sequences to classify plants more accurately.


title: Botany

text: Modern botany is a broad, multidisciplinary subject with contributions and insights from most other areas of science and technology. Research topics include the study of plant structure, growth and differentiation, reproduction, biochemistry and primary metabolism, chemical products, development, diseases, evolutionary relationships, systematics, and plant taxonomy. Dominant themes in 21st century plant science are molecular genetics and epigenetics, which study the mechanisms and control of gene expression during differentiation of plant cells and tissues. Botanical research has diverse applications in providing staple foods, materials such as timber, oil, rubber, fibre and drugs, in modern horticulture, agriculture and forestry, plant propagation, breeding and genetic modification, in the synthesis of chemicals and raw materials for construction and energy production, in environmental management, and the maintenance of biodiversity.


title: Botany

text: Botany originated as herbalism, the study and use of plants for their medicinal properties. The early recorded history of botany includes many ancient writings and plant classifications. Examples of early botanical works have been found in ancient texts from India dating back to before 1100 BCE, Ancient Egypt, in archaic Avestan writings, and in works from China purportedly from before 221 BCE.


title: Botany

text: Modern botany traces its roots back to Ancient Greece specifically to Theophrastus (–287 BCE), a student of Aristotle who invented and described many of its principles and is widely regarded in the scientific community as the "Father of Botany". His major works, "Enquiry into Plants" and "On the Causes of Plants", constitute the most important contributions to botanical science until the Middle Ages, almost seventeen centuries later.


title: Botany

text: Another work from Ancient Greece that made an early impact on botany is "De materia medica", a five-volume encyclopedia about herbal medicine written in the middle of the first century by Greek physician and pharmacologist Pedanius Dioscorides. "De materia medica" was widely read for more than 1,500 years. Important contributions from the medieval Muslim world include Ibn Wahshiyya's "Nabatean Agriculture", Abū Ḥanīfa Dīnawarī's (828–896) the "Book of Plants", and Ibn Bassal's "The Classification of Soils". In the early 13th century, Abu al-Abbas al-Nabati, and Ibn al-Baitar (d. 1248) wrote on botany in a systematic and scientific manner.


title: Botany

text: In the mid-16th century, botanical gardens were founded in a number of Italian universities. The Padua botanical garden in 1545 is usually considered to be the first which is still in its original location. These gardens continued the practical value of earlier "physic gardens", often associated with monasteries, in which plants were cultivated for medical use. They supported the growth of botany as an academic subject. Lectures were given about the plants grown in the gardens and their medical uses demonstrated. Botanical gardens came much later to northern Europe; the first in England was the University of Oxford Botanic Garden in 1621. Throughout this period, botany remained firmly subordinate to medicine.


title: Botany

text: German physician Leonhart Fuchs (1501–1566) was one of "the three German fathers of botany", along with theologian Otto Brunfels (1489–1534) and physician Hieronymus Bock (1498–1554) (also called Hieronymus Tragus). Fuchs and Brunfels broke away from the tradition of copying earlier works to make original observations of their own. Bock created his own system of plant classification.


title: Botany

text: Physician Valerius Cordus (1515–1544) authored a botanically and pharmacologically important herbal "Historia Plantarum" in 1544 and a pharmacopoeia of lasting importance, the "Dispensatorium" in 1546. Naturalist Conrad von Gesner (1516–1565) and herbalist John Gerard (1545–) published herbals covering the medicinal uses of plants. Naturalist Ulisse Aldrovandi (1522–1605) was considered the "father of natural history", which included the study of plants. In 1665, using an early microscope, Polymath Robert Hooke discovered cells, a term he coined, in cork, and a short time later in living plant tissue.


title: Botany

text: During the 18th century, systems of plant identification were developed comparable to dichotomous keys, where unidentified plants are placed into taxonomic groups (e.g. family, genus and species) by making a series of choices between pairs of characters. The choice and sequence of the characters may be artificial in keys designed purely for identification (diagnostic keys) or more closely related to the natural or phyletic order of the taxa in synoptic keys. By the 18th century, new plants for study were arriving in Europe in increasing numbers from newly discovered countries and the European colonies worldwide. In 1753, Carl von Linné (Carl Linnaeus) published his Species Plantarum, a hierarchical classification of plant species that remains the reference point for modern botanical nomenclature. This established a standardised binomial or two-part naming scheme where the first name represented the genus and the second identified the species within the genus. For the purposes of identification, Linnaeus's "Systema Sexuale" classified plants into 24 groups according to the number of their male sexual organs. The 24th group, "Cryptogamia", included all plants with concealed reproductive parts, mosses, liverworts, ferns, algae and fungi.


title: Botany

text: Increasing knowledge of plant anatomy, morphology and life cycles led to the realisation that there were more natural affinities between plants than the artificial sexual system of Linnaeus. Adanson (1763), de Jussieu (1789), and Candolle (1819) all proposed various alternative natural systems of classification that grouped plants using a wider range of shared characters and were widely followed. The Candollean system reflected his ideas of the progression of morphological complexity and the later Bentham & Hooker system, which was influential until the mid-19th century, was influenced by Candolle's approach. Darwin's publication of the "Origin of Species" in 1859 and his concept of common descent required modifications to the Candollean system to reflect evolutionary relationships as distinct from mere morphological similarity.


title: Botany

text: Botany was greatly stimulated by the appearance of the first "modern" textbook, Matthias Schleiden's "", published in English in 1849 as "Principles of Scientific Botany". Schleiden was a microscopist and an early plant anatomist who co-founded the cell theory with Theodor Schwann and Rudolf Virchow and was among the first to grasp the significance of the cell nucleus that had been described by Robert Brown in 1831.


title: Botany

text: In 1855, Adolf Fick formulated Fick's laws that enabled the calculation of the rates of molecular diffusion in biological systems.


title: Botany

text: Building upon the gene-chromosome theory of heredity that originated with Gregor Mendel (1822–1884), August Weismann (1834–1914) proved that inheritance only takes place through gametes. No other cells can pass on inherited characters. The work of Katherine Esau (1898–1997) on plant anatomy is still a major foundation of modern botany. Her books "Plant Anatomy" and "Anatomy of Seed Plants" have been key plant structural biology texts for more than half a century.


title: Botany

text: The discipline of plant ecology was pioneered in the late 19th century by botanists such as Eugenius Warming, who produced the hypothesis that plants form communities, and his mentor and successor Christen C. Raunkiær whose system for describing plant life forms is still in use today. The concept that the composition of plant communities such as temperate broadleaf forest changes by a process of ecological succession was developed by Henry Chandler Cowles, Arthur Tansley and Frederic Clements. Clements is credited with the idea of climax vegetation as the most complex vegetation that an environment can support and Tansley introduced the concept of ecosystems to biology. Building on the extensive earlier work of Alphonse de Candolle, Nikolai Vavilov (1887–1943) produced accounts of the biogeography, centres of origin, and evolutionary history of economic plants.


title: Botany

text: Particularly since the mid-1960s there have been advances in understanding of the physics of plant physiological processes such as transpiration (the transport of water within plant tissues), the temperature dependence of rates of water evaporation from the leaf surface and the molecular diffusion of water vapour and carbon dioxide through stomatal apertures. These developments, coupled with new methods for measuring the size of stomatal apertures, and the rate of photosynthesis have enabled precise description of the rates of gas exchange between plants and the atmosphere. Innovations in statistical analysis by Ronald Fisher, Frank Yates and others at Rothamsted Experimental Station facilitated rational experimental design and data analysis in botanical research. The discovery and identification of the auxin plant hormones by Kenneth V. Thimann in 1948 enabled regulation of plant growth by externally applied chemicals. Frederick Campion Steward pioneered techniques of micropropagation and plant tissue culture controlled by plant hormones. The synthetic auxin 2,4-dichlorophenoxyacetic acid or 2,4-D was one of the first commercial synthetic herbicides.


title: Botany

text: 20th century developments in plant biochemistry have been driven by modern techniques of organic chemical analysis, such as spectroscopy, chromatography and electrophoresis. With the rise of the related molecular-scale biological approaches of molecular biology, genomics, proteomics and metabolomics, the relationship between the plant genome and most aspects of the biochemistry, physiology, morphology and behaviour of plants can be subjected to detailed experimental analysis. The concept originally stated by Gottlieb Haberlandt in 1902 that all plant cells are totipotent and can be grown "in vitro" ultimately enabled the use of genetic engineering experimentally to knock out a gene or genes responsible for a specific trait, or to add genes such as GFP that report when a gene of interest is being expressed. These technologies enable the biotechnological use of whole plants or plant cell cultures grown in bioreactors to synthesise pesticides, antibiotics or other pharmaceuticals, as well as the practical application of genetically modified crops designed for traits such as improved yield.


title: Botany

text: Modern morphology recognises a continuum between the major morphological categories of root, stem (caulome), leaf (phyllome) and trichome. Furthermore, it emphasises structural dynamics. Modern systematics aims to reflect and discover phylogenetic relationships between plants. Modern Molecular phylogenetics largely ignores morphological characters, relying on DNA sequences as data. Molecular analysis of DNA sequences from most families of flowering plants enabled the Angiosperm Phylogeny Group to publish in 1998 a phylogeny of flowering plants, answering many of the questions about relationships among angiosperm families and species. The theoretical possibility of a practical method for identification of plant species and commercial varieties by DNA barcoding is the subject of active current research.


title: Botany

text: The study of plants is vital because they underpin almost all animal life on Earth by generating a large proportion of the oxygen and food that provide humans and other organisms with aerobic respiration with the chemical energy they need to exist. Plants, algae and cyanobacteria are the major groups of organisms that carry out photosynthesis, a process that uses the energy of sunlight to convert water and carbon dioxide into sugars that can be used both as a source of chemical energy and of organic molecules that are used in the structural components of cells. As a by-product of photosynthesis, plants release oxygen into the atmosphere, a gas that is required by nearly all living things to carry out cellular respiration. In addition, they are influential in the global carbon and water cycles and plant roots bind and stabilise soils, preventing soil erosion. Plants are crucial to the future of human society as they provide food, oxygen, medicine, and products for people, as well as creating and preserving soil.


title: Botany

text: Historically, all living things were classified as either animals or plants and botany covered the study of all organisms not considered animals. Botanists examine both the internal functions and processes within plant organelles, cells, tissues, whole plants, plant populations and plant communities. At each of these levels, a botanist may be concerned with the classification (taxonomy), phylogeny and evolution, structure (anatomy and morphology), or function (physiology) of plant life.


title: Botany

text: The strictest definition of "plant" includes only the "land plants" or embryophytes, which include seed plants (gymnosperms, including the pines, and flowering plants) and the free-sporing cryptogams including ferns, clubmosses, liverworts, hornworts and mosses. Embryophytes are multicellular eukaryotes descended from an ancestor that obtained its energy from sunlight by photosynthesis. They have life cycles with alternating haploid and diploid phases. The sexual haploid phase of embryophytes, known as the gametophyte, nurtures the developing diploid embryo sporophyte within its tissues for at least part of its life, even in the seed plants, where the gametophyte itself is nurtured by its parent sporophyte. Other groups of organisms that were previously studied by botanists include bacteria (now studied in bacteriology), fungi (mycology) – including lichen-forming fungi (lichenology), non-chlorophyte algae (phycology), and viruses (virology). However, attention is still given to these groups by botanists, and fungi (including lichens) and photosynthetic protists are usually covered in introductory botany courses.


title: Botany

text: Palaeobotanists study ancient plants in the fossil record to provide information about the evolutionary history of plants. Cyanobacteria, the first oxygen-releasing photosynthetic organisms on Earth, are thought to have given rise to the ancestor of plants by entering into an endosymbiotic relationship with an early eukaryote, ultimately becoming the chloroplasts in plant cells. The new photosynthetic plants (along with their algal relatives) accelerated the rise in atmospheric oxygen started by the cyanobacteria, changing the ancient oxygen-free, reducing, atmosphere to one in which free oxygen has been abundant for more than 2 billion years.


title: Botany

text: Among the important botanical questions of the 21st century are the role of plants as primary producers in the global cycling of life's basic ingredients: energy, carbon, oxygen, nitrogen and water, and ways that our plant stewardship can help address the global environmental issues of resource management, conservation, human food security, biologically invasive organisms, carbon sequestration, climate change, and sustainability.


title: Botany

text: Virtually all staple foods come either directly from primary production by plants, or indirectly from animals that eat them. Plants and other photosynthetic organisms are at the base of most food chains because they use the energy from the sun and nutrients from the soil and atmosphere, converting them into a form that can be used by animals. This is what ecologists call the first trophic level. The modern forms of the major staple foods, such as hemp, teff, maize, rice, wheat and other cereal grasses, pulses, bananas and plantains, as well as hemp, flax and cotton grown for their fibres, are the outcome of prehistoric selection over thousands of years from among wild ancestral plants with the most desirable characteristics.


title: Botany

text: Botanists study how plants produce food and how to increase yields, for example through plant breeding, making their work important to humanity's ability to feed the world and provide food security for future generations. Botanists also study weeds, which are a considerable problem in agriculture, and the biology and control of plant pathogens in agriculture and natural ecosystems. Ethnobotany is the study of the relationships between plants and people. When applied to the investigation of historical plant–people relationships ethnobotany may be referred to as archaeobotany or palaeoethnobotany. Some of the earliest plant-people relationships arose between the indigenous people of Canada in identifying edible plants from inedible plants. This relationship the indigenous people had with plants was recorded by ethnobotanists.


title: Botany

text: Plant biochemistry is the study of the chemical processes used by plants. Some of these processes are used in their primary metabolism like the photosynthetic Calvin cycle and crassulacean acid metabolism. Others make specialised materials like the cellulose and lignin used to build their bodies, and secondary products like resins and aroma compounds.


title: Botany

text: Plants and various other groups of photosynthetic eukaryotes collectively known as "algae" have unique organelles known as chloroplasts. Chloroplasts are thought to be descended from cyanobacteria that formed endosymbiotic relationships with ancient plant and algal ancestors. Chloroplasts and cyanobacteria contain the blue-green pigment chlorophyll "a". Chlorophyll "a" (as well as its plant and green algal-specific cousin chlorophyll "b") absorbs light in the blue-violet and orange/red parts of the spectrum while reflecting and transmitting the green light that we see as the characteristic colour of these organisms. The energy in the red and blue light that these pigments absorb is used by chloroplasts to make energy-rich carbon compounds from carbon dioxide and water by oxygenic photosynthesis, a process that generates molecular oxygen (O) as a by-product.


title: Botany

text: The light energy captured by chlorophyll "a" is initially in the form of electrons (and later a proton gradient) that's used to make molecules of ATP and NADPH which temporarily store and transport energy. Their energy is used in the light-independent reactions of the Calvin cycle by the enzyme rubisco to produce molecules of the 3-carbon sugar glyceraldehyde 3-phosphate (G3P). Glyceraldehyde 3-phosphate is the first product of photosynthesis and the raw material from which glucose and almost all other organic molecules of biological origin are synthesised. Some of the glucose is converted to starch which is stored in the chloroplast. Starch is the characteristic energy store of most land plants and algae, while inulin, a polymer of fructose is used for the same purpose in the sunflower family Asteraceae. Some of the glucose is converted to sucrose (common table sugar) for export to the rest of the plant.


title: Botany

text: Unlike in animals (which lack chloroplasts), plants and their eukaryote relatives have delegated many biochemical roles to their chloroplasts, including synthesising all their fatty acids, and most amino acids. The fatty acids that chloroplasts make are used for many things, such as providing material to build cell membranes out of and making the polymer cutin which is found in the plant cuticle that protects land plants from drying out.


title: Botany

text: Plants synthesise a number of unique polymers like the polysaccharide molecules cellulose, pectin and xyloglucan from which the land plant cell wall is constructed.


title: Botany

text: Vascular land plants make lignin, a polymer used to strengthen the secondary cell walls of xylem tracheids and vessels to keep them from collapsing when a plant sucks water through them under water stress. Lignin is also used in other cell types like sclerenchyma fibres that provide structural support for a plant and is a major constituent of wood. Sporopollenin is a chemically resistant polymer found in the outer cell walls of spores and pollen of land plants responsible for the survival of early land plant spores and the pollen of seed plants in the fossil record. It is widely regarded as a marker for the start of land plant evolution during the Ordovician period.


title: Botany

text: The concentration of carbon dioxide in the atmosphere today is much lower than it was when plants emerged onto land during the Ordovician and Silurian periods. Many monocots like maize and the pineapple and some dicots like the Asteraceae have since independently evolved pathways like Crassulacean acid metabolism and the carbon fixation pathway for photosynthesis which avoid the losses resulting from photorespiration in the more common carbon fixation pathway. These biochemical strategies are unique to land plants.


title: Botany

text: Phytochemistry is a branch of plant biochemistry primarily concerned with the chemical substances produced by plants during secondary metabolism. Some of these compounds are toxins such as the alkaloid coniine from hemlock. Others, such as the essential oils peppermint oil and lemon oil are useful for their aroma, as flavourings and spices (e.g., capsaicin), and in medicine as pharmaceuticals as in opium from opium poppies. Many medicinal and recreational drugs, such as tetrahydrocannabinol (active ingredient in cannabis), caffeine, morphine and nicotine come directly from plants. Others are simple derivatives of botanical natural products. For example, the pain killer aspirin is the acetyl ester of salicylic acid, originally isolated from the bark of willow trees, and a wide range of opiate painkillers like heroin are obtained by chemical modification of morphine obtained from the opium poppy. Popular stimulants come from plants, such as caffeine from coffee, tea and chocolate, and nicotine from tobacco. Most alcoholic beverages come from fermentation of carbohydrate-rich plant products such as barley (beer), rice (sake) and grapes (wine). Native Americans have used various plants as ways of treating illness or disease for thousands of years. This knowledge Native Americans have on plants has been recorded by enthnobotanists and then in turn has been used by pharmaceutical companies as a way of drug discovery.


title: Botany

text: Plants can synthesise useful coloured dyes and pigments such as the anthocyanins responsible for the red colour of red wine, yellow weld and blue woad used together to produce Lincoln green, indoxyl, source of the blue dye indigo traditionally used to dye denim and the artist's pigments gamboge and rose madder.


title: Botany

text: Sugar, starch, cotton, linen, hemp, some types of rope, wood and particle boards, papyrus and paper, vegetable oils, wax, and natural rubber are examples of commercially important materials made from plant tissues or their secondary products. Charcoal, a pure form of carbon made by pyrolysis of wood, has a long history as a metal-smelting fuel, as a filter material and adsorbent and as an artist's material and is one of the three ingredients of gunpowder. Cellulose, the world's most abundant organic polymer, can be converted into energy, fuels, materials and chemical feedstock. Products made from cellulose include rayon and cellophane, wallpaper paste, biobutanol and gun cotton. Sugarcane, rapeseed and soy are some of the plants with a highly fermentable sugar or oil content that are used as sources of biofuels, important alternatives to fossil fuels, such as biodiesel. Sweetgrass was used by Native Americans to ward off bugs like mosquitoes. These bug repelling properties of sweetgrass were later found by the American Chemical Society in the molecules phytol and coumarin.


title: Botany

text: Plant ecology is the science of the functional relationships between plants and their habitats – the environments where they complete their life cycles. Plant ecologists study the composition of local and regional floras, their biodiversity, genetic diversity and fitness, the adaptation of plants to their environment, and their competitive or mutualistic interactions with other species. Some ecologists even rely on empirical data from indigenous people that is gathered by ethnobotanists. This information can relay a great deal of information on how the land once was thousands of years ago and how it has changed over that time. The goals of plant ecology are to understand the causes of their distribution patterns, productivity, environmental impact, evolution, and responses to environmental change.


title: Botany

text: Plants depend on certain edaphic (soil) and climatic factors in their environment but can modify these factors too. For example, they can change their environment's albedo, increase runoff interception, stabilise mineral soils and develop their organic content, and affect local temperature. Plants compete with other organisms in their ecosystem for resources. They interact with their neighbours at a variety of spatial scales in groups, populations and communities that collectively constitute vegetation. Regions with characteristic vegetation types and dominant plants as well as similar abiotic and biotic factors, climate, and geography make up biomes like tundra or tropical rainforest.


title: Botany

text: Herbivores eat plants, but plants can defend themselves and some species are parasitic or even carnivorous. Other organisms form mutually beneficial relationships with plants. For example, mycorrhizal fungi and rhizobia provide plants with nutrients in exchange for food, ants are recruited by ant plants to provide protection, honey bees, bats and other animals pollinate flowers and humans and other animals act as dispersal vectors to spread spores and seeds.


title: Botany

text: Plant responses to climate and other environmental changes can inform our understanding of how these changes affect ecosystem function and productivity. For example, plant phenology can be a useful proxy for temperature in historical climatology, and the biological impact of climate change and global warming. Palynology, the analysis of fossil pollen deposits in sediments from thousands or millions of years ago allows the reconstruction of past climates. Estimates of atmospheric concentrations since the Palaeozoic have been obtained from stomatal densities and the leaf shapes and sizes of ancient land plants. Ozone depletion can expose plants to higher levels of ultraviolet radiation-B (UV-B), resulting in lower growth rates. Moreover, information from studies of community ecology, plant systematics, and taxonomy is essential to understanding vegetation change, habitat destruction and species extinction.


title: Botany

text: Inheritance in plants follows the same fundamental principles of genetics as in other multicellular organisms. Gregor Mendel discovered the genetic laws of inheritance by studying inherited traits such as shape in "Pisum sativum" (peas). What Mendel learned from studying plants has had far-reaching benefits outside of botany. Similarly, "jumping genes" were discovered by Barbara McClintock while she was studying maize. Nevertheless, there are some distinctive genetic differences between plants and other organisms.


title: Botany

text: Species boundaries in plants may be weaker than in animals, and cross species hybrids are often possible. A familiar example is peppermint, "Mentha" × "piperita", a sterile hybrid between "Mentha aquatica" and spearmint, "Mentha spicata". The many cultivated varieties of wheat are the result of multiple inter- and intra-specific crosses between wild species and their hybrids. Angiosperms with monoecious flowers often have self-incompatibility mechanisms that operate between the pollen and stigma so that the pollen either fails to reach the stigma or fails to germinate and produce male gametes. This is one of several methods used by plants to promote outcrossing. In many land plants the male and female gametes are produced by separate individuals. These species are said to be dioecious when referring to vascular plant sporophytes and dioicous when referring to bryophyte gametophytes.


title: Botany

text: Unlike in higher animals, where parthenogenesis is rare, asexual reproduction may occur in plants by several different mechanisms. The formation of stem tubers in potato is one example. Particularly in arctic or alpine habitats, where opportunities for fertilisation of flowers by animals are rare, plantlets or bulbs, may develop instead of flowers, replacing sexual reproduction with asexual reproduction and giving rise to clonal populations genetically identical to the parent. This is one of several types of apomixis that occur in plants. Apomixis can also happen in a seed, producing a seed that contains an embryo genetically identical to the parent.


title: Botany

text: Most sexually reproducing organisms are diploid, with paired chromosomes, but doubling of their chromosome number may occur due to errors in cytokinesis. This can occur early in development to produce an autopolyploid or partly autopolyploid organism, or during normal processes of cellular differentiation to produce some cell types that are polyploid (endopolyploidy), or during gamete formation. An allopolyploid plant may result from a hybridisation event between two different species. Both autopolyploid and allopolyploid plants can often reproduce normally, but may be unable to cross-breed successfully with the parent population because there is a mismatch in chromosome numbers. These plants that are reproductively isolated from the parent species but live within the same geographical area, may be sufficiently successful to form a new species. Some otherwise sterile plant polyploids can still reproduce vegetatively or by seed apomixis, forming clonal populations of identical individuals. Durum wheat is a fertile tetraploid allopolyploid, while bread wheat is a fertile hexaploid. The commercial banana is an example of a sterile, seedless triploid hybrid. Common dandelion is a triploid that produces viable seeds by apomictic seed.


title: Botany

text: As in other eukaryotes, the inheritance of endosymbiotic organelles like mitochondria and chloroplasts in plants is non-Mendelian. Chloroplasts are inherited through the male parent in gymnosperms but often through the female parent in flowering plants.


title: Botany

text: A considerable amount of new knowledge about plant function comes from studies of the molecular genetics of model plants such as the Thale cress, "Arabidopsis thaliana", a weedy species in the mustard family (Brassicaceae). The genome or hereditary information contained in the genes of this species is encoded by about 135 million base pairs of DNA, forming one of the smallest genomes among flowering plants. "Arabidopsis" was the first plant to have its genome sequenced, in 2000. The sequencing of some other relatively small genomes, of rice ("Oryza sativa") and "Brachypodium distachyon", has made them important model species for understanding the genetics, cellular and molecular biology of cereals, grasses and monocots generally.


title: Botany

text: Model plants such as "Arabidopsis thaliana" are used for studying the molecular biology of plant cells and the chloroplast. Ideally, these organisms have small genomes that are well known or completely sequenced, small stature and short generation times. Corn has been used to study mechanisms of photosynthesis and phloem loading of sugar in plants. The single celled green alga "Chlamydomonas reinhardtii", while not an embryophyte itself, contains a green-pigmented chloroplast related to that of land plants, making it useful for study. A red alga "Cyanidioschyzon merolae" has also been used to study some basic chloroplast functions. Spinach, peas, soybeans and a moss "Physcomitrella patens" are commonly used to study plant cell biology.


title: Botany

text: "Agrobacterium tumefaciens", a soil rhizosphere bacterium, can attach to plant cells and infect them with a callus-inducing Ti plasmid by horizontal gene transfer, causing a callus infection called crown gall disease. Schell and Van Montagu (1977) hypothesised that the Ti plasmid could be a natural vector for introducing the Nif gene responsible for nitrogen fixation in the root nodules of legumes and other plant species. Today, genetic modification of the Ti plasmid is one of the main techniques for introduction of transgenes to plants and the creation of genetically modified crops.


title: Botany

text: Epigenetics is the study of heritable changes in gene function that cannot be explained by changes in the underlying DNA sequence but cause the organism's genes to behave (or "express themselves") differently. One example of epigenetic change is the marking of the genes by DNA methylation which determines whether they will be expressed or not. Gene expression can also be controlled by repressor proteins that attach to silencer regions of the DNA and prevent that region of the DNA code from being expressed. Epigenetic marks may be added or removed from the DNA during programmed stages of development of the plant, and are responsible, for example, for the differences between anthers, petals and normal leaves, despite the fact that they all have the same underlying genetic code. Epigenetic changes may be temporary or may remain through successive cell divisions for the remainder of the cell's life. Some epigenetic changes have been shown to be heritable, while others are reset in the germ cells.


title: Botany

text: Epigenetic changes in eukaryotic biology serve to regulate the process of cellular differentiation. During morphogenesis, totipotent stem cells become the various pluripotent cell lines of the embryo, which in turn become fully differentiated cells. A single fertilised egg cell, the zygote, gives rise to the many different plant cell types including parenchyma, xylem vessel elements, phloem sieve tubes, guard cells of the epidermis, etc. as it continues to divide. The process results from the epigenetic activation of some genes and inhibition of others.


title: Botany

text: Unlike animals, many plant cells, particularly those of the parenchyma, do not terminally differentiate, remaining totipotent with the ability to give rise to a new individual plant. Exceptions include highly lignified cells, the sclerenchyma and xylem which are dead at maturity, and the phloem sieve tubes which lack nuclei. While plants use many of the same epigenetic mechanisms as animals, such as chromatin remodelling, an alternative hypothesis is that plants set their gene expression patterns using positional information from the environment and surrounding cells to determine their developmental fate.


title: Botany

text: Epigenetic changes can lead to paramutations, which do not follow the Mendelian heritage rules. These epigenetic marks are carried from one generation to the next, with one allele inducing a change on the other.


title: Botany

text: The chloroplasts of plants have a number of biochemical, structural and genetic similarities to cyanobacteria, (commonly but incorrectly known as "blue-green algae") and are thought to be derived from an ancient endosymbiotic relationship between an ancestral eukaryotic cell and a cyanobacterial resident.


title: Botany

text: The algae are a polyphyletic group and are placed in various divisions, some more closely related to plants than others. There are many differences between them in features such as cell wall composition, biochemistry, pigmentation, chloroplast structure and nutrient reserves. The algal division Charophyta, sister to the green algal division Chlorophyta, is considered to contain the ancestor of true plants. The Charophyte class Charophyceae and the land plant sub-kingdom Embryophyta together form the monophyletic group or clade Streptophytina.


title: Botany

text: Nonvascular land plants are embryophytes that lack the vascular tissues xylem and phloem. They include mosses, liverworts and hornworts. Pteridophytic vascular plants with true xylem and phloem that reproduced by spores germinating into free-living gametophytes evolved during the Silurian period and diversified into several lineages during the late Silurian and early Devonian. Representatives of the lycopods have survived to the present day. By the end of the Devonian period, several groups, including the lycopods, sphenophylls and progymnosperms, had independently evolved "megaspory" – their spores were of two distinct sizes, larger megaspores and smaller microspores. Their reduced gametophytes developed from megaspores retained within the spore-producing organs (megasporangia) of the sporophyte, a condition known as endospory. Seeds consist of an endosporic megasporangium surrounded by one or two sheathing layers (integuments). The young sporophyte develops within the seed, which on germination splits to release it. The earliest known seed plants date from the latest Devonian Famennian stage. Following the evolution of the seed habit, seed plants diversified, giving rise to a number of now-extinct groups, including seed ferns, as well as the modern gymnosperms and angiosperms. Gymnosperms produce "naked seeds" not fully enclosed in an ovary; modern representatives include conifers, cycads, "Ginkgo", and Gnetales. Angiosperms produce seeds enclosed in a structure such as a carpel or an ovary. Ongoing research on the molecular phylogenetics of living plants appears to show that the angiosperms are a sister clade to the gymnosperms.


title: Botany

text: Plant physiology encompasses all the internal chemical and physical activities of plants associated with life. Chemicals obtained from the air, soil and water form the basis of all plant metabolism. The energy of sunlight, captured by oxygenic photosynthesis and released by cellular respiration, is the basis of almost all life. Photoautotrophs, including all green plants, algae and cyanobacteria gather energy directly from sunlight by photosynthesis. Heterotrophs including all animals, all fungi, all completely parasitic plants, and non-photosynthetic bacteria take in organic molecules produced by photoautotrophs and respire them or use them in the construction of cells and tissues. Respiration is the oxidation of carbon compounds by breaking them down into simpler structures to release the energy they contain, essentially the opposite of photosynthesis.


title: Botany

text: Molecules are moved within plants by transport processes that operate at a variety of spatial scales. Subcellular transport of ions, electrons and molecules such as water and enzymes occurs across cell membranes. Minerals and water are transported from roots to other parts of the plant in the transpiration stream. Diffusion, osmosis, and active transport and mass flow are all different ways transport can occur. Examples of elements that plants need to transport are nitrogen, phosphorus, potassium, calcium, magnesium, and sulfur. In vascular plants, these elements are extracted from the soil as soluble ions by the roots and transported throughout the plant in the xylem. Most of the elements required for plant nutrition come from the chemical breakdown of soil minerals. Sucrose produced by photosynthesis is transported from the leaves to other parts of the plant in the phloem and plant hormones are transported by a variety of processes.


title: Botany

text: Plants are not passive, but respond to external signals such as light, touch, and injury by moving or growing towards or away from the stimulus, as appropriate. Tangible evidence of touch sensitivity is the almost instantaneous collapse of leaflets of "Mimosa pudica", the insect traps of Venus flytrap and bladderworts, and the pollinia of orchids.


title: Botany

text: The hypothesis that plant growth and development is coordinated by plant hormones or plant growth regulators first emerged in the late 19th century. Darwin experimented on the movements of plant shoots and roots towards light and gravity, and concluded "It is hardly an exaggeration to say that the tip of the radicle . . acts like the brain of one of the lower animals . . directing the several movements". About the same time, the role of auxins (from the Greek , to grow) in control of plant growth was first outlined by the Dutch scientist Frits Went. The first known auxin, indole-3-acetic acid (IAA), which promotes cell growth, was only isolated from plants about 50 years later. This compound mediates the tropic responses of shoots and roots towards light and gravity. The finding in 1939 that plant callus could be maintained in culture containing IAA, followed by the observation in 1947 that it could be induced to form roots and shoots by controlling the concentration of growth hormones were key steps in the development of plant biotechnology and genetic modification.


title: Botany

text: Cytokinins are a class of plant hormones named for their control of cell division (especially cytokinesis). The natural cytokinin zeatin was discovered in corn, "Zea mays", and is a derivative of the purine adenine. Zeatin is produced in roots and transported to shoots in the xylem where it promotes cell division, bud development, and the greening of chloroplasts. The gibberelins, such as gibberelic acid are diterpenes synthesised from acetyl CoA via the mevalonate pathway. They are involved in the promotion of germination and dormancy-breaking in seeds, in regulation of plant height by controlling stem elongation and the control of flowering. Abscisic acid (ABA) occurs in all land plants except liverworts, and is synthesised from carotenoids in the chloroplasts and other plastids. It inhibits cell division, promotes seed maturation, and dormancy, and promotes stomatal closure. It was so named because it was originally thought to control abscission. Ethylene is a gaseous hormone that is produced in all higher plant tissues from methionine. It is now known to be the hormone that stimulates or regulates fruit ripening and abscission, and it, or the synthetic growth regulator ethephon which is rapidly metabolised to produce ethylene, are used on industrial scale to promote ripening of cotton, pineapples and other climacteric crops.


title: Botany

text: Another class of phytohormones is the jasmonates, first isolated from the oil of "Jasminum grandiflorum" which regulates wound responses in plants by unblocking the expression of genes required in the systemic acquired resistance response to pathogen attack.


title: Botany

text: In addition to being the primary energy source for plants, light functions as a signalling device, providing information to the plant, such as how much sunlight the plant receives each day. This can result in adaptive changes in a process known as photomorphogenesis. Phytochromes are the photoreceptors in a plant that are sensitive to light.


title: Botany

text: Plant anatomy is the study of the structure of plant cells and tissues, whereas plant morphology is the study of their external form.


title: Botany

text: All plants are multicellular eukaryotes, their DNA stored in nuclei. The characteristic features of plant cells that distinguish them from those of animals and fungi include a primary cell wall composed of the polysaccharides cellulose, hemicellulose and pectin, larger vacuoles than in animal cells and the presence of plastids with unique photosynthetic and biosynthetic functions as in the chloroplasts. Other plastids contain storage products such as starch (amyloplasts) or lipids (elaioplasts). Uniquely, streptophyte cells and those of the green algal order Trentepohliales divide by construction of a phragmoplast as a template for building a cell plate late in cell division.


title: Botany

text: The bodies of vascular plants including clubmosses, ferns and seed plants (gymnosperms and angiosperms) generally have aerial and subterranean subsystems. The shoots consist of stems bearing green photosynthesising leaves and reproductive structures. The underground vascularised roots bear root hairs at their tips and generally lack chlorophyll. Non-vascular plants, the liverworts, hornworts and mosses do not produce ground-penetrating vascular roots and most of the plant participates in photosynthesis. The sporophyte generation is nonphotosynthetic in liverworts but may be able to contribute part of its energy needs by photosynthesis in mosses and hornworts.


title: Botany

text: The root system and the shoot system are interdependent – the usually nonphotosynthetic root system depends on the shoot system for food, and the usually photosynthetic shoot system depends on water and minerals from the root system. Cells in each system are capable of creating cells of the other and producing adventitious shoots or roots. Stolons and tubers are examples of shoots that can grow roots. Roots that spread out close to the surface, such as those of willows, can produce shoots and ultimately new plants. In the event that one of the systems is lost, the other can often regrow it. In fact it is possible to grow an entire plant from a single leaf, as is the case with plants in "Streptocarpus" sect. "Saintpaulia", or even a single cell – which can dedifferentiate into a callus (a mass of unspecialised cells) that can grow into a new plant.


title: Botany

text: In vascular plants, the xylem and phloem are the conductive tissues that transport resources between shoots and roots. Roots are often adapted to store food such as sugars or starch, as in sugar beets and carrots.


title: Botany

text: Stems mainly provide support to the leaves and reproductive structures, but can store water in succulent plants such as cacti, food as in potato tubers, or reproduce vegetatively as in the stolons of strawberry plants or in the process of layering. Leaves gather sunlight and carry out photosynthesis. Large, flat, flexible, green leaves are called foliage leaves. Gymnosperms, such as conifers, cycads, "Ginkgo", and gnetophytes are seed-producing plants with open seeds. Angiosperms are seed-producing plants that produce flowers and have enclosed seeds. Woody plants, such as azaleas and oaks, undergo a secondary growth phase resulting in two additional types of tissues: wood (secondary xylem) and bark (secondary phloem and cork). All gymnosperms and many angiosperms are woody plants. Some plants reproduce sexually, some asexually, and some via both means.


title: Botany

text: Although reference to major morphological categories such as root, stem, leaf, and trichome are useful, one has to keep in mind that these categories are linked through intermediate forms so that a continuum between the categories results. Furthermore, structures can be seen as processes, that is, process combinations.


title: Botany

text: Systematic botany is part of systematic biology, which is concerned with the range and diversity of organisms and their relationships, particularly as determined by their evolutionary history. It involves, or is related to, biological classification, scientific taxonomy and phylogenetics. Biological classification is the method by which botanists group organisms into categories such as genera or species. Biological classification is a form of scientific taxonomy. Modern taxonomy is rooted in the work of Carl Linnaeus, who grouped species according to shared physical characteristics. These groupings have since been revised to align better with the Darwinian principle of common descent – grouping organisms by ancestry rather than superficial characteristics. While scientists do not always agree on how to classify organisms, molecular phylogenetics, which uses DNA sequences as data, has driven many recent revisions along evolutionary lines and is likely to continue to do so. The dominant classification system is called Linnaean taxonomy. It includes ranks and binomial nomenclature. The nomenclature of botanical organisms is codified in the International Code of Nomenclature for algae, fungi, and plants (ICN) and administered by the International Botanical Congress.


title: Botany

text: Kingdom Plantae belongs to Domain Eukaryota and is broken down recursively until each species is separately classified. The order is: Kingdom; Phylum (or Division); Class; Order; Family; Genus (plural "genera"); Species. The scientific name of a plant represents its genus and its species within the genus, resulting in a single worldwide name for each organism. For example, the tiger lily is "Lilium columbianum". "Lilium" is the genus, and "columbianum" the specific epithet. The combination is the name of the species. When writing the scientific name of an organism, it is proper to capitalise the first letter in the genus and put all of the specific epithet in lowercase. Additionally, the entire term is ordinarily italicised (or underlined when italics are not available).


title: Botany

text: The evolutionary relationships and heredity of a group of organisms is called its phylogeny. Phylogenetic studies attempt to discover phylogenies. The basic approach is to use similarities based on shared inheritance to determine relationships. As an example, species of "Pereskia" are trees or bushes with prominent leaves. They do not obviously resemble a typical leafless cactus such as an "Echinocactus". However, both "Pereskia" and "Echinocactus" have spines produced from areoles (highly specialised pad-like structures) suggesting that the two genera are indeed related.


title: Botany

text: Judging relationships based on shared characters requires care, since plants may resemble one another through convergent evolution in which characters have arisen independently. Some euphorbias have leafless, rounded bodies adapted to water conservation similar to those of globular cacti, but characters such as the structure of their flowers make it clear that the two groups are not closely related. The cladistic method takes a systematic approach to characters, distinguishing between those that carry no information about shared evolutionary history – such as those evolved separately in different groups (homoplasies) or those left over from ancestors (plesiomorphies) – and derived characters, which have been passed down from innovations in a shared ancestor (apomorphies). Only derived characters, such as the spine-producing areoles of cacti, provide evidence for descent from a common ancestor. The results of cladistic analyses are expressed as cladograms: tree-like diagrams showing the pattern of evolutionary branching and descent.


title: Botany

text: From the 1990s onwards, the predominant approach to constructing phylogenies for living plants has been molecular phylogenetics, which uses molecular characters, particularly DNA sequences, rather than morphological characters like the presence or absence of spines and areoles. The difference is that the genetic code itself is used to decide evolutionary relationships, instead of being used indirectly via the characters it gives rise to. Clive Stace describes this as having "direct access to the genetic basis of evolution." As a simple example, prior to the use of genetic evidence, fungi were thought either to be plants or to be more closely related to plants than animals. Genetic evidence suggests that the true evolutionary relationship of multicelled organisms is as shown in the cladogram below – fungi are more closely related to animals than to plants.


title: Botany

text: In 1998, the Angiosperm Phylogeny Group published a phylogeny for flowering plants based on an analysis of DNA sequences from most families of flowering plants. As a result of this work, many questions, such as which families represent the earliest branches of angiosperms, have now been answered. Investigating how plant species are related to each other allows botanists to better understand the process of evolution in plants. Despite the study of model plants and increasing use of DNA evidence, there is ongoing work and discussion among taxonomists about how best to classify plants into various taxa. Technological developments such as computers and electron microscopes have greatly increased the level of detail studied and speed at which data can be analysed.


title: Botany

text: A few symbols are in current use in botany. A number of others are obsolete; for example, Linnaeus used planetary symbols for woody, herbaceous and perennial plants, and Willd used (Saturn) for neuter in addition to (Mercury) for hermaphroditic. The following symbols are still used:


title: Leonardo Bonucci

text: Leonardo Bonucci (; born 1 May 1987) is an Italian professional footballer who plays as a centre-back and captains both club Juventus and the Italy national team. Known for his technique and ball-playing ability, he is regarded as one of the best defenders of his generation.


title: Leonardo Bonucci

text: After beginning his career with Inter Milan in 2005, Bonucci spent the next few seasons on loan at Treviso and Pisa, before moving to Bari in 2009. His defensive performances alongside fellow Italian centre-back Andrea Ranocchia earned him a move to Juventus the following season, where he later became a key member of the club's three-man defensive line, alongside Giorgio Chiellini and Andrea Barzagli, establishing himself as one of the best defenders in world football. Among other titles, he went on to win six consecutive Serie A titles with the team between 2012 and 2017, having also played two UEFA Champions League finals in 2015 and 2017. In 2017, he moved to AC Milan, and one season later returned to Juventus, winning two more consecutive league titles.


title: Leonardo Bonucci

text: At the international level, Bonucci has earned 120 caps since his senior debut in 2010, representing Italy at two FIFA World Cups (2010 and 2014), three European Championships, (2012, 2016, and 2020), and a FIFA Confederations Cup (2013); he won Euro 2020 and earned a runners-up medal at Euro 2012 and a third-place medal at the 2013 Confederations Cup.


title: Leonardo Bonucci

text: Bonucci has also won several individual honours for his performances: he was named to the UEFA Europa League Squad of the season during the 2013–14 and 2017–18 seasons, and is a four-time member of the Serie A Team of the Year. He was named the Serie A Footballer of the Year in 2016, and was also included in the UEFA Team of the Year in the same season. In 2017, he was also included in the FIFA FIFPro World11 and the IFFHS Men's World Team, as well as the 2016–17 UEFA Champions League Team of the Season and the 2016–17 ESM Team of the Year.


title: Leonardo Bonucci

text: Bonucci started his career in the youth ranks of his hometown club Viterbese but was loaned to Inter Milan in the summer of 2005. He played a number of pre-season friendlies for the Inter first team. He then became a member of the Inter U20 team.


title: Leonardo Bonucci

text: On 14 May 2006, Bonucci made his Serie A debut in the last match of the 2005–06 season, in a 2–2 away draw against Cagliari, which was Inter's 3,500th competitive match. On 7 July 2006, Inter bought Bonucci outright.


title: Leonardo Bonucci

text: He played his first Coppa Italia match against Messina on 9 November 2006 when he came off the bench for Fabio Grosso in the 86th minute. Bonucci featured in two more Coppa Italia games for Inter that season when he was brought on for the substituted Walter Samuel at half-time during the quarter final second leg match against Empoli, and as a starter in the semi final second leg tie against Sampdoria.


title: Leonardo Bonucci

text: In January 2007, Inter sold 50% of Bonucci's registration rights to Treviso, along with 50% of the registration rights for fellow Primavera team-mate Daniel Maa Boumsong. At that time Bonucci was tagged for a peppercorn fee of €500. Bonucci subsequently remained at Inter until 30 June 2007 while Maa Boumsong returned from Treviso where he spent the first half of the season on loan. During Bonucci's last season with the Inter's youth side, he won the Campionato Nazionale Primavera (the national youth league title).


title: Leonardo Bonucci

text: On 1 July 2007, Bonucci and Maa Boumsong formally became players of Treviso after their loan contract back to Inter had expired, as well as the renewal of the co-ownerships in June 2007. At Treviso, Bonucci made 20 starts in 27 Serie B appearances as one of the regular starters.


title: Leonardo Bonucci

text: In June 2008, among the other Inter youth products, Bonucci was the only player that was bought back from Treviso. However, he was loaned back to Treviso for the 2008–09 season. According to a Treviso filing named "Tabella N°5 – circolare Co.Vi.So.C. prot. N°4051.4/GC/pc del 11 maggio 2005" in their 2007–08 financial report, Bonucci was sold for a €700,000 fee.


title: Leonardo Bonucci

text: Bonucci played 13 Serie B matches for Treviso before leaving for another Serie B struggler Pisa on loan.


title: Leonardo Bonucci

text: On 8 June 2009, Bonucci underwent a medical examination at Genoa. On 1 July, Inter officially announced that Bonucci, along with Acquafresca, Bolzoni and Meggiorini had been transferred to Genoa, as part of the deal that sent Diego Milito and Thiago Motta to Inter. Moreover, Ivan Fatić who was co-contracted ("co-owned") between Chievo and Inter, became co-contracted between Chievo and Genoa instead, according to a news article by "La Gazzetta dello Sport". Bonucci was valued at €3 million at that time.


title: Leonardo Bonucci

text: On 2 July, he was transferred to Bari from Genoa, on a co-ownership deal, for €1.75 million, along with Meggiorini (also on a co-ownership deal), Matteo Paro (on loan), Andrea Ranocchia (on loan) and Giuseppe Greco (on loan).