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Somatic cell nuclear transfer

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methods of induced pluripotent stem cells. Though recent studies have put in question how similar iPS cells are to embryonic stem cells. Epigenetic memory in iPS affects the cell lineage it can differentiate into. For instance, an iPS cell derived from a blood cell using only the yamanaka factors will be more efficient at differentiating into blood cells, while it will be less efficient at creating a neuron. Recent studies indicate however that changes to the epigenetic memory of iPSCs using small molecules can reset them to an almost naive state of pluripotency. Studies have even shown that via tetraploid complementation, an entire viable organism can be created solely from iPSCs. SCNT stem cells have been found to have similar challenges. The cause for low yields in bovine SCNT cloning has, in recent years, been attributed to the previously hidden epigenetic memory of the somatic cells that were being introduced into the oocyte.
355:), and has been proposed as a possible way to clone humans. Using SCNT in reproductive cloning has proven difficult with limited success. High fetal and neonatal death make the process very inefficient. Resulting cloned offspring are also plagued with development and imprinting disorders in non-human species. For these reasons, along with moral and ethical objections, reproductive cloning in humans is proscribed in more than 30 countries. Most researchers believe that in the foreseeable future it will not be possible to use the current cloning technique to produce a human clone that will develop to term. It remains a possibility, though critical adjustments will be required to overcome current limitations during early embryonic development in human SCNT. 594:
several eggs at once). Although such treatments have been performed for several decades now, the long-term effects have not been studied or declared safe to use on a large scale on otherwise healthy women. Longer-term treatments with much lower doses of hormones are known to increase the rate of cancer decades later. Whether hormone treatments to induce hyper-ovulation could have similar effects is unknown. There are also ethical questions surrounding paying for eggs. In general, marketing body parts is considered unethical and is banned in most countries. Human eggs have been a notable exception to this rule for some time.
317:. The imperfect embryos prevented the acquisition of human ESC. The addition of caffeine during the removal of the ovum's nucleus and fusion of the somatic cell and the egg improved blastocyst formation and ESC isolation. The ESC obtain were found to be capable of producing teratomas, expressed pluripotent transcription factors, and expressed a normal 46XX karyotype, indicating these SCNT were in fact ESC-like. This was the first instance of successfully using SCNT to reprogram human somatic cells. This study used fetal and infantile somatic cells to produce their ESC. 525:
epigenetic limitations to SCNT is regulating histone methylation. Differing regulation of these histone methylation genes can directly affect the transcription of the developing genome, causing failure of the SCNT. Another contributing factor to failure of SCNT includes the X chromosome inactivation in early development of the embryo. A non coding gene called XIST is responsible for inactivating one X chromosome during development, however in SCNT this gene can have abnormal regulation causing mortality to the developing fetus.
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nuclei into cells with pluripotent states. When the ovum's nucleus is removed, the cell loses its genetic information. This has been blamed for why enucleated eggs are hampered in their reprogramming ability. It is theorized the critical embryonic genes are physically linked to oocyte chromosomes, enucleation negatively affects these factors. Another possibility is removing the egg nucleus or inserting the somatic nucleus causes damage to the cytoplast, affecting reprogramming ability.
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transplanted for treatment. For example, to treat a man with Parkinson's disease, a cell nucleus from one of his cells would be transplanted by SCNT into an egg cell from an egg donor, creating a unique lineage of stem cells almost identical to the patient's own cells. (There would be differences. For example, the mitochondrial DNA would be the same as that of the egg donor. In comparison, his own cells would carry the mitochondrial DNA of his mother.)
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cells of the human body. Skin cells, fat cells, and liver cells are only a few examples. The genetic material of the donor egg cell is removed and discarded, leaving it 'deprogrammed.' What is left is a somatic cell and an enucleated egg cell. These are then fused by inserting the somatic cell into the 'empty' ovum. After being inserted into the egg, the somatic cell nucleus is
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differentiation. When injected into mice, cells of all three of the germ layers successfully formed. The most significant of these cells, were those who expressed insulin and were capable of secreting the hormone. These insulin producing cells could be used for replacement therapy in diabetics, demonstrating real SCNT stem cell therapeutic potential.
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tissues and organs for therapy, as there may be an immuno-response to the non-self mtDNA after transplant. Additionally, the genes found in the mitochondria’s genome need to communicate with the cell’s genome and a failure of somatic cell nuclear reprogramming can lead to non communication to the cell’s genome causing SCNT to fail.
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blastocyst; the cells fail to progress past the eight cell stage of development. This is thought to be a result from the somatic cell nucleus being unable to turn on embryonic genes crucial for proper development. These earlier experiments used procedures developed in non-primate animals with little success.
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One concern is that blastula creation in SCNT-based human stem cell research will lead to the reproductive cloning of humans. Both processes use the same first step: the creation of a nuclear transferred embryo, most likely via SCNT. Those who hold this concern often advocate for strong regulation of
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Epigenetic factors play an important role in the success or failure of SCNT attempts. The varying gene expression of a previously activated cell and its mRNAs may lead to overexpression, underexpression, or in some cases non functional genes which will affect the developing fetus. One such example of
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also has to be extremely precise, as most late term cloned fetus deaths are the result of inadequate placentation. However, by 2014, researchers were reporting success rates of 70-80% with cloning pigs and in 2016 a Korean company, Sooam Biotech, was reported to be producing 500 cloned embryos a day.
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were developed in humans in 2007. The following year, this method achieved a key goal of SCNT-based stem cell research: the derivation of pluripotent stem cell lines that have all genes linked to various diseases. Some scientists working on SCNT-based stem cell research have recently moved to the new
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An alternative technique called "handmade cloning" was described by Indian scientists in 2001. This technique requires no use of a micromanipulator and has been used for the cloning of several livestock species. Removal of the nucleus can be done chemically, by centrifuge, or with the use of a blade.
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by its host egg cell. The ovum, now containing the somatic cell's nucleus, is stimulated with a shock and will begin to divide. The egg is now viable and capable of producing an adult organism containing all necessary genetic information from just one parent. Development will ensue normally and after
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In April 2014, an international research team expanded on this break through. There remained the question of whether the same success could be accomplished using adult somatic cells. Epigenetic and age related changes were thought to possibly hinder an adult somatic cells ability to be reprogrammed.
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Taking this into account the research group applied their new technique in an attempt to produce human SCNT stem cells. In May 2013, the Oregon group reported the successful derivation of human embryonic stem cell lines derived through SCNT, using fetal and infant donor cells. Using MII oocytes from
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Yamada, M; Johannesson, B; Sagi, I; Burnett, L. C.; Kort, D. H.; Prosser, R. W.; Paull, D; Nestor, M. W.; Freeby, M; Greenberg, E; Goland, R. S.; Leibel, R. L.; Solomon, S. L.; Benvenisty, N; Sauer, M. V.; Egli, D (2014). "Human oocytes reprogram adult somatic nuclei of a type 1 diabetic to diploid
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In the United States, the practice remains legal, as it has not been addressed by federal law. However, in 2002, a moratorium on United States federal funding for SCNT prohibits funding the practice for the purposes of research. Thus, though legal, SCNT cannot be federally funded. American scholars
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Stem cell experts consider it unlikely that such large numbers of human egg donations would occur in a developed country because of the unknown long-term public health effects of treating large numbers of healthy young women with heavy doses of hormones in order to induce hyper-ovulation (ovulating
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Potentially millions of patients could benefit from stem cell therapy, and each patient would require a large number of donated eggs in order to successfully create a single custom therapeutic stem cell line. Such large numbers of donated eggs would exceed the number of eggs currently left over and
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are left behind. The resulting hybrid cells retain those mitochondrial structures which originally belonged to the egg. As a consequence, clones such as Dolly that are born from SCNT are not perfect copies of the donor of the nucleus. This fact may also hamper the potential benefits of SCNT-derived
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which typically is between domestic animals and rodents, or where there is a ready supply of oocytes and surrogate animals. However, the cloning of highly endangered or extinct species requires the use of an alternative method of cloning. Interspecies nuclear transfer utilizes a host and a donor of
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Though there has been numerous successes with cloning animals, questions remain concerning the mechanisms of reprogramming in the ovum. Despite many attempts, success in creating human nuclear transfer embryonic stem cells has been limited. There lies a problem in the human cell's ability to form a
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are undifferentiated cells of an embryo. These cells are deemed to have a pluripotent potential because they have the ability to give rise to all of the tissues found in an adult organism. This ability allows stem cells to create any cell type, which could then be transplanted to replace damaged or
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Somatic cell nuclear transfer is a technique for cloning in which the nucleus of a somatic cell is transferred to the cytoplasm of an enucleated egg. After the somatic cell transfers, the cytoplasmic factors affect the nucleus to become a zygote. The blastocyst stage is developed by the egg to help
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somatic cell nucleus and activating the recipient egg was also far from understood. Another limitation is trying to use one-cell embryos during the SCNT. When using just one-cell cloned embryos, the experiment has a 65% chance to fail in the process of making morula or blastocyst. The biochemistry
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There is also the potential for treating diseases associated with mutations in mitochondrial DNA. Recent studies show SCNT of the nucleus of a body cell afflicted with one of these diseases into a healthy oocyte prevents the inheritance of the mitochondrial disease. This treatment does not involve
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The process of somatic cell nuclear transfer involves two different cells. The first being a female gamete, known as the ovum (egg/oocyte). In human SCNT experiments, these eggs are obtained through consenting donors, utilizing ovarian stimulation. The second being a somatic cell, referring to the
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Somatic cell nuclear transfer (SCNT) can be inefficient due to stresses placed on both the egg cell and the introduced nucleus. This can result in a low percentage of successfully reprogrammed cells. For example, in 1996 Dolly the sheep was born after 277 eggs were used for SCNT, which created 29
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utilizing the principles of SCNT. In short, the experiment consisted of inducing a female specimen to ovulate, at which point her eggs were harvested. From here, the egg was enucleated using ultra-violet irradiation to disable the egg's pronucleus. At this point, the prepared egg cell and nucleus
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demonstrated SCNT procedures developed for primates successfully using skin cells. The key to their success was utilizing oocytes in metaphase II (MII) of the cell cycle. Egg cells in MII contain special factors in the cytoplasm that have a special ability in reprogramming implanted somatic cell
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One vision for successful stem cell therapies is to create custom stem cell lines for patients. Each custom stem cell line would consist of a collection of identical stem cells each carrying the patient's own DNA, thus reducing or eliminating any problems with rejection when the stem cells were
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Late April 2014, the New York Stem Cell Foundation was successful in creating SCNT stem cells derived from adult somatic cells. One of these lines of stem cells was derived from the donor cells of a type 1 diabetic. The group was then able to successfully culture these stem cells and induce
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Kim, K.; Doi, A.; Wen, B.; Ng, K.; Zhao, R.; Cahan, P.; Kim, J.; Aryee, M. J.; Ji, H.; Ehrlich, L. I. R.; Yabuuchi, A.; Takeuchi, A.; Cunniff, K. C.; Hongguang, H.; McKinney-Freeman, S.; Naveiras, O.; Yoon, T. J.; Irizarry, R. A.; Jung, N.; Seita, J.; Hanna, J.; Murakami, P.; Jaenisch, R.;
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with about 100 cells) with an identical genome to the original organism (i.e. a clone). Stem cells can then be obtained by the destruction of this clone embryo for use in therapeutic cloning or in the case of reproductive cloning the clone embryo is implanted into a host mother for further
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viable embryos, giving it a measly 0.3% efficiency. Only three of these embryos survived until birth, and only one survived to adulthood. Millie, the offspring that survived, took 95 attempts to produce. Because the procedure was not automated and had to be performed manually under a
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Another application of SCNT stem cell research is using the patient specific stem cell lines to generate tissues or even organs for transplant into the specific patient. The resulting cells would be genetically identical to the somatic cell donor, thus avoiding any complications from
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To address the problem of creating a human egg market, some stem cell researchers are investigating the possibility of creating artificial eggs. If successful, human egg donations would not be needed to create custom stem cell lines. However, this technology may be a long way off.
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and sucks out the egg cell's original nucleus using a pipette. They then make another opening to a different pipette to inject the donor nucleus. Alternatively, electric energy can be applied to fuse the empty egg cell with a donor cell containing a nucleus.
590:. Therefore, healthy young women would need to be induced to sell eggs to be used in the creation of custom stem cell lines that could then be purchased by the medical industry and sold to patients. It is so far unclear where all these eggs would come from. 574:, which can only be obtained from women. The most common source of these eggs today are eggs that are produced and in excess of the clinical need during IVF treatment. This is a minimally invasive procedure, but it does carry some health risks, such as 321:
Implementing the procedure pioneered by the Oregon research group they indeed were able to grow stem cells generated by SCNT using adult cells from two donors aged 35 and 75, indicating that age does not impede a cell's ability to be reprogrammed.
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from the donor cell were combined, and then incubation and eventual development into a tadpole proceeded. Gurdon's application of SCNT differs from more modern applications and even applications used on other model systems of the time (i.e.,
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Selokar, Naresh L.; Saini, Monika; Palta, Prabhat; Chauhan, Manmohan S.; Manik, Radhey S.; Singla, Suresh K. (April 2018). "Cloning of Buffalo, a Highly Valued Livestock Species of South and Southeast Asia: Any Achievements?".
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Interspecies nuclear transfer (iSCNT) is a means of somatic cell nuclear transfer being used to facilitate the rescue of endangered species, or even to restore species after their extinction. The technique is similar to SCNT
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donated their somatic cells, the stem cells resulting from SCNT would have genes that contribute to Parkinson's disease. The disease specific stem cell lines could then be studied in order to better understand the condition.
500:, SCNT was very resource intensive. Another reason why there is such high mortality rate with the cloned offspring is due to the fetus being larger than even other large offspring, resulting in death soon after birth. The 1572: 633:, calling on member states to "prohibit all forms of human cloning in as much as they are incompatible with human dignity and the protection of human life." This phrase may include SCNT, depending on interpretation. 2879:
Additional Protocol to the Convention for the Protection of Human Rights and Dignity of the Human Being with regard to the Application of Biology and Medicine, on the Prohibition of Cloning Human Being
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Additional Protocol to the Convention for the Protection of Human Rights and Dignity of the Human Being with regard to the Application of Biology and Medicine, on the Prohibition of Cloning Human Being
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Zhao, X. Y.; Li, W.; Lv, Z.; Liu, L.; Tong, M.; Hai, T.; Hao, J.; Guo, C. L.; Ma, Q. W.; Wang, L.; Zeng, F.; Zhou, Q. (2009). "IPS cells produce viable mice through tetraploid complementation".
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The impetus for SCNT-based stem cell research has been decreased by the development and improvement of alternative methods of generating stem cells. Methods to reprogram normal body cells into
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volunteers and their improved SCNT procedure, human clone embryos were successfully produced. These embryos were of poor quality, lacking a substantial inner cell mass and poorly constructed
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cloning but would produce a child with three genetic parents. A father providing a sperm cell, one mother providing the egg nucleus, and another mother providing the enucleated egg cell.
2326:"First cloned Bactrian camel (Camelus bactrianus) calf produced by interspecies somatic cell nuclear transfer: A step towards preserving the critically endangered wild Bactrian camels" 540: 230:
surrounds human ESC work due to the destruction of viable human embryos, leading scientists to seek alternative methods of obtaining pluripotent stem cells, SCNT is one such method.
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matched the donor organism from which they came. This gives them the ability to create patient specific pluripotent cells, which could then be used in therapies or disease research.
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model could be created, would be useful for studying that particular disease, potentially discovering its pathophysiology, and discovering therapies. For example, if a person with
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Although Dolly is generally recognized as the first animal to be cloned using this technique, earlier instances of SCNT exist as early as the 1950s. In particular, the research of
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Convention for the Protection of Human Rights and Dignity of the Human Being with regard to the Application of Biology and Medicine: Convention on Human Rights and Biomedicine
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Nuclear transfer techniques present a different set of ethical considerations than those associated with the use of other stem cells like embryonic stem cells which are
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Lanza, Robert P.; Jose B. Cibelli; Francisca A. Diaz; Carlos T. Moraes; Peter W. Farin; Charlotte E. Farin; Carolyn J. Hammer; Michael D. West; Philip Damiani (2000).
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A potential use of stem cells genetically matched to a patient would be to create cell lines that have genes linked to a patient's particular disease. By doing so, an
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have recently argued that because the product of SCNT is a clone embryo, rather than a human embryo, these policies are morally wrong and should be revised.
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create embryonic stem cells from the inner cell mass of the blastocyst. The first mammal to be developed by this technique was Dolly the sheep, in 1996.
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National Institutes of Health, Paper giving background information on cloning in general and SCNT from The Office of Science Policy Analysis.
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Wakayama, Sayaka; Mizutani, Eiji; Wakayama, Teruhiko (2010). "Production of Cloned Mice from Somatic Cells, ES Cells, and Frozen Bodies".
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became famous for being the first successful case of the reproductive cloning of a mammal. In January 2018, a team of scientists in
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for their requirement to destroy an embryo. These different considerations have led to some individuals and organizations who are
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Cunningham, Thomas V (2013). "What justifies the United States ban on federal funding for nonreproductive cloning?".
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SCNT to preclude implantation of any derived products for the intention of human reproduction, or its prohibition.
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Only a handful of the labs in the world are currently using SCNT techniques in human stem cell research. In the
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A second important concern is the appropriate source of the eggs that are needed. SCNT requires human
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opposed to human embryonic stem cell research to be concerned about, or opposed to, SCNT research.
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Somatic cell nuclear transfer can create clones for both reproductive and therapeutic purposes.
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In SCNT, not all of the donor cell's genetic information is transferred, as the donor cell's
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two different organisms that are closely related species and within the same genus. In 2000,
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are currently researching a technique to use somatic cell nuclear transfer to produce
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from a somatic (body) cell. It is used in both therapeutic and reproductive
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appear to ban SCNT of human beings. Of the Council's 45 member states, the
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SCNT involving human cells is currently legal for research purposes in the
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BTX ECM 2001 electrofusion generator used for SCNT and Cloning applications
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and skin fibroblast cells of an adult Bactrian camel as donor nuclei.
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Nuclear Transfer – Stem Cells or Somatic Cell Nuclear Transfer (SCNT)
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1346:"Cloning debate: Stem-cell researchers must stay engaged" 655:
has been signed by 29 member nations and ratified by 14.
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using somatic cell nuclear transfer, the same method as
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available from couples trying to have children through
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Tian XC, Kubota C, Enright B, Yang X (November 2003).
2894: 862: 2975: 1684:"Epigenetic memory in induced pluripotent stem cells" 1555:
Human Somatic Cell Nuclear Transfer Using Adult Cells
3197:
List of varieties of genetically modified maize/corn
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Some for Abortion Rights Lean Right in Cloning Fight
2376: 3021:Gurdon JB, Byrne JA, Simonsson S (September 2003). 2211:"First monkey clones created in Chinese laboratory" 2490:The International Journal of Developmental Biology 1881: 1987:"Human somatic cell nuclear transfer and cloning" 1633: 956:Pattinson, Shaun D.; Kind, Vanessa (2017-09-12). 414:, combining it successfully with a domestic cow, 289:has granted permission to research groups at the 3935: 3104:The International Society for Stem Cell Research 2323: 601: 391: 169:many mitotic divisions, the single cell forms a 2451: 2089: 1875: 1824: 1642:"Breakthrough of the year: Reprogramming Cells" 1528:(September 28, 2004, retrieved October 6, 2006) 762: 760: 82:announced the successful cloning of two female 3023:"Nuclear reprogramming and stem cell creation" 2733:(November 23, 2001, retrieved October 6, 2006) 2727:Cloning opponents fear loopholes in new UK law 2230: 1789: 1732: 1639: 1401:Hadjantonakis AK, Papaioannou VE (July 2002). 722: 651:has been signed by 31 and ratified by 18. The 194:The empty egg is glued to the donor cell with 3128: 2882:(January 12, 1998, retrieved October 6, 2006) 1795: 1511:(January 17, 2006, retrieved October 6, 2006) 955: 2202: 1450: 1448: 1343: 1153: 1008:; Schnieke, A. E.; McWhir, J.; Kind, A. J.; 836:"Somatic Cell Nuclear Transfer - Embryology" 757: 616:Human Fertilisation and Embryology Authority 374:, with the birth of two live female clones ( 287:Human Fertilisation and Embryology Authority 3770:Detection of genetically modified organisms 2946: 2587:International Journal of Molecular Sciences 2396:American Journal of Reproductive Immunology 2174: 1929: 1927: 1830: 1680: 1454: 1195: 1074:Journal of Animal Science and Biotechnology 951: 949: 612:Human Fertilisation and Embryology Act 1990 3135: 3121: 2814: 2699: 767:Liu, Zhen; et al. (24 January 2018). 641:Convention on Human Rights and Biomedicine 347:This technique is currently the basis for 3052: 3042: 3003: 2993: 2775: 2764:The Journal of Law, Medicine & Ethics 2761: 2687:Open Letter to US Senate on Human Cloning 2638: 2636: 2616: 2598: 2359: 2349: 2285: 2155: 2127: 2002: 1899: 1764: 1738: 1715: 1657: 1545:(January 2003, retrieved October 6, 2006) 1480: 1445: 1428: 1418: 1369: 1309: 1262: 1213: 1095: 1085: 981: 786: 766: 740: 483:Learn how and when to remove this message 425:was born through iSCNT, using oocytes of 97:" refers to the potential use of SCNT in 3083:The Basics: Stem Cells and Public Policy 2705:Lori B. Andrews et al. (March 19, 2002). 2662: 1924: 1236: 946: 538: 406:was able to produce a cloned fetus of a 154: 18: 3142: 2719: 2457: 2097: 2025: 1339: 1337: 1284: 1282: 912:Angewandte Chemie International Edition 614:. Permission must be obtained from the 336: 297:. SCNT may also be occurring in China. 3936: 2633: 2446:China cloning on an 'industrial scale' 2236: 2208: 1288: 1239:"Ethical issues in stem cell research" 905: 306:Oregon Health & Science University 267:Oregon Health & Science University 263:University of California San Francisco 181:Conventional SCNT requires the use of 3116: 2576: 2574: 2180: 1497: 1063: 1061: 1059: 618:in order to perform or attempt SCNT. 206: 125: 59:. The technique consists of taking a 2817:Medicine, Health Care and Philosophy 2736: 2549:10.1016/j.theriogenology.2022.06.030 2181:Maron, Dina Fine (24 January 2018). 2128:Cyranoski, David (24 January 2018). 1745:Cellular and Molecular Life Sciences 1531: 1334: 1279: 1196:Lomax, G. P.; Dewitt, N. D. (2013). 858: 856: 610:, having been incorporated into the 465:adding citations to reliable sources 436: 2947:Kikyo N, Wolffe AP (January 2000). 2864: 2098:Normile, Dennis (24 January 2018). 1571:Ariana Eunjung Cha (18 April 2014) 1514: 13: 3800:Genetic use restriction technology 3096:Cloning: present uses and promises 3029:. 100. Suppl 1 (90001): 11819–22. 2888: 2851: 2571: 1739:Qin, H.; Zhao, A.; Fu, X. (2017). 1311:10.1111/j.1600-0463.2005.apm_312.x 1056: 690:New Jersey legislation S1909/A2840 66:(egg cell) and implanting a donor 14: 3985: 3086:The Century Foundation, June 2005 3070: 2458:Zastrow, Mark (8 February 2016). 2444:Shukman, David (14 January 2014) 2209:Briggs, Helen (24 January 2018). 1522:Dolly scientists' human clone bid 853: 576:ovarian hyperstimulation syndrome 178:development and brought to term. 2786:10.1111/j.1748-720x.2010.00479.x 2408:10.1034/j.1600-0897.2003.00064.x 2237:Kolata, Gina (24 January 2018). 2004:10.1016/j.fertnstert.2012.06.045 1640:Gretchen Vogel (December 2008). 629:adopted a proposal submitted by 588:assisted reproductive technology 441: 134:in 1958 entailed the cloning of 3818:Cartagena Protocol on Biosafety 2897:"Somatic cell nuclear transfer" 2808: 2755: 2679: 2524: 2477: 2438: 2317: 2258: 2046: 2019: 1978: 1674: 1581: 1565: 1548: 1420:10.1186/gb-2002-3-8-reviews1023 1394: 1230: 1189: 1112: 452:needs additional citations for 201: 116: 47:strategy for creating a viable 3090:Research Cloning Basic Science 2659:. Retrieved on August 7, 2006. 998: 906:Gurdon, John B. (2013-12-05). 899: 828: 803: 716: 528: 504:involved in reprogramming the 432: 362:In 2018, the first successful 113:the two cell types truly are. 1: 3580:Somatic cell nuclear transfer 1798:Accounts of Chemical Research 1659:10.1126/science.322.5909.1766 1344:Pera, M; Trounson, A (2013). 1129:10.1016/S0076-6879(10)76009-2 710: 602:Policies regarding human SCNT 392:Interspecies nuclear transfer 37:somatic cell nuclear transfer 3027:Proc. Natl. Acad. Sci. U.S.A 2351:10.1371/journal.pone.0177800 1810:10.1021/acs.accounts.7b00020 1539:The First Cloning Superpower 351:animals (such as the famous 7: 2716:Retrieved on August 7, 2006 658: 259:Harvard Stem Cell Institute 10: 3990: 2157:10.1038/d41586-018-01027-z 1473:10.1016/j.cell.2013.05.006 1202:Stem Cells and Development 840:embryology.med.unsw.edu.au 788:10.1016/j.cell.2018.01.020 532: 421:In 2017, the first cloned 340: 304:A research group from the 150: 3903: 3872: 3826: 3808: 3755: 3725: 3693: 3638:Genetically modified food 3630: 3623: 3593: 3540: 3531: 3494: 3447: 3397: 3388: 3318: 3300: 3282: 3264: 3241: 3223: 3205: 3172: 3163: 3150: 2829:10.1007/s11019-013-9465-5 2685:Lori B. Andrews, et al., 1757:10.1007/s00018-017-2586-x 1589:pluripotent stem cells". 1578:. Retrieved 18 April 2014 1562:. Retrieved 18 April 2014 1237:Lo, B; Parham, L (2009). 1087:10.1186/s40104-015-0043-y 962:Medical Law International 295:Newcastle Centre for Life 2982:Reprod. Biol. Endocrinol 1505:Cloning race is on again 974:10.1177/0968533217726350 275:Advanced Cell Technology 3944:Cell culture techniques 3044:10.1073/pnas.1834207100 2645:. (February 18, 2002). 2296:10.1089/152045500436104 2114:10.1126/science.aat1066 1991:Fertility and Sterility 815:Encyclopedia Britannica 516:that contain their own 271:Stemagen (La Jolla, CA) 248:immune system rejection 88:Zhong Zhong and Hua Hua 2995:10.1186/1477-7827-1-98 2955:. 113. (Pt 1): 11–20. 2668:Sheryl Gay Stolberg, " 1901:10.1096/fj.201900578RR 1175:10.1089/cell.2017.0051 1163:Cellular Reprogramming 924:10.1002/anie.201306722 729:Cloning and Stem Cells 680:In vitro fertilisation 552: 330:pluripotent stem cells 160: 90:) from foetal nuclei. 24: 3969:1996 in biotechnology 3251:Roundup ready soybean 2647:"Fusion Biopolitics". 2502:10.1387/ijdb.180324mc 1215:10.1089/scd.2013.0402 742:10.1089/clo.2008.0041 700:Stem cell controversy 542: 535:Stem cell controversy 533:Further information: 158: 99:regenerative medicine 33:developmental biology 22: 3790:Reverse transfection 3565:Genetic transduction 2961:10.1242/jcs.113.1.11 2600:10.3390/ijms23041969 1455:Tachibana M (2013). 1255:10.1210/er.2008-0031 461:improve this article 376:crab-eating macaques 337:Reproductive cloning 279:embryonic stem cells 257:, scientists at the 223:Embryonic stem cells 107:embryonic stem cells 84:crab-eating macaques 3780:Genetics in fiction 3712:Genetic enhancement 3514:Hepatitis B vaccine 3144:Genetic engineering 3035:2003PNAS..10011819G 2924:10.1038/nature01079 2916:2002Natur.419..583W 2870:Council of Europe, 2342:2017PLoSO..1277800W 2188:Scientific American 2148:2018Natur.553..387C 1948:1996Natur.380...64C 1853:10.1038/nature08267 1845:2009Natur.461...86Z 1708:10.1038/nature09342 1700:2010Natur.467..285K 1652:(5909): 1766–1767. 1611:10.1038/nature13287 1603:2014Natur.510..533Y 1362:2013Natur.498..159P 1026:1997Natur.385..810W 918:(52): 13890–13899. 885:10.1242/dev.8.4.437 653:Additional Protocol 638:Council of Europe's 239:Parkinson's disease 196:phytohaemagglutinin 95:Therapeutic cloning 3964:Stem cell research 3954:Induced stem cells 3890:Stem cell research 3509:Ice-minus bacteria 2748:2007-07-06 at the 2712:2006-09-30 at the 2696:, (March 19, 2002) 2692:2010-11-22 at the 2676:(January 24, 2002) 2652:2009-06-16 at the 2496:(3–4–5): 123–130. 2244:The New York Times 1537:Charles C. Mann, " 1503:Elizabeth Weise, " 1413:(8): REVIEWS1023. 1010:Campbell, K. H. S. 705:Stem cell research 685:Induced stem cells 553: 213:stem cell research 207:Stem cell research 161: 126:Early 20th-Century 25: 3931: 3930: 3895:Synthetic biology 3785:Human enhancement 3775:Genetic pollution 3751: 3750: 3619: 3618: 3527: 3526: 3490: 3489: 3384: 3383: 2857:United Nations, " 2142:(7689): 387–388. 1751:(19): 3553–3575. 1304:(11–12): 743–50. 1243:Endocrine Reviews 1208:(Suppl 1): 25–8. 1138:978-0-12-374775-4 1020:(6619): 810–813. 781:(4): 881–887.e7. 518:mitochondrial DNA 493: 492: 485: 226:destroyed cells. 183:micromanipulators 3981: 3668:Dow AgroSciences 3628: 3627: 3538: 3537: 3395: 3394: 3170: 3169: 3161: 3160: 3137: 3130: 3123: 3114: 3113: 3066: 3056: 3046: 3017: 3007: 2997: 2972: 2943: 2901: 2883: 2868: 2862: 2855: 2849: 2848: 2812: 2806: 2805: 2779: 2759: 2753: 2740: 2734: 2723: 2717: 2703: 2697: 2683: 2677: 2666: 2660: 2640: 2631: 2630: 2620: 2602: 2578: 2569: 2568: 2528: 2522: 2521: 2481: 2475: 2474: 2472: 2470: 2455: 2449: 2442: 2436: 2435: 2387: 2374: 2373: 2363: 2353: 2321: 2315: 2314: 2312: 2310: 2289: 2271: 2262: 2256: 2255: 2253: 2251: 2234: 2228: 2227: 2225: 2223: 2206: 2200: 2199: 2197: 2195: 2178: 2172: 2169: 2159: 2124: 2122: 2120: 2093: 2087: 2086: 2050: 2044: 2043: 2023: 2017: 2016: 2006: 1982: 1976: 1975: 1956:10.1038/380064a0 1931: 1922: 1921: 1903: 1894:(1): 1637–1651. 1879: 1873: 1872: 1828: 1822: 1821: 1804:(5): 1202–1211. 1793: 1787: 1786: 1768: 1736: 1730: 1729: 1719: 1694:(7313): 285–90. 1678: 1672: 1671: 1661: 1637: 1631: 1630: 1585: 1579: 1569: 1563: 1552: 1546: 1535: 1529: 1518: 1512: 1501: 1495: 1494: 1484: 1452: 1443: 1442: 1432: 1422: 1398: 1392: 1391: 1373: 1356:(7453): 159–61. 1341: 1332: 1331: 1313: 1295: 1286: 1277: 1276: 1266: 1234: 1228: 1227: 1217: 1193: 1187: 1186: 1157: 1151: 1150: 1116: 1110: 1109: 1099: 1089: 1065: 1054: 1053: 1034:10.1038/385810a0 1002: 996: 995: 985: 953: 944: 943: 903: 897: 896: 860: 851: 850: 848: 846: 832: 826: 825: 823: 821: 807: 801: 800: 790: 764: 755: 754: 744: 720: 675:Handmade cloning 488: 481: 477: 474: 468: 445: 437: 388:) was reported. 291:Roslin Institute 173:(an early stage 3989: 3988: 3984: 3983: 3982: 3980: 3979: 3978: 3934: 3933: 3932: 3927: 3899: 3868: 3822: 3804: 3757: 3747: 3721: 3717:Genetic testing 3699: 3689: 3615: 3601:Recombinant DNA 3589: 3560:Electroporation 3523: 3519:Oncolytic virus 3498: 3486: 3443: 3429:Herman the Bull 3380: 3314: 3296: 3278: 3260: 3237: 3219: 3201: 3155: 3153: 3146: 3141: 3073: 2910:(6907): 583–6. 2899: 2891: 2889:Further reading 2886: 2869: 2865: 2856: 2852: 2813: 2809: 2777:10.1.1.475.1709 2760: 2756: 2750:Wayback Machine 2741: 2737: 2725:Andy Coghlan, " 2724: 2720: 2714:Wayback Machine 2704: 2700: 2694:Wayback Machine 2684: 2680: 2667: 2663: 2654:Wayback Machine 2641: 2634: 2579: 2572: 2529: 2525: 2482: 2478: 2468: 2466: 2456: 2452: 2443: 2439: 2388: 2377: 2336:(5): e0177800. 2322: 2318: 2308: 2306: 2287:10.1.1.455.5842 2269: 2263: 2259: 2249: 2247: 2235: 2231: 2221: 2219: 2207: 2203: 2193: 2191: 2179: 2175: 2118: 2116: 2094: 2090: 2067:10.1038/nrg1205 2055:Nat. 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Genet 2051: 2047: 2024: 2020: 1983: 1979: 1932: 1925: 1880: 1876: 1839:(7260): 86–90. 1829: 1825: 1794: 1790: 1737: 1733: 1679: 1675: 1638: 1634: 1597:(7506): 533–6. 1586: 1582: 1576:Washington Post 1570: 1566: 1553: 1549: 1536: 1532: 1519: 1515: 1502: 1498: 1453: 1446: 1399: 1395: 1371:10.1038/498159a 1342: 1335: 1293: 1289:Semb H (2005). 1287: 1280: 1235: 1231: 1194: 1190: 1158: 1154: 1139: 1117: 1113: 1066: 1057: 1003: 999: 954: 947: 904: 900: 861: 854: 844: 842: 834: 833: 829: 819: 817: 809: 808: 804: 765: 758: 721: 717: 713: 661: 604: 549:inner cell mass 537: 531: 489: 478: 472: 469: 458: 446: 435: 427:dromedary camel 394: 372:Dolly the sheep 353:Dolly the sheep 345: 339: 209: 204: 153: 132:Sir John Gurdon 128: 119: 76:Dolly the sheep 17: 12: 11: 5: 3987: 3977: 3976: 3971: 3966: 3961: 3959:Life extension 3956: 3951: 3946: 3929: 3928: 3926: 3925: 3920: 3915: 3910: 3904: 3901: 3900: 3898: 3897: 3892: 3887: 3882: 3876: 3874: 3873:Similar fields 3870: 3869: 3867: 3866: 3861: 3856: 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Index


genetics
developmental biology
laboratory
embryo
body cell
egg cell
denucleated
oocyte
nucleus
cloning
Dolly the sheep
Shanghai
crab-eating macaques
Zhong Zhong and Hua Hua
regenerative medicine
issues
embryonic stem cells
homologous
Sir John Gurdon
Xenopus laevis
Rana pipiens

reprogrammed
blastocyst
embryo
micromanipulators
zona pellucida
phytohaemagglutinin
stem cell research

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