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124-527: Fertilisation or fertilization (see spelling differences ), also known as generative fertilisation , syngamy and impregnation , is the fusion of gametes to give rise to a zygote and initiate its development into a new individual organism or offspring. While processes such as insemination or pollination , which happen before the fusion of gametes, are also sometimes informally referred to as fertilisation, these are technically separate processes. The cycle of fertilisation and development of new individuals
248-691: A consonant followed by an unstressed -re (pronounced /ə(r)/ ). In modern American English, most of these words have the ending -er . The difference is most common for words ending in -bre or -tre : British spellings calibre , centre , fibre , goitre , litre , lustre , manoeuvre , meagre , metre (length) , mitre , nitre , ochre , reconnoitre , sabre , saltpetre , sepulchre , sombre , spectre , theatre (see exceptions ) and titre all have -er in American spelling. In Britain, both -re and -er spellings were common before Johnson's 1755 dictionary
372-475: A distinctive set of Canadian English spellings is viewed by many Canadians as one of the unique aspects of Canadian culture (especially when compared to the United States). In Australia, -or endings enjoyed some use throughout the 19th century and in the early 20th century. Like Canada, though, most major Australian newspapers have switched from " -or " endings to " -our " endings. The " -our " spelling
496-495: A emium , and a enigma . In others, it is kept in all varieties: for example, phoenix , and usually subpoena , but Phenix in Virginia . This is especially true of names: Aegean (the sea), Caesar , Oedipus , Phoebe , etc., although "caesarean section" may be spelled as "cesarean section". There is no reduction of Latin -ae plurals (e.g., larv ae ); nor where the digraph <ae>/<oe> does not result from
620-466: A eon , an a emia , an a esthesia , c a ecum , c a esium , c o eliac , diarrh o ea , encyclop a edia , f a eces , f o etal , gyn a ecology , h a emoglobin , h a emophilia , leuk a emia , o esophagus , o estrogen , orthop a edic , pal a eontology , p a ediatric , p a edophile . Oenology is acceptable in American English but is deemed
744-479: A family of glycoproteins called TTS proteins that enhanced growth of pollen tubes. Pollen tubes in a sugar free pollen germination medium and a medium with purified TTS proteins both grew. However, in the TTS medium, the tubes grew at a rate 3x that of the sugar-free medium. TTS proteins were also placed on various locations of semi in vivo pollinated pistils, and pollen tubes were observed to immediately extend toward
868-532: A minor variant of enology , whereas although archeology and ameba exist in American English, the British versions amoeba and archaeology are more common. The chemical haem (named as a shortening of h a emoglobin ) is spelled heme in American English, to avoid confusion with hem . Canadian English mostly follows American English in this respect, although it is split on gynecology (e.g. Society of Obstetricians and Gynaecologists of Canada vs.
992-569: A mistaken etymology. The etymologically correct original spelling fetus reflects the Latin original and is the standard spelling in medical journals worldwide; the Oxford English Dictionary notes that "In Latin manuscripts both fētus and foetus are used". The Ancient Greek diphthongs <αι> and <οι> were transliterated into Latin as <ae> and <oe>. The ligatures æ and œ were introduced when
1116-454: A mode called by him as epigenetic . In 1784, Spallanzani established the need of interaction between the female's ovum and male's sperm to form a zygote in frogs. In 1827, Karl Ernst von Baer observed a therian mammalian egg for the first time. Oscar Hertwig (1876), in Germany, described the fusion of nuclei of spermatozoa and of ova from sea urchin . The evolution of fertilisation
1240-515: A short period lasting some days; a queen may mate with eight or more drones . She then stores the sperm for the rest of her life, perhaps for five years or more. In many fungi (except chytrids ), as in some protists, fertilisation is a two step process. First, the cytoplasms of the two gamete cells fuse (called plasmogamy ), producing a dikaryotic or heterokaryotic cell with multiple nuclei. This cell may then divide to produce dikaryotic or heterokaryotic hyphae . The second step of fertilisation
1364-507: A study suggested that self-fertilisation evolved roughly a million years ago or more in A. thaliana . In long-established self-fertilising plants, the masking of deleterious mutations and the production of genetic variability is infrequent and thus unlikely to provide a sufficient benefit over many generations to maintain the meiotic apparatus. Consequently, one might expect self-fertilisation to be replaced in nature by an ameiotic asexual form of reproduction that would be less costly. However
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#17327718352401488-567: A subject of semantic arguments about the beginning of pregnancy , typically in the context of the abortion debate. Upon gastrulation , which occurs around 16 days after fertilisation, the implanted blastocyst develops three germ layers, the endoderm, the ectoderm and the mesoderm, and the genetic code of the father becomes fully involved in the development of the embryo; later twinning is impossible. Additionally, interspecies hybrids survive only until gastrulation and cannot further develop. However, some human developmental biology literature refers to
1612-667: A suffix for agentive ( reader , user , winner ) and comparative ( louder , nicer ) forms. One outcome is the British distinction of meter for a measuring instrument from metre for the unit of length. However, while " poetic metre " is often spelled as -re , pentameter , hexameter , etc. are always -er . Many other words have -er in British English. These include Germanic words, such as anger , mother , timber and water , and such Romance-derived words as danger , quarter and river . The ending -cre , as in acre , lucre , massacre , and mediocre ,
1736-400: A thin spike at the head of the sperm called the acrosomal process . The sperm binds to the egg through another ligand reaction between receptors on the vitelline membrane . The sperm surface protein bindin, binds to a receptor on the vitelline membrane identified as EBR1. Fusion of the plasma membranes of the sperm and egg are likely mediated by bindin. At the site of contact, fusion causes
1860-447: Is karyogamy , the fusion of the nuclei to form a diploid zygote. In chytrid fungi, fertilisation occurs in a single step with the fusion of gametes, as in animals and plants. There are three types of fertilisation processes in protozoa: Algae, like some land plants, undergo alternation of generations . Some algae are isomorphic, where both the sporophyte (2n) and gameteophyte (n) are the same morphologically. When algae reproduction
1984-432: Is - /s/ for the noun and - /z/ for the verb). For licence / license or practice / practise , British English also keeps the noun–verb distinction graphically (although phonetically the two words in each pair are homophones with - /s/ pronunciation). On the other hand, American English uses license and practice for both nouns and verbs (with - /s/ pronunciation in both cases too). American English has kept
2108-434: Is a curated online information source for Arabidopsis thaliana genetic and molecular biology research, and The Arabidopsis Book is an online compilation of invited chapters on Arabidopsis thaliana biology. (Note that as of 2013 no further chapters will be published.) In Europe, the model organism resource centre for Arabidopsis thaliana germplasm , bioinformatics and molecular biology resources (including GeneChips )
2232-406: Is a genus in the family Brassicaceae . They are small flowering plants related to cabbage and mustard . This genus is of great interest since it contains thale cress ( Arabidopsis thaliana ), one of the model organisms used for studying plant biology and the first plant to have its entire genome sequenced. Changes in thale cress are easily observed, making it a very useful model. Currently,
2356-607: Is also estimated that about 42% of flowering plants exhibit a mixed mating system in nature. In the most common kind of mixed mating system, individual plants produce a single type of flower and fruits may contain self-fertilised, outcrossed or a mixture of progeny types. The transition from cross-fertilisation to self-fertilisation is the most common evolutionary transition in plants, and has occurred repeatedly in many independent lineages. About 10-15% of flowering plants are predominantly self-fertilising. Under circumstances where pollinators or mates are rare, self-fertilisation offers
2480-399: Is called sexual reproduction . During double fertilisation in angiosperms , the haploid male gamete combines with two haploid polar nuclei to form a triploid primary endosperm nucleus by the process of vegetative fertilisation. In antiquity, Aristotle conceived the formation of new individuals through fusion of male and female fluids, with form and function emerging gradually, in
2604-402: Is described as oogamous, the male and female gametes are different morphologically, where there is a large non-motile egg for female gametes, and the male gamete are uniflagellate (motile). Via the process of syngamy, these will form a new zygote, regenerating the sporophyte generation again. Meiosis results in a random segregation of the genes that each parent contributes. Each parent organism
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#17327718352402728-404: Is diploid, resulting from the fusion of a sperm with one, rather than two, maternal nuclei. It is believed that early in the development of angiosperm lineages, there was a duplication in this mode of reproduction, producing seven-celled/eight-nucleate female gametophytes, and triploid endosperms with a 2:1 maternal to paternal genome ratio. In many plants, the development of the flesh of the fruit
2852-399: Is dropped for other derivations, for example, central , fibrous , spectral . However, the existence of related words without e before the r is not proof for the existence of an -re British spelling: for example, entry and entrance come from enter , which has not been spelled entre for centuries. The difference relates only to root words; -er rather than -re is universal as
2976-502: Is generally preferred over oe and often over ae , but oe and ae are sometimes found in academic and scientific writing as well as government publications (for example, the fee schedule of the Ontario Health Insurance Plan ) and some words such as palaeontology or aeon . In Australia, it can go either way, depending on the word: for instance, medieval is spelled with the e rather than ae , following
3100-601: Is known as the Pacific Parlour car, not Pacific Parlor . Proper names such as Pearl Harbor or Sydney Harbour are usually spelled according to their native-variety spelling vocabulary. The name of the herb savory is spelled thus everywhere, although the related adjective savo(u)ry , like savo(u)r , has a u in the UK. Honor (the name) and arbor (the tool) have -or in Britain, as mentioned above, as does
3224-404: Is more common for ejaculation to precede ovulation than vice versa. When sperm are deposited into the anterior vagina, they are not capable of fertilisation (i.e., non-capacitated) and are characterised by slow linear motility patterns. This motility, combined with muscular contractions enables sperm transport towards the uterus and oviducts . There is a pH gradient within the micro-environment of
3348-471: Is proportional to the percentage of fertilised ovules. For example, with watermelon , about a thousand grains of pollen must be delivered and spread evenly on the three lobes of the stigma to make a normal sized and shaped fruit. Outcrossing , or cross-fertilisation, and self-fertilisation represent different strategies with differing benefits and costs. An estimated 48.7% of plant species are either dioecious or self-incompatible obligate outcrossers. It
3472-492: Is related to the origin of meiosis , as both are part of sexual reproduction , originated in eukaryotes . One hypothesis states that meiosis originated from mitosis. The gametes that participate in fertilisation of plants are the sperm (male) and the egg (female) cell. Various plant groups have differing methods by which the gametes produced by the male and female gametophytes come together and are fertilised. In bryophytes and pteridophytic land plants, fertilisation of
3596-529: Is similar to the role of the vitelline membrane in sea urchins, binds the sperm. Unlike sea urchins, the sperm binds to the egg before the acrosomal reaction. ZP3 , a glycoprotein in the zona pellucida, is responsible for egg/sperm adhesion in humans. The receptor galactosyltransferase (GalT) binds to the N-acetylglucosamine residues on the ZP3 and is important for binding with the sperm and activating
3720-613: Is sometimes used. The ratio between -ise and -ize stood at 3:2 in the British National Corpus up to 2002. The spelling -ise is more commonly used in UK mass media and newspapers, including The Times (which switched conventions in 1992), The Daily Telegraph , The Economist and the BBC . The Government of the United Kingdom additionally uses -ise , stating "do not use Americanisms" justifying that
3844-714: Is standard worldwide and complection is rare. However, the adjective complected (as in "dark-complected"), although sometimes proscribed, is on equal ground in the U.S. with complexioned. It is not used in this way in the UK, although there exists a rare alternative meaning of complicated . In some cases, words with "old-fashioned" spellings are retained widely in the U.S. for historical reasons (cf. connexionalism ). Many words, especially medical words, that are written with ae/æ or oe/œ in British English are written with just an e in American English. The sounds in question are /iː/ or /ɛ/ (or, unstressed, /i/ , /ɪ/ or /ə/ ). Examples (with non-American letter in bold ):
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3968-607: Is taught in schools nationwide as part of the Australian curriculum. The most notable countrywide use of the -or ending is for one of the country's major political parties, the Australian Labor Party , which was originally called "the Australian Labour Party" (name adopted in 1908), but was frequently referred to as both "Labour" and "Labor". The "Labor" was adopted from 1912 onward due to
4092-878: Is the Nottingham Arabidopsis Stock Centre (NASC) whilst in North America germplasm services are provided by the Arabidopsis Biological Resource Center (ABRC) based at Ohio State University . The ordering system for ABRC was incorporated into the TAIR database in June 2001 whilst NASC has always (since 1991) hosted its own ordering system and genome browser. In 1982, the crew of the Soviet Salyut 7 space station grew some Arabidopsis, thus becoming
4216-549: Is the usual form of the male given name, as a surname both the spellings Peter and Petre (the latter notably borne by a British lord ) are found. For British accoutre , the American practice varies: the Merriam-Webster Dictionary prefers the -re spelling, but The American Heritage Dictionary of the English Language prefers the -er spelling. More recent French loanwords keep
4340-484: Is used in both British and American English to show that the c is pronounced /k/ rather than /s/ . The spellings euchre and ogre are also the same in both British and American English. Fire and its associated adjective fiery are the same in both British and American English, although the noun was spelled fier in Old and Middle English. Theater is the prevailing American spelling used to refer to both
4464-598: Is usually identical save for a fraction of their genes; each gamete is therefore genetically unique. At fertilisation, parental chromosomes combine. In humans , (2²²)² = 17.6x10 chromosomally different zygotes are possible for the non-sex chromosomes, even assuming no chromosomal crossover . If crossover occurs once, then on average (4²²)² = 309x10 genetically different zygotes are possible for every couple, not considering that crossover events can take place at most points along each chromosome. The X and Y chromosomes undergo no crossover events and are therefore excluded from
4588-591: The -meter suffix (from Ancient Greek - μέτρον métron , via French -mètre ) normally had the -re spelling from earliest use in English but were superseded by -er . Examples include thermometer and barometer . The e preceding the r is kept in American-inflected forms of nouns and verbs, for example, fibers , reconnoitered , centering , which are fibres , reconnoitred , and centring respectively in British English. According to
4712-558: The conceptus and such medical literature refers to the "products of conception" as the post-implantation embryo and its surrounding membranes. The term "conception" is not usually used in scientific literature because of its variable definition and connotation. Insects in different groups, including the Odonata ( dragonflies and damselflies ) and the Hymenoptera ( ants , bees , and wasps ) practise delayed fertilisation. Among
4836-560: The -ise form as an alternative. Publications by Oxford University Press (OUP)—such as Henry Watson Fowler 's A Dictionary of Modern English Usage , Hart's Rules , and The Oxford Guide to English Usage —also recommend -ize . However, Robert Allan's Pocket Fowler's Modern English Usage considers either spelling to be acceptable anywhere but the U.S. American spelling avoids -ise endings in words like organize , realize and recognize . British spelling mostly uses -ise ( organise , realise , recognise ), though -ize
4960-687: The -ise form is preferred in Australian English at a ratio of about 3:1 according to the Macquarie Dictionary . In Canada, the -ize ending is more common, although the Ontario Public School Spelling Book spelled most words in the -ize form, but allowed for duality with a page insert as late as the 1970s, noting that, although the -ize spelling was in fact the convention used in the OED ,
5084-439: The -ise variation, once more common amongst older Canadians, is employed less and less often in favour of the -ize spelling. (The alternate convention offered as a matter of choice may have been due to the fact that although there were an increasing number of American- and British-based dictionaries with Canadian Editions by the late 1970s, these were largely only supplemental in terms of vocabulary with subsequent definitions. It
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5208-450: The -re spelling in American English. These are not exceptions when a French-style pronunciation is used ( /rə/ rather than /ə(r)/ ), as with double entendre , genre and oeuvre . However, the unstressed /ə(r)/ pronunciation of an -er ending is used more (or less) often with some words, including cadre , macabre , maître d' , Notre Dame , piastre , and timbre . The -re endings are mostly standard throughout
5332-495: The Canadian Medical Association 's Canadian specialty profile of Obstetrics/gynecology ). Pediatrician is preferred roughly 10 to 1 over paediatrician , while foetal and oestrogen are similarly uncommon. Words that can be spelled either way in American English include a esthetics and arch a eology (which usually prevail over esthetics and archeology ), as well as pal a estra , for which
5456-548: The OED , centring is a "word ... of 3 syllables (in careful pronunciation)" (i.e., /ˈsɛntərɪŋ/ ), yet there is no vowel in the spelling corresponding to the second syllable ( /ə/ ). The OED third edition (revised entry of June 2016) allows either two or three syllables. On the Oxford Dictionaries Online website, the three-syllable version is listed only as the American pronunciation of centering . The e
5580-740: The Organisation for Economic Co-operation and Development ). The European Union 's style guides require the usage of - ise . Proofreaders at the EU's Publications Office ensure consistent spelling in official publications such as the Official Journal of the European Union (where legislation and other official documents are published), but the -ize spelling may be found in other documents. Arabidopsis See text Cardaminopsis (C.A.Mey.) Hayek Arabidopsis ( rockcress )
5704-557: The Simplified Spelling Board in the early 20th century, but most were not adopted. In Britain, the influence of those who preferred the Norman (or Anglo-French ) spellings of words proved to be decisive. Later spelling adjustments in the United Kingdom had little effect on today's American spellings and vice versa. For the most part, the spelling systems of most Commonwealth countries and Ireland closely resemble
5828-432: The endosperm , a nutrient -rich tissue , inside the seed . The two central-cell maternal nuclei (polar nuclei) that contribute to the endosperm arise by mitosis from the single meiotic product that also gave rise to the egg. Therefore, maternal contribution to the genetic constitution of the triploid endosperm is double that of the embryo. One primitive species of flowering plant, Nuphar polysepala , has endosperm that
5952-543: The theatre spelling. (The word "theater" in American English is a place where both stage performances and screenings of films take place, but in British English a "theatre" is where stage performances take place but not film screenings – these take place in a cinema, or "picture theatre" in Australia.) In the United States, the spelling theatre is sometimes used when referring to the art form of theatre, while
6076-519: The u has since been dropped: ambassadour , emperour , errour , governour , horrour , inferiour , mirrour , perturbatour , superiour , tenour , terrour , tremour . Johnson, unlike Webster, was not an advocate of spelling reform, but chose the spelling best derived, as he saw it, from among the variations in his sources. He preferred French over Latin spellings because, as he put it, "the French generally supplied us". English speakers who moved to
6200-435: The u : In American usage, derivatives and inflected forms are built by simply adding the suffix in all cases (for example, favorite , savory etc.) since the u is absent to begin with. American usage, in most cases, keeps the u in the word glamour , which comes from Scots , not Latin or French. Glamor is sometimes used in imitation of the spelling reform of other -our words to -or . Nevertheless,
6324-652: The ‑our/or group do not have a Latin counterpart that ends in ‑or ; for example, armo(u)r , behavio(u)r , harbo(u)r , neighbo(u)r ; also arbo(u)r , meaning "shelter", though senses "tree" and "tool" are always arbor , a false cognate of the other word. The word arbor would be more accurately spelled arber or arbre in the US and the UK, respectively, the latter of which is the French word for "tree". Some 16th- and early 17th-century British scholars indeed insisted that ‑or be used for words from Latin (e.g., color ) and ‑our for French loans; however, in many cases,
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#17327718352406448-441: The 17th and 18th centuries, whereas there are thousands of examples of their -our counterparts. One notable exception is honor . Honor and honour were equally frequent in Britain until the 17th century; honor only exists in the UK now as the spelling of Honor Oak , a district of London, and of the occasional given name Honor . In derivatives and inflected forms of the -our/or words, British usage depends on
6572-615: The 1970s, but had by then been overtaken by connection in regular usage (for example, in more popular newspapers). Connexion (and its derivatives connexional and connexionalism ) is still in use by the Methodist Church of Great Britain to refer to the whole church as opposed to its constituent districts, circuits and local churches, whereas the US-majority United Methodist Church uses Connection . Complexion (which comes from complex )
6696-743: The American National Theatre was referred to by The New York Times as the "American National Theater ", but the organization uses "re" in the spelling of its name. The John F. Kennedy Center for the Performing Arts in Washington, D.C. has the more common American spelling theater in its references to the Eisenhower Theater, part of the Kennedy Center. Some cinemas outside New York also use
6820-480: The American and British varieties of English. However, English-language spelling reform has rarely been adopted otherwise. As a result, modern English orthography varies only minimally between countries and is far from phonemic in any country. In the early 18th century, English spelling was inconsistent. These differences became noticeable after the publication of influential dictionaries . Today's British English spellings mostly follow Johnson's A Dictionary of
6944-413: The American usage along with numerous other words such as eon or fetus , while other words such as oestrogen or paediatrician are spelled the British way. The Macquarie Dictionary also notes a growing tendency towards replacing ae and oe with e worldwide and with the exception of manoeuvre, all British or American spellings are acceptable variants. Elsewhere, the British usage prevails, but
7068-587: The Anglo-French spelling for defense and offense , which are defence and offence in British English. Likewise, there are the American pretense and British pretence ; but derivatives such as defensive , offensive , and pretension are always thus spelled in both systems. Australian and Canadian usages generally follow British usage. The spelling connexion is now rare in everyday British usage, its use lessening as knowledge of Latin attenuates, and it has almost never been used in
7192-440: The British system. In Canada, the spelling system can be said to follow both British and American forms, and Canadians are somewhat more tolerant of foreign spellings when compared with other English-speaking nationalities. Australian English mostly follows British spelling norms but has strayed slightly, with some American spellings incorporated as standard. New Zealand English is almost identical to British spelling, except in
7316-668: The British usage of -our . This coincided with a renewed interest in Canadian English, and the release of the updated Gage Canadian Dictionary in 1997 and the first Canadian Oxford Dictionary in 1998. Historically, most libraries and educational institutions in Canada have supported the use of the Oxford English Dictionary rather than the American Webster's Dictionary . Today, the use of
7440-471: The Commonwealth. The -er spellings are recognized as minor variants in Canada, partly due to United States influence. They are sometimes used in proper names (such as Toronto's controversially named Centerpoint Mall ). For advice / advise and device / devise , American English and British English both keep the noun–verb distinction both graphically and phonetically (where the pronunciation
7564-521: The English Language (1755), while many American English spellings follow Webster's An American Dictionary of the English Language ("ADEL", "Webster's Dictionary", 1828). Webster was a proponent of English spelling reform for reasons both philological and nationalistic. In A Companion to the American Revolution (2008), John Algeo notes: "it is often assumed that characteristically American spellings were invented by Noah Webster. He
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#17327718352407688-488: The Greek-style ligature as, for example, in maelstrom or toe ; the same is true for the British form aeroplane (compare other aero- words such as aerosol ) . The now chiefly North American airplane is not a respelling but a recoining, modelled after airship and aircraft . The word airplane dates from 1907, at which time the prefix aero- was trisyllabic, often written aëro- . In Canada, e
7812-505: The Odonata, females may mate with multiple males, and store sperm until the eggs are laid. The male may hover above the female during egg-laying (oviposition) to prevent her from mating with other males and replacing his sperm; in some groups such as the darters, the male continues to grasp the female with his claspers during egg-laying, the pair flying around in tandem. Among social Hymenoptera, honeybee queens mate only on mating flights, in
7936-518: The US: the more common connection has become the standard worldwide. According to the Oxford English Dictionary , the older spelling is more etymologically conservative, since the original Latin word had -xio- . The American usage comes from Webster , who abandoned -xion and preferred -ction . Connexion was still the house style of The Times of London until the 1980s and was still used by Post Office Telecommunications for its telephone services in
8060-497: The United States took these preferences with them. In the early 20th century, H. L. Mencken notes that " honor appears in the 1776 Declaration of Independence , but it seems to have been put there rather by accident than by design". In Jefferson 's original draft it is spelled "honour". In Britain, examples of behavior , color , flavor , harbor , and neighbor rarely appear in Old Bailey court records from
8184-423: The United States use Centre in their names. Examples include the villages of Newton Centre and Rockville Centre , the city of Centreville , Centre County and Centre College . Sometimes, these places were named before spelling changes but more often the spelling serves as an affectation. Proper names are usually spelled according to their native-variety spelling vocabulary; so, for instance, although Peter
8308-476: The United States. A "British standard" began to emerge following the 1755 publication of Samuel Johnson 's A Dictionary of the English Language , and an "American standard" started following the work of Noah Webster and, in particular, his An American Dictionary of the English Language , first published in 1828. Webster's efforts at spelling reform were effective in his native country, resulting in certain well-known patterns of spelling differences between
8432-468: The Vegetable Kingdom (pages 466-467) summed up his findings in the following way. "It has been shown in the present volume that the offspring from the union of two distinct individuals, especially if their progenitors have been subjected to very different conditions, have an immense advantage in height, weight, constitutional vigour and fertility over the self-fertilised offspring from one of
8556-467: The ability to store sperm for extended periods of time and can fertilise their eggs at their own desire. Oviparous animals producing eggs with thin tertiary membranes or no membranes at all, on the other hand, use external fertilisation methods. Such animals may be more precisely termed ovuliparous. External fertilisation is advantageous in that it minimises contact (which decreases the risk of disease transmission), and greater genetic variation. Sperm find
8680-440: The acrosome reaction. ZP3 is sufficient though unnecessary for sperm/egg binding. Two additional sperm receptors exist: a 250kD protein that binds to an oviduct secreted protein, and SED1, which independently binds to the zona. After the acrosome reaction, the sperm is believed to remain bound to the zona pellucida through exposed ZP2 receptors. These receptors are unknown in mice but have been identified in guinea pigs. In mammals,
8804-405: The actual persistence of meiosis and self-fertilisation as a form of reproduction in long-established self-fertilising plants may be related to the immediate benefit of efficient recombinational repair of DNA damage during formation of germ cells provided by meiosis at each generation. The mechanics behind fertilisation has been studied extensively in sea urchins and mice. This research addresses
8928-617: The adjective glamorous often drops the first "u". Saviour is a somewhat common variant of savior in the US. The British spelling is very common for honour (and favour ) in the formal language of wedding invitations in the US. The name of the Space Shuttle Endeavour has a u in it because the spacecraft was named after British Captain James Cook 's ship, HMS Endeavour . The (former) special car on Amtrak 's Coast Starlight train
9052-426: The advantage of reproductive assurance . Self-fertilisation can therefore result in improved colonisation ability. In some species, self-fertilisation has persisted over many generations. Capsella rubella is a self-fertilising species that became self-compatible 50,000 to 100,000 years ago. Arabidopsis thaliana is a predominantly self-fertilising plant with an out-crossing rate in the wild of less than 0.3%;
9176-466: The binding of the spermatozoon to the GalT initiates the acrosome reaction . This process releases the hyaluronidase that digests the matrix of hyaluronic acid in the vestments around the oocyte. Additionally, heparin-like glycosaminoglycans (GAGs) are released near the oocyte that promote the acrosome reaction. Fusion between the oocyte plasma membranes and sperm follows and allows the sperm nucleus ,
9300-518: The building itself, as noted above, generally is spelled theater . For example, the University of Wisconsin–Madison has a "Department of Theatre and Drama", which offers courses that lead to the "Bachelor of Arts in Theatre ", and whose professed aim is "to prepare our graduate students for successful 21st Century careers in the theatre both as practitioners and scholars". Some placenames in
9424-465: The calculation. The mitochondrial DNA is only inherited from the maternal parent. Shortly after the sperm fuse with the egg, the two sperm centrioles form the embryo first centrosome and microtubule aster . The sperm centriole, found near the male pronucleus, recruit egg Pericentriolar material proteins forming the zygote first centrosome. This centrosome nucleates microtubules in the shape of stars called astral microtubules. The microtubules span
9548-456: The choice to spell such words in the -ise form was a matter of personal preference ; however, a pupil having made the decision, one way or the other, thereafter ought to write uniformly not only for a given word, but to apply that same uniformity consistently for all words where the option is found. Just as with -yze spellings, however, in Canada the ize form remains the preferred or more common spelling, though both can still be found, yet
9672-426: The close historic, economic, and cultural relationship with the United States, -or endings are also sometimes used. Throughout the late 19th and early to mid-20th century, most Canadian newspapers chose to use the American usage of -or endings, originally to save time and money in the era of manual movable type . However, in the 1990s, the majority of Canadian newspapers officially updated their spelling policies to
9796-403: The dramatic arts and buildings where stage performances and screenings of films take place (i.e., " movie theaters "); for example, a national newspaper such as The New York Times would use theater in its entertainment section. However, the spelling theatre appears in the names of many New York City theatres on Broadway (cf. Broadway theatre) and elsewhere in the United States. In 2003,
9920-446: The egg cell (at the bottom of the gametophyte near the micropyle), forming a diploid (2n) zygote . This is the point when fertilisation actually occurs; pollination and fertilisation are two separate processes. The nucleus of the other sperm cell fuses with two haploid polar nuclei (contained in the central cell) in the centre of the gametophyte. The resulting cell is triploid (3n). This triploid cell divides through mitosis and forms
10044-555: The egg to develop without fertilisation or syngamy. The sperm may enter the egg. Hybridogenesis : One genome is eliminated to produce haploid eggs. Canina meiosis : (sometimes called "permanent odd polyploidy") one genome is transmitted in the Mendelian fashion, others are transmitted clonally. The major benefit of cross-fertilisation is generally thought to be the avoidance of inbreeding depression . Charles Darwin , in his 1876 book The Effects of Cross and Self Fertilisation in
10168-405: The eggs via chemotaxis , a type of ligand/receptor interaction. Resact is a 14 amino acid peptide purified from the jelly coat of A. punctulata that attracts the migration of sperm. After finding the egg, the sperm penetrates the jelly coat through a process called sperm activation. In another ligand/receptor interaction, an oligosaccharide component of the egg binds and activates a receptor on
10292-715: The ending became ‑our to match the later Old French spelling. The ‑our ending was used not only in new English borrowings, but was also applied to the earlier borrowings that had used ‑or . However, ‑or was still sometimes found. The first three folios of Shakespeare 's plays used both spellings before they were standardised to ‑our in the Fourth Folio of 1685. After the Renaissance , new borrowings from Latin were taken up with their original ‑or ending, and many words once ending in ‑our (for example, chancellour and governour ) reverted to ‑or . A few words of
10416-406: The etymology was not clear, and therefore some scholars advocated ‑or only and others ‑our only. Webster's 1828 dictionary had only -or and is given much of the credit for the adoption of this form in the United States. By contrast, Johnson's 1755 (pre-U.S. independence and establishment) dictionary used -our for all words still so spelled in Britain (like colour ), but also for words where
10540-469: The female reproductive tract such that the pH near the vaginal opening is lower (approximately 5) than the oviducts (approximately 8). The sperm-specific pH-sensitive calcium transport protein called CatSper increases the sperm cell permeability to calcium as it moves further into the reproductive tract. Intracellular calcium influx contributes to sperm capacitation and hyperactivation, causing a more violent and rapid non-linear motility pattern as sperm approach
10664-676: The first plants to flower and produce seeds in space . They had a life span of 40 days. Arabidopsis thaliana seeds were taken to the Moon on the Chang'e 4 lander in 2019, as part of a student experiment. As of May 2022 Arabidopsis thaliana has successfully been grown in samples of lunar soil. Arabidopsis is quite similar to the Boechera genus. The following species previously placed in Arabidopsis are not currently considered part of
10788-441: The formation of a diploid cell called a zygote . The zygote divides to form a blastocyst and, upon entering the uterus, implants in the endometrium, beginning pregnancy . Embryonic implantation not in the uterine wall results in an ectopic pregnancy that can kill the mother. In such animals as rabbits, coitus induces ovulation by stimulating the release of the pituitary hormone gonadotropin; this release greatly increases
10912-447: The formation of a fertilisation cone . Mammals internally fertilise through copulation . After a male ejaculates , many sperm move to the upper vagina (via contractions from the vagina) through the cervix and across the length of the uterus to meet the ovum. In cases where fertilisation occurs, the female usually ovulates during a period that extends from hours before copulation to a few days after; therefore, in most mammals, it
11036-567: The genus Arabidopsis has nine species and a further eight subspecies recognised. This delimitation is quite recent and is based on morphological and molecular phylogenies by O'Kane and Al-Shehbaz and others. Their findings confirm the species formerly included in Arabidopsis made it polyphyletic . The most recent reclassification moves two species previously placed in Cardaminopsis and Hylandra and three species of Arabis into Arabidopsis , but excludes 50 that have been moved into
11160-448: The genus. Cytogenetic analysis has shown the haploid chromosome number (n) is variable and varies across species in the genus: A. thaliana is n=5 and the DNA sequencing of this species was completed in 2001. A. lyrata has n=8 but some subspecies or populations are tetraploid. Various subspecies A. arenosa have n=8 but can be either 2n (diploid) or 4n (tetraploid). A. suecica
11284-685: The influence of the American labor movement and King O'Malley . On top of that, some place names in South Australia such as Victor Harbor , Franklin Harbor or Outer Harbor are usually spelled with the -or spellings. Aside from that, -our is now almost universal in Australia but the -or endings remain a minority variant. New Zealand English , while sharing some words and syntax with Australian English , follows British usage. In British English, some words from French, Latin or Greek end with
11408-539: The likelihood of pregnancy. Fertilisation in humans is the union of a human egg and sperm , usually occurring in the ampulla of the fallopian tube , producing a single celled zygote , the first stage of life in the development of a genetically unique organism, and initiating embryonic development . Scientists discovered the dynamics of human fertilisation in the nineteenth century. The term conception commonly refers to "the process of becoming pregnant involving fertilisation or implantation or both". Its use makes it
11532-415: The mother, or in some cases genetically differ from her but inherit only part of her DNA. Parthenogenesis occurs in many plants and animals and may be induced in others through a chemical or electrical stimulus to the egg cell. In 2004, Japanese researchers led by Tomohiro Kono succeeded after 457 attempts to merge the ova of two mice by blocking certain proteins that would normally prevent the possibility;
11656-414: The nature of the suffix used. The u is kept before English suffixes that are freely attachable to English words (for example in humourless , neighbourhood , and savoury ) and suffixes of Greek or Latin origin that have been adopted into English (for example in behaviourism , favourite , and honourable ). However, before Latin suffixes that are not freely attachable to English words,
11780-488: The new genera Beringia , Crucihimalaya , Ianhedgea , Olimarabidopsis , and Pseudoarabidopsis . All of the species in Arabidopsis are indigenous to Europe , while two of the species have broad ranges also extending into North America and Asia . In the last two decades, Arabidopsis thaliana has gained much interest from the scientific community as a model organism for research on numerous aspects of plant biology. The Arabidopsis Information Resource (TAIR)
11904-446: The oocyte. The capacitated spermatozoon and the oocyte meet and interact in the ampulla of the fallopian tube . Rheotaxis, thermotaxis and chemotaxis are known mechanisms that guide sperm towards the egg during the final stage of sperm migration. Spermatozoa respond (see Sperm thermotaxis ) to the temperature gradient of ~2 °C between the oviduct and the ampulla, and chemotactic gradients of progesterone have been confirmed as
12028-415: The ovary starts to swell and develop into the fruit . With multi-seeded fruits, multiple grains of pollen are necessary for syngamy with each ovule. The growth of the pollen tube is controlled by the vegetative (or tube) cytoplasm. Hydrolytic enzymes are secreted by the pollen tube that digest the female tissue as the tube grows down the stigma and style; the digested tissue is used as a nutrient source for
12152-557: The ovule is the most receptive to pollen tubes, and lowest during times of development and following fertilisation. High amounts of ROS activate Calcium ion channels in the pollen tube, causing these channels to take up Calcium ions in large amounts. This increased uptake of calcium causes the pollen tube to rupture, and release its sperm into the ovule. Pistil feeding assays in which plants were fed diphenyl iodonium chloride (DPI) suppressed ROS concentrations in Arabidopsis , which in turn prevented pollen tube rupture. After being fertilised,
12276-414: The pollen tube as it grows. During pollen tube growth towards the ovary, the generative nucleus divides to produce two separate sperm nuclei (haploid number of chromosomes) – a growing pollen tube therefore contains three separate nuclei, two sperm and one tube. The sperms are interconnected and dimorphic, the large one, in a number of plants, is also linked to the tube nucleus and the interconnected sperm and
12400-416: The pollen tube, and a second fertilisation event occurs involving the second sperm cell and the central cell of the ovule, which is a second female gamete. Unlike animal sperm which is motile, the sperm of most seed plants is immotile and relies on the pollen tube to carry it to the ovule where the sperm is released. The pollen tube penetrates the stigma and elongates through the extracellular matrix of
12524-579: The proteins. Transgenic plants lacking the ability to produce TTS proteins had slower pollen tube growth and reduced fertility. The rupture of the pollen tube to release sperm in Arabidopsis has been shown to depend on a signal from the female gametophyte. Specific proteins called FER protein kinases present in the ovule control the production of highly reactive derivatives of oxygen called reactive oxygen species (ROS). ROS levels have been shown via GFP to be at their highest during floral stages when
12648-546: The question of how the sperm and the appropriate egg find each other and the question of how only one sperm gets into the egg and delivers its contents. There are three steps to fertilisation that ensure species-specificity: Consideration as to whether an animal (more specifically a vertebrate) uses internal or external fertilisation is often dependent on the method of birth. Oviparous animals laying eggs with thick calcium shells, such as chickens , or thick leathery shells generally reproduce via internal fertilisation so that
12772-462: The resulting embryo normally developed into a mouse. Allogamy , which is also known as cross-fertilisation, refers to the fertilisation of an egg cell from one individual with the male gamete of another. Autogamy which is also known as self-fertilisation, occurs in such hermaphroditic organisms as plants and flatworms; therein, two gametes from one individual fuse. Some relatively unusual forms of reproduction are: Gynogenesis : A sperm stimulates
12896-456: The same parents. And this fact is amply sufficient to account for the development of the sexual elements, that is, for the genesis of the two sexes." In addition, it is thought by some, that a long-term advantage of out-crossing in nature is increased genetic variability that promotes adaptation or avoidance of extinction (see Genetic variability ). American and British English spelling differences#-ise, -ize (-isation, -ization) Despite
13020-408: The signal emanating from the cumulus oophorus cells surrounding rabbit and human oocytes. Capacitated and hyperactivated sperm respond to these gradients by changing their behaviour and moving towards the cumulus-oocyte complex. Other chemotactic signals such as formyl Met-Leu-Phe (fMLF) may also guide spermatozoa. The zona pellucida , a thick layer of extracellular matrix that surrounds the egg and
13144-475: The simplified form palestra is described by Merriam-Webster as "chiefly Brit[ish]." This is a reverse of the typical rule, where British spelling uses the ae / oe and American spelling simply uses e . Words that can be spelled either way in British English include cham a eleon , encyclop a edia , hom o eopathy , medi a eval (a minor variant in both AmE and BrE ), f o etid and f o etus . The spellings f o etus and f o etal are Britishisms based on
13268-410: The sounds became monophthongs , and later applied to words not of Greek origin, in both Latin (for example, cœli ) and French (for example, œuvre ). In English, which has adopted words from all three languages, it is now usual to replace Æ/æ with Ae/ae and Œ/œ with Oe/oe . In many words, the digraph has been reduced to a lone e in all varieties of English: for example, o economics , pr
13392-695: The spelling "is often seen as such". The -ize form is known as Oxford spelling and is used in publications of the Oxford University Press, most notably the Oxford English Dictionary , and of other academic publishers such as Nature , the Biochemical Journal and The Times Literary Supplement . It can be identified using the IETF language tag en-GB-oxendict (or, historically, by en-GB-oed ). In Ireland, India, Australia, and New Zealand -ise spellings strongly prevail:
13516-608: The spellings with just e are increasingly used. Manoeuvre is the only spelling in Australia, and the most common one in Canada, where maneuver and manoeuver are also sometimes found. The -ize spelling is often incorrectly seen in Britain as an Americanism. It has been in use since the 15th century, predating the -ise spelling by over a century. The verb-forming suffix -ize comes directly from Ancient Greek -ίζειν ( -ízein ) or Late Latin -izāre , while -ise comes via French -iser . The Oxford English Dictionary ( OED ) recommends -ize and lists
13640-435: The sperm and causes the acrosomal reaction . The acrosomal vesicles of the sperm fuse with the plasma membrane and are released. In this process, molecules bound to the acrosomal vesicle membrane, such as bindin, are exposed on the surface of the sperm. These contents digest the jelly coat and eventually the vitelline membrane. In addition to the release of acrosomal vesicles, there is explosive polymerisation of actin to form
13764-444: The sperm and egg takes place within the archegonium . In seed plants , the male gametophyte is formed within a pollen grain. After pollination , the pollen grain germinates , and a pollen tube grows and penetrates the ovule through a tiny pore called a micropyle . The sperm are transferred from the pollen through the pollen tube to the ovule where the egg is fertilised. In flowering plants , two sperm cells are released from
13888-501: The sperm fertilises the egg without having to pass through the thick, protective, tertiary layer of the egg. Ovoviviparous and viviparous animals also use internal fertilisation. Although some organisms reproduce via amplexus , they may still use internal fertilisation, as with some salamanders. Advantages of internal fertilisation include minimal waste of gametes, greater chance of individual egg fertilisation, longer period of egg protection, and selective fertilisation. Many females have
14012-406: The style before reaching the ovary. Then near the receptacle, it breaks through the ovule through the micropyle (an opening in the ovule wall) and the pollen tube "bursts" into the embryo sac, releasing sperm. The growth of the pollen tube has been believed to depend on chemical cues from the pistil, however these mechanisms were poorly understood until 1995. Work done on tobacco plants revealed
14136-427: The tube nucleus form the "male germ unit". Double fertilisation is the process in angiosperms (flowering plants) in which two sperm from each pollen tube fertilise two cells in a female gametophyte (sometimes called an embryo sac) that is inside an ovule. After the pollen tube enters the gametophyte, the pollen tube nucleus disintegrates and the two sperm cells are released; one of the two sperm cells fertilises
14260-486: The typical centriole , and atypical centriole that is attached to the flagellum , but not the mitochondria , to enter the oocyte. The protein CD9 likely mediates this fusion in mice (the binding homolog). The egg " activates " itself upon fusing with a single sperm cell and thereby changes its cell membrane to prevent fusion with other sperm. Zinc atoms are released during this activation. This process ultimately leads to
14384-605: The various English dialects spoken from country to country and within different regions of the same country, there are only slight regional variations in English orthography , the two most notable variations being British and American spelling. Many of the differences between American and British or Commonwealth English date back to a time before spelling standards were developed. For instance, some spellings seen as "American" today were once commonly used in Britain, and some spellings seen as "British" were once commonly used in
14508-412: The vowel is unreduced in pronunciation (e.g., devour , contour , flour , hour , paramour , tour , troubadour , and velour ), the spelling is uniform everywhere. Most words of this kind came from Latin, where the ending was spelled ‑or . They were first adopted into English from early Old French , and the ending was spelled ‑our , ‑or or ‑ur . After the Norman conquest of England ,
14632-533: The whole valium of the egg, allowing the egg pronucleus to use the cables to get to the male pronucleus. As the male and female pronuclei approach each other, the single centrosome split into two centrosomes located in the interphase between the pronuclei. Then the centrosome via the astral microtubules polarises the genome inside the pronuclei. Organisms that normally reproduce sexually can also reproduce via parthenogenesis , wherein an unfertilised female gamete produces viable offspring. These offspring may be clones of
14756-624: The word fiord (instead of fjord ) . There is an increasing use of macrons in words that originated in Māori and an unambiguous preference for -ise endings (see below). Most words ending in an unstressed ‑our in British English (e.g., behaviour , colour , favour , flavour , harbour , honour , humour , labour , neighbour , rumour , splendour ) end in ‑or in American English ( behavior , color , favor , flavor , harbor , honor , humor , labor , neighbor , rumor , splendor ). Wherever
14880-428: The word pallor . As a general noun, rigour / ˈ r ɪ ɡ ər / has a u in the UK; the medical term rigor (sometimes / ˈ r aɪ ɡ ər / ) does not, such as in rigor mortis , which is Latin. Derivations of rigour / rigor such as rigorous , however, are typically spelled without a u , even in the UK. Words with the ending -irior , -erior or similar are spelled thus everywhere. The word armour
15004-666: Was not until the mid-1990s that Canadian-based dictionaries became increasingly common.) Worldwide, -ize endings prevail in scientific writing and are commonly used by many international organizations, such as United Nations Organizations (such as the World Health Organization and the International Civil Aviation Organization ) and the International Organization for Standardization (but not by
15128-428: Was once somewhat common in American usage but has disappeared except in some brand names such as Under Armour . The agent suffix -or ( separator , elevator , translator , animator , etc.) is spelled thus both in American and British English. Commonwealth countries normally follow British usage. Canadian English most commonly uses the -our ending and -our- in derivatives and inflected forms. However, owing to
15252-929: Was published. Following this, -re became the most common usage in Britain. In the United States, following the publication of Webster's Dictionary in the early 19th century, American English became more standardized, exclusively using the -er spelling. In addition, spelling of some words have been changed from -re to -er in both varieties. These include September , October , November , December , amber, blister , cadaver , chamber , chapter , charter , cider , coffer , coriander , cover , cucumber , cylinder , diaper , disaster , enter , fever , filter , gender , leper , letter , lobster , master , member , meter (measuring instrument) , minister , monster , murder , number , offer , order , oyster , powder , proper , render , semester , sequester , sinister , sober , surrender , tender , and tiger . Words using
15376-437: Was very influential in popularizing certain spellings in the United States, but he did not originate them. Rather [...] he chose already existing options such as center, color and check for the simplicity, analogy or etymology". William Shakespeare 's first folios , for example, used spellings such as center and color as much as centre and colour . Webster did attempt to introduce some reformed spellings, as did
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