Chimera (genetics) - Knowledge (XXG)

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embryo and diploid embryo, ES cell and tetraploid embryo, diploid embryo and tetraploid embryo, ES cells and ES cells. The combination of embryonic stem cell and diploid embryo is a common technique used for the making of chimeric mice, since gene targeting can be done in the embryonic stem cell. These kinds of chimeras can be made through either aggregation of stem cells and the diploid embryo or injection of the stem cells into the diploid embryo. If embryonic stem cells are to be used for gene targeting to make a chimera, the following procedure is common: a construct for homologous recombination for the gene targeted will be introduced into cultured mouse embryonic stem cells from the donor mouse, by way of electroporation; cells positive for the recombination event will have antibiotic resistance, provided by the insertion cassette used in the gene targeting; and be able to be positively selected for. ES cells with the correct targeted gene are then injected into a diploid host mouse blastocyst. Then, these injected blastocysts are implanted into a pseudo pregnant female surrogate mouse, which will bring the embryos to term and give birth to a mouse whose germline is derived from the donor mouse's ES cells. This same procedure can be achieved through aggregation of ES cells and diploid embryos, diploid embryos are cultured in aggregation plates in wells where single embryos can fit, to these wells ES cells are added the aggregates are cultured until a single embryo is formed and has progressed to the blastocyst stage, and can then be transferred to the surrogate mouse.

957:. Each of these parts of the blastocyst gives rise to different parts of the embryo; the inner cell mass gives rise to the embryo proper, while the trophectoderm and primitive endoderm give rise to extra embryonic structures that support growth of the embryo. Two- to eight-cell-stage embryos are competent for making chimeras, since at these stages of development, the cells in the embryos are not yet committed to give rise to any particular cell lineage, and could give rise to the inner cell mass or the trophectoderm. In the case where two diploid eight-cell-stage embryos are used to make a chimera, chimerism can be later found in the 785: 273: 334:

to be maternally-derived. However, his form of microchimerism was found to be due to a vanished twin, and it is unknown whether microchimerism from a vanished twin might predispose individuals to autoimmune diseases as well. Mothers often also have a few cells genetically identical to those of their children, and some people also have some cells genetically identical to those of their siblings (maternal siblings only, since these cells are passed to them because their mother retained them).

589: 289: 900: 45: 620: 928:(ES cells). Chimeras can be derived from mouse embryos that have not yet implanted in the uterus as well as from implanted embryos. ES cells from the inner cell mass of an implanted blastocyst can contribute to all cell lineages of a mouse including the germ line. ES cells are a useful tool in chimeras because genes can be mutated in them through the use of 3770: 330:

retain higher numbers of cells genetically identical to their mother's have been observed to have higher rates of some autoimmune diseases, presumably because the immune system is responsible for destroying these cells and a common immune defect prevents it from doing so and also causes autoimmune problems.

711:, plastids may sort-out from a mixed egg or mixed zygote respectively. This type of chimera is recognized at the time of origin by the sorting-out pattern in the leaves. After sorting-out is complete, periclinal chimeras are distinguished from similar looking nuclear gene-differential chimeras by their 333:

The higher rates of autoimmune diseases due to the presence of maternally-derived cells is why in a 2010 study of a 40-year-old man with scleroderma-like disease (an autoimmune rheumatic disease), the female cells detected in his blood stream via FISH (fluorescence in situ hybridization) were thought

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receptors, the male searches until it locates a female anglerfish. The male, less than an inch in length, bites into her skin and releases an enzyme that digests the skin of both his mouth and her body, fusing the pair down to the blood-vessel level. While this attachment has become necessary for the

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contributions to chimeras have been developed. ES cells can be used in combination with eight-cell-and two-cell-stage embryos to make chimeras and exclusively give rise to the embryo proper. Embryos that are to be used in chimeras can be further genetically altered to specifically contribute to only

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Chimerism has been found in some species of marine sponges. Four distinct genotypes have been found in a single individual, and there is potential for even greater genetic heterogeneity. Each genotype functions independently in terms of reproduction, but the different intra-organism genotypes behave

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The US and Western Europe have strict codes of ethics and regulations in place that expressly forbid certain subsets of experimentation using human cells, though there is a vast difference in the regulatory framework. Through the creation of human chimeras comes the question: where does society now

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The ability to make mouse chimeras comes from an understanding of early mouse development. Between the stages of fertilization of the egg and the implantation of a blastocyst into the uterus, different parts of the mouse embryo retain the ability to give rise to a variety of cell lineages. Once the

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Chimeric mice are important animals in biological research, as they allow for the investigation of a variety of biological questions in an animal that has two distinct genetic pools within it. These include insights into problems such as the tissue specific requirements of a gene, cell lineage, and

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are obligate chimeras, the first known such case. In this species, the queens have arisen from fertilized eggs with a genotype of RR (Reproductive x Reproductive), the sterile female workers show a RW arrangement (Reproductive x Worker), and the males instead of being haploid as is usually the case

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In the current embryological (which is also the classical) sense, a "chimaera" is an organism whose cells derive from two or more distinct zygote lineages, and this is the sense which the term "genetical chimaera" is here intended to convey. "Genetical mosaic" is less suitable, because a mosaic is

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Normally, genetic chimerism is not visible on casual inspection; however, it has been detected in the course of proving parentage. More practically, in agronomy Chimera indicates a plant or portion of a plant whose tissues are made up of two or more types of cells with different genetic makeup; it

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There are a variety of combinations that can give rise to a successful chimera mouse and – according to the goal of the experiment – an appropriate cell and embryo combination can be picked; they are generally but not limited to diploid embryo and ES cells, diploid

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and a straight thumb, eyes of slightly different colors, differential hair growth on opposite sides of the body, etc.) or completely undetectable. Chimeras may also show, under a certain spectrum of UV light, distinctive marks on the back resembling that of arrow points pointing downward from the

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Microchimerism is the presence of a small number of cells that are genetically distinct from those of the host individual. Most people are born with a few cells genetically identical to their mothers' and the proportion of these cells goes down in healthy individuals as they get older. People who

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if they pose a threat to humans. If a chimpanzee is genetically altered to be more similar to a human, it may blur the ethical line between animal and human. Legal debate would be the next step in the process to determine whether certain chimeras should be granted legal rights. Along with issues

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It is possible to dissect the embryo at other stages so as to accordingly give rise to one lineage of cells from an embryo selectively and not the other. For example, subsets of blastomeres can be used to give rise to chimera with specified cell lineage from one embryo. The Inner Cell Mass of a

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in the chick embryo stage, the neural tube and the neural crest cells of the chick were ablated, and replaced with the same parts from a quail. Once hatched, the quail feathers were visibly apparent around the wing area, whereas the rest of the chick's body was made of its own chicken cells.

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one part of chimera. An example is the chimera built off of ES cells and tetraploid embryos, which are artificially made by electrofusion of two two-cell diploid embryos. The tetraploid embryo will exclusively give rise to the trophectoderm and primitive endoderm in the chimera.

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individual. Sometimes in this process, more than one male will attach to a single female as a symbiote. In this case, they will all be consumed into the body of the larger female angler. Once fused to a female, the males will reach sexual maturity, developing large

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diploid blastocyst, for example, can be used to make a chimera with another blastocyst of eight-cell diploid embryo; the cells taken from the inner cell mass will give rise to the primitive endoderm and to the epiblast in the chimera mouse. From this knowledge,

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ability. This mutated tissue is unable to survive on its own, but it is kept alive by its partnership with normal photosynthetic tissue. Sometimes chimeras are also found with layers differing in respect of both their nuclear and their plastid genes.

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in the 1960s through the aggregation of eight-cell-stage embryos. Injection on the other hand was pioneered by Richard Gardner and Ralph Brinster who injected cells into blastocysts to create chimeric mice with germ lines fully derived from injected

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Mosaicism originates by intrinsic genetic variations caused, among other processes, by somatic mutations, while chimerism originates from allogenic fusion or grafting. As such, chimerism is much rarer and involves a much larger genetic change than

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Rakosy-Tican, E.; Aurori, C.M.; Dijkstra, C.; Thieme, R.; Aurori, A. & Davey, M.R. (2007). "The usefulness of the gfp reporter gene for monitoring Agrobacterium-mediated transformation of potato dihaploid and tetraploid genotypes".

557:, where they state, "... where a twin pregnancy evolves into one child, is currently believed to be one of the rarer forms. However, we know that 20 to 30% of singleton pregnancies were originally a twin or a multiple pregnancy". 837:

four-cell morulas; although the cells never fused, they worked together to form organs. It was discovered that one of these primates, Roku, was a sexual chimera; as four percent of Roku's blood cells contained two x chromosomes.

440:. Innate chimeras are formed from at least four parent cells (two fertilised eggs or early embryos fused together). Each population of cells keeps its own character and the resulting organism is a mixture of tissues. Cases of 489:. Specific types of transplants that could induce this condition include bone marrow transplants and organ transplants, as the recipient's body essentially works to permanently incorporate the new blood stem cells into it. 763:

Untransformed cells should be easy to detect and remove to avoid chimeras. This is because it is important to maintain the stable ability of the transgenic plants across different generations. Reporter genes such as

413:, tetragametic chimeras can be very conspicuous, as the resulting bird will have an obvious split between two colour types – often divided bilaterally down the centre. These individuals are known as 772:(GFP) are used in combination with plant selective markers (herbicide, antibody etc.) However, GUS expression depends on the plant development stage and GFP may be influenced by the green tissue autofluorescence. 1665:

Hadjiathanasiou, C. G.; Brauner, R.; Lortat-Jacob, S.; Nivot, S.; Jaubert, F.; Fellous, M.; Nihoul-Fékété, C.; Rappaport, R. (November 1994). "True hermaphroditism: genetic variants and clinical management".

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Cao, Jing; Li, Wenjuan; Li, Jie; Mazid, Md. Abdul; Li, Chunyang; Jiang, Yu; Jia, Wenqi; Wu, Liang; Liao, Zhaodi; Sun, Shiyu; Song, Weixiang; Fu, Jiqiang; Wang, Yan; Lu, Yong; Xu, Yuting (November 2023).

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These chimeras arise by spontaneous or induced mutation of a nuclear gene to a dominant or recessive allele. As a rule, one character is affected at a time in the leaf, flower, fruit, or other parts.

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to form a single growing organism that preserves both types of tissue in a single shoot. Just as the constituent species are likely to differ in a wide range of features, so the behavior of their

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composed of cells with a second set of chromosomes. This has occurred in humans, and at one time was thought to be extremely rare although more recent evidence suggests that this is not the case.

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can derive from a bud mutation or, more rarely, at the grafting point, from the concrescence of cells of the two bionts; in this case it is improperly known with the name of "graft hybrid".

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Schoenle, E.; Schmid, W.; Schinzel, A.; Mahler, M.; Ritter, M.; Schenker, T.; Metaxas, M.; Froesch, P.; Froesch, E. R. (1983-07-01). "46,XX/46,XY chimerism in a phenotypically normal man".

891:, an average of 67% and a highest of 92% of the cells across the 26 tested tissues were descendants of the donor stem cells against 0.1–4.5% from previous experiments on chimeric monkeys. 707:

These chimeras arise by spontaneous or induced mutation of a plastid gene, followed by the sorting-out of two kinds of plastid during vegetative growth. Alternatively, after selfing or

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Shah, V. C.; Krishna Murthy, D. S.; Roy, S.; Contractor, P. M.; Shah, A. V. (November 1982). "True hermaphrodite: 46, XX/46, XY, clinical cytogenetic and histopathological studies".

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draw the line of humanity? This question poses serious legal and moral issues, along with creating controversy. Chimpanzees, for example, are not offered any legal standing, and are

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The general methods for creating chimeric mice can be summarized either by injection or aggregation of embryonic cells from different origins. The first chimeric mouse was made by

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for ants also display a RW genotype, but for them the egg R and the sperm W do not fuse so they develop as a chimera with some cells carrying an R and others carrying a W genome.

691:. More commonly cytochimeras have simple multiple of the normal chromosome complement in the changed layer. There are various effects on cell size and growth characteristics. 887:

In 2023 a study reported the first chimeric monkey using embryonic stem cell lines, it was the only live birth from 12 pregnancies resulting from 40 implanted embryos of the

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Tetragametic chimerism is a form of congenital chimerism. This condition occurs through fertilizing two separate ova by two sperm, followed by aggregation of the two at the

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Zhu, X.; Zhao, M.; Ma, S.; Ge, Y.; Zhang, M. & Chen, L. (2007). "Induction and origin of adventitious shoots from chimeras of Brassica juncea and Brassica oleracea".

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ova to create the first human chimeric embryos. The embryos were allowed to develop for several days in a laboratory setting, and then destroyed to harvest the resulting

803: 671:, and "Family" trees, where multiple varieties of apple or pear are grafted onto the same tree. Many fruit trees are cultivated by grafting the body of a sapling onto a 354:

can carry the reproductive cells of their (fraternal) twin siblings due to placental fusion during development. (Marmosets almost always give birth to fraternal twins.)

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L. M. Repas-Humpe; A. Humpe; R. Lynen; B. Glock; E. M. Dauber; G. Simson; W. R. Mayr; M. Köhler; S. Eber (1999). "A Dispermic Chimerism in a 2-year-old Caucasian Boy".

236:. This process allows for sperm to be in constant supply when the female produces an egg, so that the chimeric fish is able to have a greater number of offspring. 466:

of the genetic material in a cell, giving rise to a subset of cells that are different from the rest. Natural chimerism is the fusion of more than one fertilized

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or zygote stages. This results in the development of an organism with intermingled cell lines. Put another way, the chimera is formed from the merging of two

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A chimera is an organism whose cells are derived from two or more zygotes as opposed to a mosaic whose different cell lines are derived from a single zygote

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regarding the rights of chimeras, individuals have expressed concern about whether or not creating human chimeras diminishes the "dignity" of being human.

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To research the developmental biology of the bird embryo, researchers produced artificial quail-chick chimeras in 1987. By using transplantation and

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Some level of chimerism occurs naturally in the wild in many animal species, however in some cases may be a required (obligate) part of their life cycle.

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Gengozian, N.; Batson, JS; Eide, P. (1964). "Hematologic and Cytogenetic Evidence for Hematopoietic Chimerism in the Marmoset, Tamarinus Nigricollis".

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Thompson, J.D.; Herre, E.A.; Hamrick, J.L. & Stone, J.L. (1991). "Genetic Mosaics in strangler Fig Trees: Implication for Tropical Conservation".

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In contrast, an individual where each cell contains genetic material from two organisms of different breeds, varieties, species or genera is called a

546:. Chimeras can often breed, but the fertility and type of offspring depend on which cell line gave rise to the ovaries or testes; varying degrees of 3953: 718:

All plastid gene- and some nuclear gene-differential chimeras affect the color of the plasmids within the leaves, and these are grouped together as

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and is in fact a natural and essential part of their life cycle. Once the male achieves adulthood, it begins its search for a female. Using strong

822:, which infect plants. The study surprised scientists, because DNA and RNA viruses vary and the way the chimera came together was not understood. 481:

that was inherited genetically at the time of the formation of the human embryo and the other that was introduced through a procedure, including

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In contrast, a human where each cell contains genetic material from two organisms of different breeds, varieties, species or genera is called a

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The first known primate chimeras are the rhesus monkey twins, Roku and Hex, each having six genomes. They were created by mixing cells from

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Artificial chimerism refers to examples of chimerism that are accidentally produced by humans, either for research or commercial purposes.

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Pappaioannou, V.; Johnson, R. (1993). Joyner, A. (ed.). "Production of chimeras and genetically defined offspring from targeted ES cells".

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Binkhorst, Mathijs; de Leeuw, Nicole; Otten, Barto J. (January 2009). "A healthy, female chimera with 46,XX/46,XY karyotype".

1132:"Temporal Change in Blood Group after Bone Marrow Transplant: A Case of Successful ABO-Incompatible Deceased Donor Transplant" 4124: 3715:

Yu N, Kruskall MS, Yunis JJ, Knoll JH, Uhl L, Alosco S, Ohashi M, Clavijo O, Husain Z, Yunis EJ, Yunis JJ, Yunis EJ (2002).

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Phytohormones in Plant Biotechnology and Agriculture: Proceedings of the NATO-Russia Workshop held in Moscow, 12–16 May 2002

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Park SH, Rose SC, Zapata C, Srivatanakul M (1998). "Cross-protection and selectable marker genes in plant transformation".

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Tanaka, M; Hadjantonakis, AK; Nagy, A (2001). "Aggregation Chimeras: Combining ES Cells, Diploid and Tetraploid Embryos".

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Most human chimeras will go through life without realizing they are chimeras. The difference in phenotypes may be subtle (

2782:"Using quantitative real-time PCR to detect chimeras in transgenic tobacco and apricot and to monitor their dissociation" 2345: 1513:

Farag, T I; Al-Awadi, S A; Tippett, P; el-Sayed, M; Sundareshan, T S; Al-Othman, S A; el-Badramany, M H (December 1987).

752:(3) A self-protection mechanism (cross protection). Transformed cells serve as guards to protect the untransformed ones. 1021: 687:

constitution. Occasionally, chimeras arise from loss or gain of individual chromosomes or chromosome fragments owing to

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Bellefon, L.; Heiman, P.; Kanaan, S.; Azzouz, D.; Rak, J.; Martin, M.; Roudier, J.; Roufosse, F.; Lambert, C. (2010).

936:. Since this discovery occurred in 1988, ES cells have become a key tool in the generation of specific chimeric mice. 708: 31: 2040: 722:

chimeras, or preferably as variegated leaf chimeras. For most variegation, the mutation involved is the loss of the

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or different species (which may belong to different genera). The tissues may be partially fused together following

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Norris, R.; Smith, R.H. & Vaughn, K.C. (1983). "Plant chimeras used to establish de novo origin of shoots".

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Gencík, A.; Genciková, A.; Hrubisko, M.; Mergancová, O. (1980). "Chimerism 46,XX/46,XY in a phenotypic female".

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In 2012, the first example of a naturally-occurring RNA-DNA hybrid virus was unexpectedly discovered during a

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Boklage, C.E. How New Humans Are Made. Hackensack, NJ; London: World Scientific Publishing Co. Pte. Ltd; 2010

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Another form of chimera is the mosaic, which is a composite individual derived from a single fertilized egg.

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Other viral chimeras have also been found, and the group is known as the CHIV viruses ("chimeric viruses").

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Rossant, J. (1976). "Postimplantation development of blastomeres isolated from 4- and 8-cell mouse eggs".

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Rinkevich, B. (June 2001). "Human natural chimerism: an acquired character or a vestige of evolution?".

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in 1901, the genetic science took until the 1930s to approach a vocabulary for the phenomenon. The term

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Chimera's Children: Ethical, Philosophical and Religious Perspectives on Human-Nonhuman Experimentation

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There are multiple reasons to explain the occurrence of plant chimera during the plant recovery stage:

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Boklage argues that many human 'mosaic' cell lines will be "found to be chimeric if properly tested".

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Tam, P.L.; Rossant, J. (2003). "Mouse embryonic chimeras: tools for studying mammalian development".

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Belgovskii, ML (1944). "The Causes of Mosaicism Associated With Heterochromatic Chromosome Regions".

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cells) of an organism are not genetically identical to its own. It has been recently discovered that

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This condition is either innate or it is synthetic, acquired for example through the infusion of

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differences may result if one set of cells is genetically female and another genetically male.

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Nagy, A.; Rossant, J. (1999). Joyner, A. (ed.). "Production of Es-cell aggregation chimeras".

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Ralston, A; Rossant, J (2005). "Genetic regulation of stem cell origins in the mouse embryo".

3270: 3176: 3101: 3093: 3046: 2925: 2887:"Could a newly discovered viral genome change what we thought we knew about virus evolution?". 2813: 2754: 2675: 2632: 2583:"Growing Fruit: Grafting Fruit Trees in the Home Orchard [fact sheet] | UNH Extension" 2555: 2518: 2466: 2465:(873). Wageningen, Netherlands: Centre for Agricultural Publishing and Documentation (PUDOC). 2430: 2405: 2282: 2241: 2233: 2194: 2186: 2140: 2098: 2090: 1996: 1961: 1810: 1792: 1740: 1691: 1683: 1647: 1626:"A True Hermaphrodite Chimera Resulting from Embryo Amalgamation after in Vitro Fertilization" 1595: 1587: 1552: 1534: 1487: 1479: 1436: 1428: 1385: 1377: 1334: 1291: 1283: 1236: 1228: 1163: 1091: 1003: 884:

School of Medicine created a sheep whose blood contained 15% human cells and 85% sheep cells.

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Boiling Springs Lake, California, is where the first natural chimeric virus was found in 2012.

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Robertson, EJ (1986). "Pluripotential stem cell lines as a route into the mouse germ line".

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Mintz, B.; Silvers, W. K. (1967). "'Intrinsic' Immunological Tolerance in Allophenic Mice".

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to be a form of chimerism, while others consider them to be distinct. Mosaicism involves a

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Single organism composed of two or more different populations of genetically distinct cells

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involves distinct types of tissue that originated from the same zygote but differ due to

3491: 3246: 3164: 3081: 2855:"Hot spring yields hybrid genome: Researchers discover natural chimaeric DNA-RNA virus". 2836: 2620: 2506: 1778: 1271: 1254:

Darras, H.; Berney, C.; Hasin, S.; Drescher, J.; Feldhaar, H.; Keller, L. (2023-04-07).

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male's survival, it will eventually consume him, as both anglerfish fuse into a single

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chimeras is like to be highly variable. The first such known chimera was probably the

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shoulders down to the lower back; this is one expression of pigment unevenness called

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embryo has reached the blastocyst stage, it is composed of several parts, mainly the

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in the mutated tissue, so that part of the plant tissue has no green pigment and no

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On January 22, 2019 the National Society of Genetic Counselors released an article:

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Chimera Mouse production by blastocyst injection, Wellcome trust Sanger Institute,

3127: 1499: 1448: 577: 478: 3172: 2628: 2545: 2459:"Mutation Breeding Techniques and Behaviour of Irradiated Shoot Apices of Potato" 2458: 2424: 4098: 3857: 3840: 3595: 3025:"Live birth of chimeric monkey with high contribution from embryonic stem cells" 2514: 2387: 2167:"Mosaicism and chimerism as components of intraorganismal genetic heterogeneity" 1642: 1625: 1042: 1016: 908: 819: 799: 723: 719: 651: 542:. The likelihood of offspring being a chimera is increased if it is created via 183: 3839:

Strain, Lisa; Dean, John C.S.; Hamilton, Mark P.R.; Bonthron, David T. (1998).

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transplantation, giving one individual tissues that developed from a different

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Anderson, D.; Billingham, R. E.; Lampkin, G. H.; Medawar, P. B. (1951-12-01).

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A major milestone in chimera experimentation occurred in 1984 when a chimeric

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that is composed of two or more different populations of genetically distinct

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Strain, Lisa; John C.S. Dean; Mark P. R. Hamilton; David T. Bonthron (1998).

1591: 1538: 1483: 1432: 1381: 1287: 1255: 1232: 1197:"'Living Together Apart': The Hidden Genetic Diversity of Sponge Populations" 3904: 3751: 3500: 3420:

Kubiak, J; Tarkowski, A. (1985). "Electrofusion of mouse blastomeres. Exp".

3255: 2798: 2353: 2277: 2120: 1787: 1279: 1213: 1196: 996: 904: 877: 672: 643: 639: 395: 390:. Recent research shows most marmosets are chimeras, sharing DNA with their 308: 219: 3823: 3742: 3682:"S.659 – Human Chimera Prohibition Act of 2005 (Introduced in Senate - IS)" 3613: 3560: 3352: 3309: 3105: 3050: 2817: 2758: 2679: 2636: 2522: 2286: 2245: 2198: 2102: 2000: 1965: 1814: 1744: 1726: 1440: 1338: 1295: 1240: 1167: 1148: 619: 245:

as a single large individual in terms of ecological responses like growth.

3866: 3815: 3519: 3441: 3387: 3274: 3180: 2929: 2346:"Chimerism Explained: How One Person Can Unknowingly Have Two Sets of DNA" 1695: 1651: 1599: 1556: 1530: 1491: 1389: 30:

This article is about Chimera genetics. For DNA sequencing artifacts, see

4003: 3733: 3716: 2904:"Origins and evolution of viruses of eukaryotes: The ultimate modularity" 1711:"Cells from a vanished twin as a source of microchimerism 40 years later" 962: 958: 861: 663: 635: 631: 547: 463: 421: 387: 351: 229: 149: 137: 17: 3717:"Disputed maternity leading to identification of tetragametic chimerism" 2136: 1763:"Germ-line chimerism and paternal care in marmosets (Callithrix kuhlii)" 1130:

Lam, Susanna; Hultin, Sebastian; Preston, John; Campbell, Scott (2020).

872:

in China reported that they had successfully fused human skin cells and

630:

These are produced by grafting genetically different parents, different

555:

Chimerism Explained: How One Person Can Unknowingly Have Two Sets of DNA

538:

In nonidentical twins, innate chimerism occurs by means of blood vessel

3948: 3128:"First live birth of a chimeric monkey using embryonic stem cell lines" 2714: 1892: 1583: 1475: 1373: 1223: 974: 233: 3344: 2323:. United States Department of Commerce - Office of Technical Services. 1992: 3651:"Defining Humanity: The Ethics of Chimeric Animals and Organ Growing" 1881:"Which Half Is Mommy?: Tetragametic Chimerism and Trans-Subjectivity" 1026: 873: 846: 811: 467: 429: 380: 175: 141: 136:) is a single organism composed of cells with more than one distinct 2902:

Koonina, Eugene V.; Doljab, Valerian V.; Krupovic, Mart (May 2015).

1830:"In the Marmoset Family, Things Really Do Appear to Be All Relative" 1515:"Unilateral true hermaphrodite with 46,XX/46,XY dispermic chimerism" 2261:"Embryogenesis of chimeras, twins and anterior midline asymmetries" 3963: 898: 854: 783: 623: 618: 587: 376: 343: 287: 271: 43: 3954:

Embryogenesis of chimeras, twins and anterior midline asymmetries

3911:

Chimera: Apical Origin, Ontogeny and Consideration in Propagation

2426:

The plastids, their chemistry, structure, growth, and inheritance

1064:"The Stranger-Than-Fiction Story Of A Woman Who Was Her Own Twin" 140:. Animal chimeras can be produced by the merger of two (or more) 4011: 3994: 3590:. Methods in Molecular Biology. Vol. 158. pp. 135–54. 3575:

http://www.eucomm.org/docs/protocols/mouse_protocol_1_Sanger.pdf

3231:"Targeted mutation of the Hp gene in mouse embryonic stem cells" 2423:

Kirk, John Thomas Osmond; Tilney-Bassett, Richard A. E. (1978).

850: 347: 301: 3967: 307:

The tetragametic state has important implications for organ or

116: 78: 2429:(Rev. 2d ed.). Elsevier/North Holland Biomedical Press. 715:. The majority of variegated-leaf chimeras are of this kind. 601:

The distinction between sectorial, mericlinal and periclinal

517:

has been used at least since the 1944 article of Belgovskii.

342:

Germline chimerism occurs when the germ cells (for example,

3066:"This hybrid baby monkey is made of cells from two embryos" 119: 81: 69: 379:

composed of cells with one set of chromosomes and have a

107: 2333: 683:

These are chimeras in which the layers differ in their

776:

could be an alternative method for chimera detection.

394:. 95% of marmoset fraternal twins trade blood through 170:

Another way that chimerism can occur in animals is by

3758:

Nelson, J. Lee (Scientific American, February 2008).

2703:

In Vitro Cellular & Developmental Biology - Plant

2376:"Understanding Genetics: Human Health and the Genome" 128: 104: 90: 66: 3874:

Jones, David Albert; MacKellar, Calum, eds. (2012).

1319:"Crazy ants' strange genomes are a biological first" 1084:

Machácková, Ivana; Romanov, Georgy A. (2013-11-11).

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Journal of Pediatric Endocrinology & Metabolism

304:. As such, they can be male, female, or intersex. 113: 110: 75: 72: 818:, which usually infect birds and pigs, and a RNA 367:As the organism develops, it can come to possess 3229:Doetschman, T.; Maeda, N.; Smithies, O. (1988). 2835:Diemer, Geoffrey S., Kenneth M. (11 June 2013). 2150:formed of the cells of a single zygote lineage. 2041:"Chimaera - an overview | ScienceDirect Topics" 1855:"Marmosets may carry their sibling's sex cells" 1767:Proceedings of the National Academy of Sciences 1195:Blanquer, Andrea; Uriz, Maria-J. (2011-04-15). 477:In artificial chimerism, an individual has one 3474:Jasin, M; Moynahan, ME; Richardson, C (1996). 2306:The Embryo Project at Arizona State University 1256:"Obligate chimerism in male yellow crazy ants" 1117:The Embryo Project at Arizona State University 3979: 654:. Well-known examples of a graft-chimera are 8: 2300:Sarah Taddeo, Jason S. Robert (2014-11-04). 1190: 1188: 1111:Sarah Taddeo, Jason S. Robert (2014-11-04). 2831: 2829: 2827: 1761:Ross, C. N.; J. A. French; G. Orti (2007). 1756: 1754: 802:study of the acidic extreme environment of 592:Ficus with chlorophyll-deficient cell zones 3986: 3972: 3964: 2976:"Developmental Biology Cinema, Le Douarin" 2402:"The Twin Inside Me: Extraordinary People" 3856: 3789:U.S. Denies Patent for a too-human hybrid 3767:Cloning yields human-rabbit hybrid embryo 3732: 3550: 3509: 3499: 3264: 3254: 3040: 2919: 2807: 2797: 2780:Faize, M.; Faize, L.; Burgos, L. (2010). 2276: 2084: 1955: 1926:Masahito Tachibana, Michelle Sparman and 1874: 1872: 1870: 1804: 1786: 1734: 1641: 1619: 1617: 1546: 1222: 1212: 1157: 1147: 1090:. Springer Science & Business Media. 311:transplantation. Chimeras typically have 276:Chimeric trait distribution by generation 186:often determines the recipient's ensuing 3755:, Volume 19, Number 2, March–April 2007. 3535:"Embryonic Stem Cell and Gene Targeting" 3001:"Animal-Human Hybrids Spark Controversy" 2885:BioMed Central Limited (18 April 2012). 3927:Ainsworth, Claire (November 15, 2003). 1932:"Generation of Chimeric Rhesus Monkeys" 1054: 2544:Tilney-Bassett, Richard A. E. (1991). 2384:Stanford University School of Medicine 2350:National Society of Genetic Counselors 746:starts from the multicellular origin. 375:. For example, the chimera may have a 144:. In plants and some animal chimeras, 3749:Appel, Jacob M. "The Monster's Law", 2160: 2158: 7: 3457:Gene Targeting: A Practical Approach 3403:Gene Targeting: A Practical Approach 2073:European Journal of Medical Genetics 2016:"Half-siders: A tale of two birdies" 1879:Norton, Aaron; Ozzie Zehner (2008). 292:African violets exhibiting chimerism 255:Yellow crazy ant § Reproduction 211:Chimerism occurs naturally in adult 2871:"Scientists discover hybrid virus". 1630:The New England Journal of Medicine 868:In August 2003, researchers at the 474:. It is much rarer than mosaicism. 3634:Futehally, Ilmas, Beyond Biology, 2999:Mott, Maryann (January 25, 2005). 2952:. 27 February 1984. Archived from 2374:Starr, Barry (November 30, 2004). 2069:"Natural human chimeras: A review" 870:Shanghai Second Medical University 703:Plastid gene-differential chimeras 695:Nuclear gene-differential chimeras 386:This is particularly true for the 25: 4030:Twin-to-twin transfusion syndrome 3786:Weiss, Rick (February 13, 2005). 2853:Thompson, Helen (20 April 2012). 965:, and trophectoderm of the mouse 207:Symbiotic chimerism in anglerfish 4035:Twin reversed arterial perfusion 3959:Natural human chimeras: A review 3916:Plant Chimeras in Tissue Culture 3905:Chimerism and cellular mosaicism 3680:Brownback, Samuel (2005-03-17). 3302:10.1111/j.1399-0004.2005.00478.x 2183:10.1111/j.1420-9101.2004.00813.x 100: 62: 3845:New England Journal of Medicine 3764:Weiss, Rick (August 14, 2003). 2171:Journal of Evolutionary Biology 1853:Hooper, Rowan (26 March 2007). 1201:Molecular Biology and Evolution 428:that originated from different 3686:The Library of Congress THOMAS 2874:Journal of Young Investigators 2869:Devor, Caitlin (12 July 2012). 2408:. 9 March 2006. Archived from 1887:. Fall/Winter (3–4): 106–127. 420:An animal chimera is a single 199:Natural and obligate chimerism 1: 3649:Bruch, Quinton (2014-02-20). 3173:10.1126/science.158.3807.1484 2629:10.1126/science.254.5035.1214 2463:Agricultural Research Reports 2230:10.1016/s0198-8859(01)00249-x 2165:Santelices, B. (2004-11-01). 2067:Madan, Kamlesh (2020-09-01). 1680:10.1016/s0022-3476(94)70067-2 1317:Callaway, Ewen (2023-04-06). 880:. In 2007, scientists at the 857:, and survived to adulthood. 808:Lassen Volcanic National Park 3434:10.1016/0014-4827(85)90143-0 3216:10.1016/0168-9525(86)90161-7 3064:Wong, Carissa (2023-11-16). 2259:Boklage, Charles E. (2006). 1572:Indian Journal of Pediatrics 1022:Genetic chimerism in fiction 371:that have different sets of 3878:. London: Continuum Books. 3858:10.1056/NEJM199801153380305 2892:. Retrieved March 31, 2020. 2515:10.1126/science.220.4592.75 1828:Zimmer, Carl (2007-03-27). 1643:10.1056/NEJM199801153380305 1519:Journal of Medical Genetics 709:nucleic acid thermodynamics 646:, which is a fusion of the 409:, due to the many existing 32:Chimera (molecular biology) 4156: 3090:10.1038/d41586-023-03473-w 3042:10.1016/j.cell.2023.10.005 2876:. Retrieved 31 March 2020. 2860:. Retrieved 27 March 2020. 2844:. Retrieved 29 March 2020. 2552:Cambridge University Press 2457:van Harten, A. M. (1978). 2086:10.1016/j.ejmg.2020.103971 1948:10.1016/j.cell.2011.12.007 1331:10.1038/d41586-023-01002-3 994: 903:A chimeric mouse with her 845:was produced by combining 791: 661:, caused by a fusion of a 612: 451: 322: 252: 36: 29: 3636:Strategic Foresight Group 3596:10.1385/1-59259-220-1:135 3552:10.1017/S0958067000021059 2921:10.2535/ofaj1936.41.5_285 2751:10.1007/s00299-006-0273-8 2672:10.1007/s00299-007-0398-4 1885:Women's Studies Quarterly 1668:The Journal of Pediatrics 1425:10.1515/jpem.2009.22.1.97 770:Green Fluorescent Protein 742:(1) The process of shoot 713:non-mendelian inheritance 576:Another case was that of 411:plumage colour variations 3376:J. Embryol. Exp. Morphol 3005:National Geographic News 930:homologous recombination 48:Two-colored rose chimera 37:Not to be confused with 4063:Craniopagus parasiticus 3938:(subscription required) 3760:Your Cells Are My Cells 3655:The Triple Helix Online 3588:Gene Knockout Protocols 3539:Experimental Physiology 3501:10.1073/pnas.93.17.8804 3476:"Targeted transgenesis" 3461:Oxford University Press 3407:Oxford University Press 3256:10.1073/pnas.85.22.8583 2799:10.1186/1472-6750-10-53 1788:10.1073/pnas.0607426104 1280:10.1126/science.adf0419 470:in the early stages of 259:It has been shown that 1727:10.4161/chim.1.2.14294 991:Ethics and legislation 912: 909:agouti coat color gene 789: 627: 593: 544:in vitro fertilisation 444:have been documented. 415:half-sider budgerigars 293: 284:Tetragametic chimerism 277: 232:as their other organs 49: 4125:Intersex and medicine 3929:"The Stranger Within" 3900:"Chimerism Explained" 3816:10.1007/s002770050543 3533:Ledermann, B (2000). 3235:Proc. Natl. Acad. Sci 3035:(23): 4996–5014.e24. 2278:10.1093/humrep/dei370 2045:www.sciencedirect.com 1531:10.1136/jmg.24.12.784 1214:10.1093/molbev/msr096 982:Methods of production 902: 814:is related to a DNA 787: 622: 591: 483:organ transplantation 398:fusions, making them 313:immunologic tolerance 291: 275: 47: 3804:Annals of Hematology 3734:10.1056/NEJMoa013452 3011:on January 27, 2005. 2956:on February 16, 2008 1149:10.1155/2020/7461052 926:embryonic stem cells 882:University of Nevada 804:Boiling Springs Lake 679:Chromosomal chimeras 472:prenatal development 315:to both cell lines. 268:Artificial chimerism 220:olfactory (or smell) 4022:Monochorionic twins 3795:The Washington Post 3780:The Washington Post 3492:1996PNAS...93.8804J 3247:1988PNAS...85.8583D 3165:1967Sci...158.1484M 3082:2023Natur.623..468W 2621:1991Sci...254.1214T 2615:(5035): 1214–1216. 2507:1983Sci...220...75N 2137:10.1038/hdy.1951.38 1928:Shoukhrat Mitalipov 1779:2007PNAS..104.6278R 1674:(5 Pt 1): 738–744. 1272:2023Sci...380...55D 1136:Case Rep Transplant 911:; note her pink eye 889:crab-eating macaque 497:human–animal hybrid 448:Chimerism in humans 434:sexual reproduction 39:Chimera (mythology) 4040:Monoamniotic twins 3947:. (Reprinted here 3943:2008-10-22 at the 3921:2013-10-03 at the 3773:2012-10-25 at the 2715:10.1007/BF02822775 2265:Human Reproduction 1893:10.1353/wsq.0.0115 1834:The New York Times 1584:10.1007/bf02976984 1476:10.1007/bf00290226 1374:10.1007/BF00289485 1062:Friedman, Lauren. 955:primitive endoderm 940:Underlying biology 913: 907:, which carry the 790: 628: 594: 566:hitchhiker's thumb 338:Germline chimerism 302:nonidentical twins 294: 278: 50: 4130:Genetic anomalies 4107: 4106: 3605:978-1-59259-220-3 3486:(17): 8804–8808. 3345:10.1242/dev.00893 3339:(25): 6155–6163. 3241:(22): 8583–8587. 3076:(7987): 468–469. 2786:BMC Biotechnology 2666:(10): 1727–1732. 2589:. 17 January 2018 2587:extension.unh.edu 2561:978-0-521-42787-6 2472:978-90-220-0667-2 1993:10.1159/000129828 1773:(15): 6278–6282. 1097:978-94-017-2664-1 648:Florentine citron 605:are widely used. 487:blood transfusion 261:yellow crazy ants 54:genetic chimerism 16:(Redirected from 4147: 3988: 3981: 3974: 3965: 3939: 3889: 3870: 3860: 3835: 3746: 3736: 3702: 3701: 3699: 3697: 3688:. Archived from 3677: 3671: 3670: 3668: 3666: 3657:. Archived from 3646: 3640: 3632: 3626: 3625: 3583: 3577: 3571: 3565: 3564: 3554: 3530: 3524: 3523: 3513: 3503: 3471: 3465: 3464: 3452: 3446: 3445: 3417: 3411: 3410: 3398: 3392: 3391: 3371: 3365: 3364: 3328: 3322: 3321: 3285: 3279: 3278: 3268: 3258: 3226: 3220: 3219: 3199: 3193: 3192: 3159:(3807): 1484–6. 3148: 3142: 3141: 3139: 3138: 3124: 3118: 3117: 3061: 3055: 3054: 3044: 3019: 3013: 3012: 3007:. Archived from 2996: 2990: 2989: 2987: 2986: 2972: 2966: 2965: 2963: 2961: 2940: 2934: 2933: 2923: 2899: 2893: 2883: 2877: 2867: 2861: 2851: 2845: 2833: 2822: 2821: 2811: 2801: 2777: 2771: 2770: 2733: 2727: 2726: 2698: 2692: 2691: 2655: 2649: 2648: 2604: 2598: 2597: 2595: 2594: 2579: 2573: 2572: 2570: 2568: 2541: 2535: 2534: 2490: 2484: 2483: 2481: 2479: 2454: 2448: 2447: 2445: 2443: 2420: 2414: 2413: 2412:on May 26, 2006. 2406:Channel 5 TV, UK 2398: 2392: 2391: 2386:. Archived from 2380:Ask a Geneticist 2371: 2365: 2364: 2362: 2361: 2352:. Archived from 2342: 2336: 2331: 2325: 2324: 2316: 2310: 2309: 2297: 2291: 2290: 2280: 2256: 2250: 2249: 2218:Human Immunology 2213: 2207: 2206: 2177:(6): 1187–1188. 2162: 2153: 2152: 2116: 2110: 2109: 2088: 2064: 2058: 2057: 2052: 2051: 2037: 2031: 2030: 2028: 2026: 2011: 2005: 2004: 1976: 1970: 1969: 1959: 1930:(January 2012). 1923: 1914: 1911: 1905: 1904: 1876: 1865: 1864: 1850: 1844: 1843: 1841: 1840: 1825: 1819: 1818: 1808: 1790: 1758: 1749: 1748: 1738: 1706: 1700: 1699: 1662: 1656: 1655: 1645: 1621: 1612: 1611: 1578:(401): 885–890. 1567: 1561: 1560: 1550: 1510: 1504: 1503: 1459: 1453: 1452: 1408: 1402: 1401: 1357: 1351: 1350: 1314: 1308: 1307: 1251: 1245: 1244: 1226: 1216: 1207:(9): 2435–2438. 1192: 1183: 1178: 1172: 1171: 1161: 1151: 1127: 1121: 1120: 1108: 1102: 1101: 1081: 1075: 1074: 1072: 1070: 1059: 932:, thus allowing 916:cell potential. 774:Quantitative PCR 571:Blaschko's lines 511:Blaschko's lines 152:during ordinary 132: 126: 125: 122: 121: 118: 115: 112: 109: 106: 94: 88: 87: 84: 83: 80: 77: 74: 71: 68: 21: 4155: 4154: 4150: 4149: 4148: 4146: 4145: 4144: 4110: 4109: 4108: 4103: 4077: 4050:Conjoined twins 4044: 4016: 3998: 3992: 3945:Wayback Machine 3937: 3923:Wayback Machine 3896: 3886: 3873: 3838: 3801: 3775:Wayback Machine 3727:(20): 1545–52. 3714: 3711: 3709:Further reading 3706: 3705: 3695: 3693: 3679: 3678: 3674: 3664: 3662: 3648: 3647: 3643: 3633: 3629: 3606: 3585: 3584: 3580: 3572: 3568: 3532: 3531: 3527: 3473: 3472: 3468: 3459:. 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For example, 130: 103: 99: 92: 65: 61: 42: 35: 28: 23: 22: 15: 12: 11: 5: 4153: 4151: 4143: 4142: 4137: 4132: 4127: 4122: 4112: 4111: 4105: 4104: 4102: 4101: 4096: 4091: 4089:Vanishing twin 4085: 4083: 4079: 4078: 4076: 4075: 4074: 4073: 4065: 4058:Parasitic twin 4054: 4052: 4046: 4045: 4043: 4042: 4037: 4032: 4026: 4024: 4018: 4017: 4015: 4014: 4008: 4006: 4000: 3999: 3993: 3991: 3990: 3983: 3976: 3968: 3962: 3961: 3956: 3951: 3925: 3913: 3908: 3902: 3895: 3894:External links 3892: 3891: 3890: 3884: 3871: 3836: 3810:(9): 431–434. 3799: 3784: 3762: 3756: 3747: 3710: 3707: 3704: 3703: 3672: 3641: 3627: 3604: 3578: 3566: 3545:(6): 603–613. 3525: 3466: 3447: 3428:(2): 561–566. 3412: 3393: 3382:(2): 283–290. 3366: 3323: 3296:(2): 106–112. 3280: 3221: 3194: 3143: 3119: 3056: 3014: 2991: 2967: 2935: 2914:(5): 285–293. 2894: 2878: 2862: 2846: 2841:Biology Direct 2823: 2772: 2745:(5): 661–671. 2739:Plant Cell Rep 2728: 2709:(2): 117–121. 2693: 2660:Plant Cell Rep 2650: 2599: 2574: 2560: 2547:Plant Chimeras 2536: 2485: 2471: 2449: 2435: 2415: 2393: 2390:on 2011-07-24. 2366: 2337: 2326: 2311: 2292: 2271:(3): 579–591. 2251: 2224:(6): 651–657. 2208: 2154: 2131:(3): 379–397. 2111: 2059: 2032: 2006: 1987:(6): 384–393. 1971: 1915: 1906: 1866: 1845: 1820: 1750: 1701: 1657: 1636:(3): 166–169. 1613: 1562: 1505: 1470:(3): 407–408. 1464:Human Genetics 1454: 1403: 1362:Human Genetics 1352: 1309: 1246: 1184: 1173: 1122: 1103: 1096: 1076: 1053: 1052: 1050: 1047: 1046: 1045: 1040: 1037:X-inactivation 1034: 1032:Vanishing twin 1029: 1024: 1019: 1012: 1009: 992: 989: 983: 980: 941: 938: 934:gene targeting 921:Beatrice Mintz 896: 895:Work with mice 893: 830: 827: 792:Main article: 781: 778: 760: 757: 736: 733: 728:photosynthetic 704: 701: 696: 693: 680: 677: 657:Laburnocytisus 615:Graft-chimaera 613:Main article: 610: 609:Graft chimeras 607: 603:plant chimeras 598: 595: 585: 582: 458:Some consider 452:Main article: 449: 446: 442:human chimeras 364: 361: 359: 356: 339: 336: 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