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108-634: The clementine is a spontaneous citrus hybrid that arose in the late 19th century in Misserghin , Algeria , in the garden of the orphanage of the French Missionary Brother Clément Rodier , for whom it would be formally named in 1902. Some sources have attributed an earlier origin for the hybrid, pointing to similar fruit native to the provinces of Guangxi and Guangdong in present-day China , but these are likely distinct mandarin hybrids, and genomic analysis of

216-475: A synthetic population . In horticulture , the term stable hybrid is used to describe an annual plant that, if grown and bred in a small monoculture free of external pollen (e.g., an air-filtered greenhouse) produces offspring that are "true to type" with respect to phenotype; i.e., a true-breeding organism. Hybridization can occur in the hybrid zones where the geographical ranges of species, subspecies, or distinct genetic lineages overlap. For example,

324-463: A bactrian camel and a dromedary . There are many examples of felid hybrids , including the liger . The oldest-known animal hybrid bred by humans is the kunga equid hybrid produced as a draft animal and status symbol 4,500 years ago in Umm el-Marra , present-day Syria . The first known instance of hybrid speciation in marine mammals was discovered in 2014. The clymene dolphin ( Stenella clymene )

432-416: 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

540-564: A bear shot by a hunter in the Northwest Territories confirmed the existence of naturally occurring and fertile grizzly–polar bear hybrids . Hybridization between reproductively isolated species often results in hybrid offspring with lower fitness than either parental. However, hybrids are not, as might be expected, always intermediate between their parents (as if there were blending inheritance), but are sometimes stronger or perform better than either parental lineage or variety,

648-515: A considerable seed yield advantage over open pollinated varieties. Hybrid seed dominates the commercial maize seed market in the United States, Canada and many other major maize-producing countries. In a hybrid, any trait that falls outside the range of parental variation (and is thus not simply intermediate between its parents) is considered heterotic. Positive heterosis produces more robust hybrids, they might be stronger or bigger; while

756-509: A cross between a female horse and a male donkey, and the hinny, a cross between a female donkey and a male horse. Pairs of complementary types like the mule and hinny are called reciprocal hybrids. Polar bears and brown bears are another case of a hybridizing species pairs, and introgression among non-sister species of bears appears to have shaped the Ursidae family tree. Among many other mammal crosses are hybrid camels , crosses between

864-509: A degree that none of the originally genetically distinct population remains. In agriculture and animal husbandry , the Green Revolution 's use of conventional hybridization increased yields by breeding high-yielding varieties . The replacement of locally indigenous breeds, compounded with unintentional cross-pollination and crossbreeding (genetic mixing), has reduced the gene pools of various wild and indigenous breeds resulting in

972-602: A donkey as the father. A variety of mechanisms limit the success of hybridization, including the large genetic difference between most species. Barriers include morphological differences, differing times of fertility, mating behaviors and cues, and physiological rejection of sperm cells or the developing embryo. Some act before fertilization; others after it. In plants, some barriers to hybridization include blooming period differences, different pollinator vectors, inhibition of pollen tube growth, somatoplastic sterility, cytoplasmic-genic male sterility and structural differences of

1080-407: A few days after; therefore, in most mammals, it 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

1188-548: A group of about 50 natural hybrids between Australian blacktip shark and the larger common blacktip shark was found by Australia's eastern coast in 2012. Russian sturgeon and American paddlefish were hybridized in captivity when sperm from the paddlefish and eggs from the sturgeon were combined, unexpectedly resulting in viable offspring. This hybrid is called a sturddlefish . The two genera Asymmetron and Branchiostoma are able to produce viable hybrid offspring, even if none have lived into adulthood so far, despite

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1296-500: A hybrid is interpreted differently in animal and plant breeding, where there is interest in the individual parentage. In genetics , attention is focused on the numbers of chromosomes . In taxonomy, a key question is how closely related the parent species are. Species are reproductively isolated by strong barriers to hybridization, which include genetic and morphological differences, differing times of fertility, mating behaviors and cues, and physiological rejection of sperm cells or

1404-439: A long time, both intentionally for purposes such as biological control , and unintentionally, as with accidental escapes of individuals. Introductions can drastically affect populations, including through hybridization. There is a kind of continuum with three semi-distinct categories dealing with anthropogenic hybridization: hybridization without introgression, hybridization with widespread introgression (backcrossing with one of

1512-498: A mate among the species that raised it, instead of the species of its biological parents. Cagebird breeders sometimes breed bird hybrids known as mules between species of finch , such as goldfinch × canary . Among amphibians, Japanese giant salamanders and Chinese giant salamanders have created hybrids that threaten the survival of Japanese giant salamanders because of competition for similar resources in Japan. Among fish,

1620-420: A more violent and rapid non-linear motility pattern as sperm approach 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

1728-432: A phenomenon called heterosis, hybrid vigour, or heterozygote advantage . This is most common with plant hybrids. A transgressive phenotype is a phenotype that displays more extreme characteristics than either of the parent lines. Plant breeders use several techniques to produce hybrids, including line breeding and the formation of complex hybrids. An economically important example is hybrid maize (corn), which provides

1836-517: 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

1944-465: 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 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

2052-405: A species, such as between different breeds . Single cross hybrids result from the cross between two true-breeding organisms which produces an F1 hybrid (first filial generation). The cross between two different homozygous lines produces an F1 hybrid that is heterozygous ; having two alleles , one contributed by each parent and typically one is dominant and the other recessive . Typically,

2160-567: 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

2268-478: A thick layer of extracellular matrix that surrounds the egg and 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

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2376-400: 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

2484-427: 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 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

2592-448: 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

2700-577: Is a hybrid of two Atlantic species, the spinner and striped dolphins . In 2019, scientists confirmed that a skull found 30 years earlier was a hybrid between the beluga whale and narwhal , dubbed the narluga . Hybridization between species is common in birds. Hybrid birds are purposefully bred by humans, but hybridization is also common in the wild. Waterfowl have a particularly high incidence of hybridization, with at least 60% of species known to produce hybrids with another species. Among ducks , mallards widely hybridize with many other species, and

2808-407: Is a hybridization test widely used in genetics to determine whether two separately isolated mutants that have the same (or similar) phenotype are defective in the same gene or in different genes (see Complementation (genetics) article). If a hybrid organism containing the genomes of two different mutant parental organisms displays a wild type phenotype, it is ordinarily considered that

2916-448: Is a pH gradient within the micro-environment of 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

3024-571: Is a particularly common mechanism for speciation in plants, and is now known to be fundamental to the evolutionary history of plants. Plants frequently form polyploids , individuals with more than two copies of each chromosome. Whole genome doubling has occurred repeatedly in plant evolution. When two plant species hybridize, the hybrid may double its chromosome count by incorporating the entire nuclear genome of both parents, resulting in offspring that are reproductively incompatible with either parent because of different chromosome counts. Human impact on

3132-610: 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

3240-659: Is also more occasionally done in the livestock and pet trades; some well-known wild × domestic hybrids are beefalo and wolfdogs . Human selective breeding of domesticated animals and plants has also resulted in the development of distinct breeds (usually called cultivars in reference to plants); crossbreeds between them (without any wild stock ) are sometimes also imprecisely referred to as "hybrids". Hybrid humans existed in prehistory. For example, Neanderthals and anatomically modern humans are thought to have interbred as recently as 40,000 years ago. Mythological hybrids appear in human culture in forms as diverse as

3348-451: Is always sterile worker ants (and because ants are haplodiploid , unfertilized eggs become males). Without mating with males of the other species, the queens are unable to produce workers, and will fail to establish a colony of their own. Plant species hybridize more readily than animal species, and the resulting hybrids are fertile more often. Many plant species are the result of hybridization, combined with polyploidy , which duplicates

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3456-552: Is derived from Latin hybrida , used for crosses such as of a tame sow and a wild boar. The term came into popular use in English in the 19th century, though examples of its use have been found from the early 17th century. Conspicuous hybrids are popularly named with portmanteau words , starting in the 1920s with the breeding of tiger–lion hybrids ( liger and tigon ). From the point of view of animal and plant breeders, there are several kinds of hybrid formed from crosses within

3564-402: 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

3672-405: 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

3780-419: 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 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

3888-489: Is in significant content (59% of the Daily Value ) in a 100 gram reference serving, with all other nutrients in low amounts. A 2017 study indicated that clementine phytochemicals may interact with drugs in a manner similar to those of grapefruit . A follow-up study in 2019, however, has called these results into question. Hybrid (biology) In biology , a hybrid is the offspring resulting from combining

3996-473: 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

4104-427: Is that hybrid individuals can form a "bridge" transmitting potentially helpful genes from one species to another when the hybrid backcrosses with one of its parent species, a process called introgression . Hybrids can also cause speciation , either because the hybrids are genetically incompatible with their parents and not each other, or because the hybrids occupy a different niche than either parent. Hybridization

4212-413: 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 the egg cell (at the bottom of the gametophyte near

4320-515: Is the proper time to give up on a population that is becoming a hybrid swarm, or to try and save the still existing pure individuals. Once a population becomes a complete mixture, the goal becomes to conserve those hybrids to avoid their loss. Conservationists treat each case on its merits, depending on detecting hybrids within the population. It is nearly impossible to formulate a uniform hybridization policy, because hybridization can occur beneficially when it occurs "naturally", and when hybrid swarms are

4428-601: 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

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4536-559: 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

4644-653: The Minotaur , blends of animals, humans and mythical beasts such as centaurs and sphinxes , and the Nephilim of the Biblical apocrypha described as the wicked sons of fallen angels and attractive women. Hybridization between species plays an important role in evolution, though there is much debate about its significance. Roughly 25% of plants and 10% of animals are known to form hybrids with at least one other species. One example of an adaptive benefit to hybridization

4752-436: 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 the pollen tube as it grows. During pollen tube growth towards

4860-472: The sea urchin Strongylocentrotus purpuratus (female) and the sand dollar Dendraster excentricus (male). When two distinct types of organisms breed with each other, the resulting hybrids typically have intermediate traits (e.g., one plant parent has red flowers, the other has white, and the hybrid, pink flowers). Commonly, hybrids also combine traits seen only separately in one parent or

4968-438: The "suture region". It is at these regions that the subspecies were formed. Other hybrid zones have formed between described species of plants and animals. From the point of view of genetics, several different kinds of hybrid can be distinguished. A genetic hybrid carries two different alleles of the same gene , where for instance one allele may code for a lighter coat colour than the other. A structural hybrid results from

5076-506: The F1 generation is also phenotypically homogeneous, producing offspring that are all similar to each other. Double cross hybrids result from the cross between two different F1 hybrids (i.e., there are four unrelated grandparents). Three-way cross hybrids result from the cross between an F1 hybrid and an inbred line. Triple cross hybrids result from the crossing of two different three-way cross hybrids. Top cross (or "topcross") hybrids result from

5184-506: 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

5292-469: 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

5400-557: The ZP3 and is important for binding with the sperm and activating 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,

5508-469: 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

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5616-406: 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

5724-400: 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

5832-467: 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 ,

5940-524: The butterfly Limenitis arthemis has two major subspecies in North America, L. a. arthemis (the white admiral) and L. a. astyanax (the red-spotted purple). The white admiral has a bright, white band on its wings, while the red-spotted purple has cooler blue-green shades. Hybridization occurs between a narrow area across New England, southern Ontario, and the Great Lakes,

6048-467: 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

6156-426: The chromosomes. A few animal species are the result of hybridization. The Lonicera fly is a natural hybrid. The American red wolf appears to be a hybrid of the gray wolf and the coyote , although its taxonomic status has been a subject of controversy. The European edible frog is a semi-permanent hybrid between pool frogs and marsh frogs ; its population requires the continued presence of at least one of

6264-547: The chromosomes. Chromosome duplication allows orderly meiosis and so viable seed can be produced. Plant hybrids are generally given names that include an "×" (not in italics), such as Platanus × hispanica for the London plane, a natural hybrid of P. orientalis (oriental plane) and P. occidentalis (American sycamore). The parent's names may be kept in their entirety, as seen in Prunus persica × Prunus americana , with

6372-562: The clementine has shown it to have arisen from a cross between a sweet orange ( Citrus × sinensis ) and the Mediterranean willowleaf mandarin ( Citrus × deliciosa ), consistent with Algerian origin. There are three types of clementines: seedless clementines, clementines (maximum of 10 seeds), and Monreal (more than 10 seeds). Clementines resemble other citrus varieties such as the satsuma and tangerines . Clementines differ from other citrus in having lower heat requirement, which means

6480-481: The common pheasant and hen golden pheasant ( Chrysolophus pictus ). Spurs are absent in hybrids of the former type, although present in both parents. Hybridization is greatly influenced by human impact on the environment, through effects such as habitat fragmentation and species introductions. Such impacts make it difficult to conserve the genetics of populations undergoing introgressive hybridization . Humans have introduced species worldwide to environments for

6588-400: The crossing of a top quality or pure-bred male and a lower quality female, intended to improve the quality of the offspring, on average. Population hybrids result from the crossing of plants or animals in one population with those of another population. These include interspecific hybrids or crosses between different breeds. In biology, the result of crossing of two populations is called

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6696-444: The developing embryo . Some act before fertilization and others after it. Similar barriers exist in plants, with differences in flowering times, pollen vectors, inhibition of pollen tube growth, somatoplastic sterility, cytoplasmic-genic male sterility and the structure of the chromosomes. A few animal species and many plant species, however, are the result of hybrid speciation , including important crop plants such as wheat , where

6804-407: The diverse Heliconius butterflies , but that is disputed. The two closely related harvester ant species Pogonomyrmex barbatus and Pogonomyrmex rugosus have evolved to depend on hybridization. When a queen fertilizes her eggs with sperm from males of her own species, the offspring is always new queens. And when she fertilizes the eggs with sperm from males of the other species, the offspring

6912-413: The doubling of chromosome sets, causing immediate genetic isolation. Hybridization may be important in speciation in some plant groups. However, homoploid hybrid speciation (not increasing the number of sets of chromosomes) may be rare: by 1997, only eight natural examples had been fully described. Experimental studies suggest that hybridization offers a rapid route to speciation, a prediction confirmed by

7020-555: 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

7128-405: 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

7236-611: The environment has resulted in an increase in the interbreeding between regional species, and the proliferation of introduced species worldwide has also resulted in an increase in hybridization. This has been referred to as genetic pollution out of concern that it may threaten many species with extinction. Similarly, genetic erosion from monoculture in crop plants may be damaging the gene pools of many species for future breeding. The conservation impacts of hybridization between species are highly debated. While hybridization could potentially threaten rare species or lineages by "swamping"

7344-429: The fact that early generation hybrids and ancient hybrid species have matching genomes, meaning that once hybridization has occurred, the new hybrid genome can remain stable. Many hybrid zones are known where the ranges of two species meet, and hybrids are continually produced in great numbers. These hybrid zones are useful as biological model systems for studying the mechanisms of speciation. Recently DNA analysis of

7452-512: The female parent's name given first, or if not known, the parent's names given alphabetically. Fertilization This is an accepted version of this page 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

7560-441: 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

7668-399: 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

7776-479: The formation of new individuals through fusion of male and female fluids, with form and function emerging gradually, in 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

7884-565: The fusion of gametes that have differing structure in at least one chromosome, as a result of structural abnormalities . A numerical hybrid results from the fusion of gametes having different haploid numbers of chromosomes . A permanent hybrid results when only the heterozygous genotype occurs, as in Oenothera lamarckiana , because all homozygous combinations are lethal. In the early history of genetics, Hugo de Vries supposed these were caused by mutation . Genetic complementation

7992-466: 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 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

8100-432: The fusion of nuclei of spermatozoa and of ova from sea urchin . The evolution of fertilisation 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

8208-406: The gametes produced by the male and female gametophytes come together and are fertilised. In bryophytes and pteridophytic land plants, fertilisation of 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

8316-507: The genetic relationships between ducks are further complicated by the widespread gene flow between wild and domestic mallards. One of the most common interspecific hybrids in geese occurs between Greylag and Canada geese ( Anser anser x Branta canadensis ). One potential mechanism for the occurrence of hybrids in these geese is interspecific nest parasitism , where an egg is laid in the nest of another species to be raised by non-biological parents. The chick imprints upon and eventually seeks

8424-630: The genetically "pure" individuals with hybrids, hybridization could also save a rare lineage from extinction by introducing genetic diversity. It has been proposed that hybridization could be a useful tool to conserve biodiversity by allowing organisms to adapt, and that efforts to preserve the separateness of a "pure" lineage could harm conservation by lowering the organisms' genetic diversity and adaptive potential, particularly in species with low populations. While endangered species are often protected by law, hybrids are often excluded from protection, resulting in challenges to conservation. The term hybrid

8532-401: The hybrids are more fit and have breeding advantages over the indigenous ecotype or species. These hybridization events can result from the introduction of non-native genotypes by humans or through habitat modification, bringing previously isolated species into contact. Genetic mixing can be especially detrimental for rare species in isolated habitats, ultimately affecting the population to such

8640-541: 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

8748-400: The loss of genetic diversity . Since the indigenous breeds are often well-adapted to local extremes in climate and have immunity to local pathogens, this can be a significant genetic erosion of the gene pool for future breeding. Therefore, commercial plant geneticists strive to breed "widely adapted" cultivars to counteract this tendency. Familiar examples of equid hybrids are the mule,

8856-411: 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 the endosperm ,

8964-415: 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;

9072-452: The number of chromosomes has been doubled. A form of often intentional human-mediated hybridization is the crossing of wild and domesticated species. This is common in both traditional horticulture and modern agriculture ; many commercially useful fruits, flowers, garden herbs, and trees have been produced by hybridization. One such flower, Oenothera lamarckiana , was central to early genetics research into mutationism and polyploidy. It

9180-420: The only remaining evidence of prior species, they need to be conserved as well. Regionally developed ecotypes can be threatened with extinction when new alleles or genes are introduced that alter that ecotype. This is sometimes called genetic mixing. Hybridization and introgression, which can happen in natural and hybrid populations, of new genetic material can lead to the replacement of local genotypes if

9288-438: The other (e.g., a bird hybrid might combine the yellow head of one parent with the orange belly of the other). Interspecific hybrids are bred by mating individuals from two species, normally from within the same genus. The offspring display traits and characteristics of both parents, but are often sterile , preventing gene flow between the species. Sterility is often attributed to the different number of chromosomes between

9396-421: 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 the tube nucleus form the "male germ unit". Double fertilisation

9504-444: The oviduct and the ampulla, and chemotactic gradients of progesterone have been confirmed as 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 ,

9612-573: The parent species), and hybrid swarms (highly variable populations with much interbreeding as well as backcrossing with the parent species). Depending on where a population falls along this continuum, the management plans for that population will change. Hybridization is currently an area of great discussion within wildlife management and habitat management. Global climate change is creating other changes such as difference in population distributions which are indirect causes for an increase in anthropogenic hybridization. Conservationists disagree on when

9720-537: The parent species. Cave paintings indicate that the European bison is a natural hybrid of the aurochs and the steppe bison . Plant hybridization is more commonplace compared to animal hybridization. Many crop species are hybrids, including notably the polyploid wheats : some have four sets of chromosomes (tetraploid) or six (hexaploid), while other wheat species have (like most eukaryotic organisms) two sets ( diploid ), so hybridization events likely involved

9828-506: The parents' common ancestor living tens of millions of years ago. Among insects, so-called killer bees were accidentally created during an attempt to breed a strain of bees that would both produce more honey and be better adapted to tropical conditions. It was done by crossing a European honey bee and an African bee . The Colias eurytheme and C. philodice butterflies have retained enough genetic compatibility to produce viable hybrid offspring. Hybrid speciation may have produced

9936-637: The point of view of taxonomy , hybrids differ according to their parentage. Hybrids between different subspecies (such as between the dog and Eurasian wolf ) are called intra-specific hybrids. Interspecific hybrids are the offspring from interspecies mating ; these sometimes result in hybrid speciation. Intergeneric hybrids result from matings between different genera, such as between sheep and goats . Interfamilial hybrids, such as between chickens and guineafowl or pheasants , are reliably described but extremely rare. Interordinal hybrids (between different orders) are few, but have been engineered between

10044-449: 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 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,

10152-491: 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, the ovary starts to swell and develop into

10260-580: The qualities of two organisms of different varieties , subspecies , species or genera through sexual reproduction . Generally, it means that each cell has genetic material from two different organisms, whereas an individual where some cells are derived from a different organism is called a chimera . Hybrids are not always intermediates between their parents such as in blending inheritance (a now discredited theory in modern genetics by particulate inheritance ), but can show hybrid vigor , sometimes growing larger or taller than either parent. The concept of

10368-546: 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

10476-465: 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

10584-435: 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

10692-501: 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

10800-470: The term negative heterosis refers to weaker or smaller hybrids. Heterosis is common in both animal and plant hybrids. For example, hybrids between a lion and a tigress (" ligers ") are much larger than either of the two progenitors, while " tigons " (lioness × tiger) are smaller. Similarly, the hybrids between the common pheasant ( Phasianus colchicus ) and domestic fowl ( Gallus gallus ) are larger than either of their parents, as are those produced between

10908-513: The tolerance to fruit maturity and sensitivity to unfavorable conditions during the flowering and fruit-setting period is higher. However, in regions of high total heat, the Clementine bears fruit early; only slightly later than satsuma mandarins. These regions such as North Africa, Mediterranean basin, and California, also favor maximizing the Clementine size and quality. It was introduced into California commercial agriculture in 1914, though it

11016-495: 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 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

11124-400: The two parental mutant organisms are defective in different genes. If the hybrid organism displays a distinctly mutant phenotype, the two mutant parental organisms are considered to be defective in the same gene. However, in some cases the hybrid organism may display a phenotype that is only weakly (or partially) wild-type, and this may reflect intragenic (interallelic) complementation. From

11232-462: The two species. For example, donkeys have 62 chromosomes , horses have 64 chromosomes, and mules or hinnies have 63 chromosomes. Mules, hinnies, and other normally sterile interspecific hybrids cannot produce viable gametes, because differences in chromosome structure prevent appropriate pairing and segregation during meiosis , meiosis is disrupted, and viable sperm and eggs are not formed. However, fertility in female mules has been reported with

11340-488: 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

11448-534: 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

11556-537: The wild of less than 0.3%; 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

11664-716: Was grown at the Citrus Research Center (now part of the University of California, Riverside ) as early as 1909. Clementines lose their desirable seedless characteristic when they are cross-pollinated with other fruit. In 2006, to prevent this, growers such as Paramount Citrus in California threatened to sue local beekeepers to keep bees away from their crops. A clementine contains 87% water, 12% carbohydrates , and negligible amounts of fat and protein (table). Among micronutrients , only vitamin C

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