Embryonic stem cell

600:, PhD. A previous experiment had shown an improvement in locomotor recovery in spinal cord-injured rats after a 7-day delayed transplantation of human ESCs that had been pushed into an oligodendrocytic lineage. The phase I clinical study was designed to enroll about eight to ten paraplegics who have had their injuries no longer than two weeks before the trial begins, since the cells must be injected before scar tissue is able to form. The researchers emphasized that the injections were not expected to fully cure the patients and restore all mobility. Based on the results of the rodent trials, researchers speculated that restoration of myelin sheathes and an increase in mobility might occur. This first trial was primarily designed to test the safety of these procedures and if everything went well, it was hoped that it would lead to future studies that involve people with more severe disabilities. The trial was put on hold in August 2009 due to FDA concerns regarding a small number of microscopic cysts found in several treated rat models but the hold was lifted on July 30, 2010. 647:

presented the results from phase 1 clinical trial testing of a low dose of AST-OPC1 in patients with neurologically complete thoracic spinal cord injury. The results showed that AST-OPC1 was successfully delivered to the injured spinal cord site. Patients followed 2–3 years after AST-OPC1 administration showed no evidence of serious adverse events associated with the cells in detailed follow-up assessments including frequent neurological exams and MRIs. Immune monitoring of subjects through one year post-transplantation showed no evidence of antibody-based or cellular immune responses to AST-OPC1. In four of the five subjects, serial MRI scans performed throughout the 2–3 year follow-up period indicate that reduced spinal cord cavitation may have occurred and that AST-OPC1 may have had some positive effects in reducing spinal cord tissue deterioration. There was no unexpected neurological degeneration or improvement in the five subjects in the trial as evaluated by the International Standards for Neurological Classification of Spinal Cord Injury (ISNCSCI) exam.

392:, the use of stem cells are known to be of importance. In order to successfully engineer a tissue, the cells used must be able to perform specific biological functions such as secretion of cytokines, signaling molecules, interacting with neighboring cells, and producing an extracellular matrix in the correct organization. Stem cells demonstrates these specific biological functions along with being able to self-renew and differentiate into one or more types of specialized cells. Embryonic stem cells is one of the sources that are being considered for the use of tissue engineering. The use of human embryonic stem cells have opened many new possibilities for tissue engineering, however, there are many hurdles that must be made before human embryonic stem cell can even be utilized. It is theorized that if embryonic stem cells can be altered to not evoke the immune response when implanted into the patient then this would be a revolutionary step in tissue engineering. Embryonic stem cells are not limited to tissue engineering. 907:, the process in which antibodies are bound to the trophectoderm and removed by another solution, and mechanical dissection are performed to achieve separation. The resulting inner cell mass cells are plated onto cells that will supply support. The inner cell mass cells attach and expand further to form a human embryonic cell line, which are undifferentiated. These cells are fed daily and are enzymatically or mechanically separated every four to seven days. For differentiation to occur, the human embryonic stem cell line is removed from the supporting cells to form embryoid bodies, is co-cultured with a serum containing necessary signals, or is grafted in a three-dimensional scaffold to result. 558:

in the G1 phase of the cell cycle (i.e. after metaphase/cell division but prior the next round of replication) have only one copy of each chromosome (i.e. sister chromosomes aren't present). Mouse ES cells lack a G1 checkpoint and do not undergo cell cycle arrest upon acquiring DNA damage. Rather they undergo programmed cell death (apoptosis) in response to DNA damage. Apoptosis can be used as a fail-safe strategy to remove cells with un-repaired DNA damages in order to avoid mutation and progression to cancer. Consistent with this strategy, mouse ES stem cells have a mutation frequency about 100-fold lower than that of isogenic mouse somatic cells.

813:. This experiment instituted the proposition that specialized adult cells obtain the genetic makeup to perform a specific task; which established a basis for further research within a variety of cloning techniques. The Dolly experiment was performed by obtaining the mammalian udder cells from a sheep (Dolly) and differentiating these cells until division was concluded. An egg cell was then procured from a different sheep host and the nucleus was removed. An udder cell was placed next to the egg cell and connected by electricity causing this cell to share DNA. This egg cell differentiated into an 146: 4525: 824:(James A. Thomson, Joseph Itskovitz-Eldor, Sander S. Shapiro, Michelle A. Waknitz, Jennifer J. Swiergiel, Vivienne S. Marshall, and Jeffrey M. Jones) publish a paper titled "Embryonic Stem Cell Lines Derived From Human Blastocysts". The researchers behind this study not only created the first embryonic stem cells, but recognized their pluripotency, as well as their capacity for self-renewal. The abstract of the paper notes the significance of the discovery with regards to the fields of developmental biology and drug discovery. 747: 537:. The techniques are now used by many pregnant women and prospective parents, especially couples who have a history of genetic abnormalities or where the woman is over the age of 35 (when the risk of genetically related disorders is higher). In addition, by allowing parents to select an embryo without genetic disorders, they have the potential of saving the lives of siblings that already had similar disorders and diseases using cells from the disease free offspring. 1079:. This approach has also sometimes been referred to as "therapeutic cloning" because SCNT bears similarity to other kinds of cloning in that nuclei are transferred from a somatic cell into an enucleated zygote. However, in this case SCNT was used to produce embryonic stem cell lines in a lab, not living organisms via a pregnancy. The "therapeutic" part of the name is included because of the hope that SCNT produced embryonic stem cells could have clinical utility. 5645: 42: 5673: 31: 799: 5685: 138: 705:

the 5-day-old mass of cells is too young to achieve personhood or that the embryo, if donated from an IVF clinic (where labs typically acquire embryos), would otherwise go to medical waste anyway. Opponents of ESC research claim that an embryo is a human life, therefore destroying it is murder and the embryo must be protected under the same ethical view as a more developed human being.

381: 1135:) to differentiated cells. Utilizing the four genes previously listed, the differentiated cells are "reprogrammed" into pluripotent stem cells, allowing for the generation of pluripotent/embryonic stem cells without the embryo. The morphology and growth factors of these lab induced pluripotent cells, are equivalent to embryonic stem cells, leading these cells to be known as 1037:(BMPs) that are necessary to prevent ES cells from differentiating. These factors are extremely important for the efficiency of deriving ES cells. Furthermore, it has been demonstrated that different mouse strains have different efficiencies for isolating ES cells. Current uses for mouse ES cells include the generation of 682:

eliminating c-Myc expression are unlikely to preserve the cells' "stemness". However, N-myc and L-myc have been identified to induce iPS cells instead of c-myc with similar efficiency. Later protocols to induce pluripotency bypass these problems completely by using non-integrating RNA viral vectors such as

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ESCs grown in serum-free 2i conditions do express hypo-phosphorylated active Retinoblastoma proteins and have an elongated G1 phase. Despite this difference in the cell cycle when compared to ESCs grown in media containing serum these cells have similar pluripotent characteristics. Pluripotency factors

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in Worcester, MA) stating that his team had found a way to extract embryonic stem cells without destroying the actual embryo. This technical achievement would potentially enable scientists to work with new lines of embryonic stem cells derived using public funding in the US, where federal funding was

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to repair DSBs. This type of repair depends on the interaction of the two sister chromosomes formed during S phase and present together during the G2 phase of the cell cycle. HRR can accurately repair DSBs in one sister chromosome by using intact information from the other sister chromosome. Cells

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Several new studies have started to address the concept of modeling genetic disorders with embryonic stem cells. Either by genetically manipulating the cells, or more recently, by deriving diseased cell lines identified by prenatal genetic diagnosis (PGD), modeling genetic disorders is something that

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are hyperphosphorylated and inactivated in ESCs, leading to continual expression of proliferation genes. These changes result in accelerated cycles of cell division. Although high expression levels of pro-proliferative proteins and a shortened G1 phase have been linked to maintenance of pluripotency,

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Under defined conditions, embryonic stem cells are capable of self-renewing indefinitely in an undifferentiated state. Self-renewal conditions must prevent the cells from clumping and maintain an environment that supports an unspecialized state. Typically this is done in the lab with media containing

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Embryonic stem cells (ESCs), derived from the blastocyst stage of early mammalian embryos, are distinguished by their ability to differentiate into any embryonic cell type and by their ability to self-renew. It is these traits that makes them valuable in the scientific and medical fields. ESCs have a

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published a study on January 24, 2005, which stated that the human embryonic stem cells available for federally funded research are contaminated with non-human molecules from the culture medium used to grow the cells. It is a common technique to use mouse cells and other animal cells to maintain the

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Due to the nature of embryonic stem cell research, there are a lot of controversial opinions on the topic. Since harvesting embryonic stem cells usually necessitates destroying the embryo from which those cells are obtained, the moral status of the embryo comes into question. Some people claim that

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ES cells use a different strategy to deal with DSBs. Because ES cells give rise to all of the cell types of an organism including the cells of the germ line, mutations arising in ES cells due to faulty DNA repair are a more serious problem than in differentiated somatic cells. Consequently, robust

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cells. However, the derivation of such cell types from ESCs is not without obstacles, therefore research has focused on overcoming these barriers. For example, studies are underway to differentiate ESCs into tissue specific cardiomyocytes and to eradicate their immature properties that distinguish

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on March 8, 2005, detailed information about a new stem cell line that was derived from human embryos under completely cell- and serum-free conditions. After more than 6 months of undifferentiated proliferation, these cells demonstrated the potential to form derivatives of all three embryonic germ

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This may enable the generation of patient specific ES cell lines that could potentially be used for cell replacement therapies. In addition, this will allow the generation of ES cell lines from patients with a variety of genetic diseases and will provide invaluable models to study those diseases.

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generates multiple embryos. The surplus of embryos is not clinically used or is unsuitable for implantation into the patient, and therefore may be donated by the donor with consent. Human embryonic stem cells can be derived from these donated embryos or additionally they can also be extracted from

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ESC-derived hepatocytes are also useful models that could be used in the preclinical stages of drug discovery. However, the development of hepatocytes from ESCs has proven to be challenging and this hinders the ability to test drug metabolism. Therefore, research has focused on establishing fully

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Zikuan Leng 1 2, Dongming Sun 2, Zihao Huang 3, Iman Tadmori 2, Ning Chiang 2, Nikhit Kethidi 2, Ahmed Sabra 2, Yoshihiro Kushida 4, Yu-Show Fu 3, Mari Dezawa 4, Xijing He 1, Wise Young 2Quantitative Analysis of SSEA3+ Cells from Human Umbilical Cord after Magnetic SortingCell Transplant . 2019

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found in adults. They were discovered in 2010 by Mari Dezawa and her research group. Muse cells reside in the connective tissue of nearly every organ including the umbilical cord, bone marrow and peripheral blood. They are collectable from commercially obtainable mesenchymal cells such as human

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The Strategic Partnership III grant from CIRM will provide funding to Asterias to support the next clinical trial of AST-OPC1 in subjects with spinal cord injury, and for Asterias' product development efforts to refine and scale manufacturing methods to support later-stage trials and eventually

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AST-OPC1 is a population of cells derived from human embryonic stem cells (hESCs) that contains oligodendrocyte progenitor cells (OPCs). OPCs and their mature derivatives called oligodendrocytes provide critical functional support for nerve cells in the spinal cord and brain. Asterias recently

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Ogura, Fumitaka; Wakao, Shohei; Kuroda, Yasumasa; Tsuchiyama, Kenichiro; Bagheri, Mozhdeh; Heneidi, Saleh; Chazenbalk, Gregorio; Aiba, Setsuya; Dezawa, Mari (2014). "Human Adipose Tissue Possesses a Unique Population of Pluripotent Stem Cells with Nontumorigenic and Low Telomerase Activities:

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BioTime company Asterias Biotherapeutics (NYSE MKT: AST) was granted a $ 14.3 million Strategic Partnership Award by the California Institute for Regenerative Medicine (CIRM) to re-initiate the world's first embryonic stem cell-based human clinical trial, for spinal cord injury. Supported by

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because they are not genetically modified with genes such as c-Myc that are linked to cancer. Nonetheless, ESCs express very high levels of the iPS inducing genes and these genes including Myc are essential for ESC self-renewal and pluripotency, and potential strategies to improve safety by

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The main strategy to enhance the safety of ESCs for potential clinical use is to differentiate the ESCs into specific cell types (e.g. neurons, muscle, liver cells) that have reduced or eliminated ability to cause tumors. Following differentiation, the cells are subjected to sorting by

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allows federal funding to support research on roughly 60—at this time, already existing—lines of embryonic stem cells. Seeing as the limited lines that Bush allowed research on had already been established, this law supported embryonic stem cell research without raising any

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Hori, Emiko; Hayakawa, Yumiko; Hayashi, Tomohide; Hori, Satoshi; Okamoto, Soushi; Shibata, Takashi; Kubo, Michiya; Horie, Yukio; Sasahara, Masakiyo; Kuroda, Satoshi (2016). "Mobilization of Pluripotent Multilineage-Differentiating Stress-Enduring Cells in Ischemic Stroke".

347:, etc. In addition to their potential in regenerative medicine, embryonic stem cells provide a possible alternative source of tissue/organs which serves as a possible solution to the donor shortage dilemma. There are some ethical controversies surrounding this though (see 664:

The major concern with the possible transplantation of ESCs into patients as therapies is their ability to form tumors including teratomas. Safety issues prompted the FDA to place a hold on the first ESC clinical trial, however no tumors were observed.

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In November 2011 Geron announced it was halting the trial and dropping out of stem cell research for financial reasons, but would continue to monitor existing patients, and was attempting to find a partner that could continue their research. In 2013

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Hanna, J.; Wernig, M.; Markoulaki, S.; Sun, C.-W.; Meissner, A.; Cassady, J. P.; Beard, C.; Brambrink, T.; Wu, L.-C.; Townes, T. M.; Jaenisch, R. (2007). "Treatment of Sickle Cell Anemia Mouse Model with iPS Cells Generated from Autologous Skin".

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commercialization. CIRM funding will be conditional on FDA approval for the trial, completion of a definitive agreement between Asterias and CIRM, and Asterias' continued progress toward the achievement of certain pre-defined project milestones.

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On January 16, 2008, a California-based company, Stemagen, announced that they had created the first mature cloned human embryos from single skin cells taken from adults. These embryos can be harvested for patient matching embryonic stem cells.

1045:. For human treatment, there is a need for patient specific pluripotent cells. Generation of human ES cells is more difficult and faces ethical issues. So, in addition to human ES cell research, many groups are focused on the generation of 4564:

Wernig, Marius; Meissner, Alexander; Foreman, Ruth; Brambrink, Tobias; Ku, Manching; Hochedlinger, Konrad; Bernstein, Bradley E.; Jaenisch, Rudolf (2007-07-19). "In vitro reprogramming of fibroblasts into a pluripotent ES-cell-like state".

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The award provides funding for Asterias to reinitiate clinical development of AST-OPC1 in subjects with spinal cord injury and to expand clinical testing of escalating doses in the target population intended for future pivotal trials.

549:, an error prone DNA repair process, as the primary pathway for repairing double-strand breaks (DSBs) during all cell cycle stages. Because of its error-prone nature, NHEJ tends to produce mutations in a cell's clonal descendants. 981:

derived and cultured her ES cells differently. She removed the embryos from the donor mother at approximately 76 hours after copulation and cultured them overnight in a medium containing serum. The following day, she removed the

722:. These cells were isolated from the teratocarcinoma replicated and grew in cell culture as a stem cell and are now known as embryonal carcinoma (EC) cells. Although similarities in morphology and differentiating potential ( 5007:

Kuroda, Y.; Kitada, M.; Wakao, S.; Nishikawa, K.; Tanimura, Y.; Makinoshima, H.; Goda, M.; Akashi, H.; Inutsuka, A.; Niwa, A.; Shigemoto, T.; Nabeshima, Y.; Nakahata, T.; Nabeshima, Y.-i.; Fujiyoshi, Y.; Dezawa, M. (2010).

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d'Amour, KA; Bang, AG; Eliazer, S; Kelly, OG; Agulnick, AD; Smart, NG; Moorman, MA; Kroon, E; Carpenter, MK; Baetge, EE (2006). "Production of pancreatic hormone-expressing endocrine cells from human embryonic stem cells".

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Tachibana, Masahito; Amato, Paula; Sparman, Michelle; Gutierrez, Nuria Marti; Tippner-Hedges, Rebecca; Ma, Hong; Kang, Eunju; Fulati, Alimujiang; Lee, Hyo-Sang; Sritanaudomchai, Hathaitip; Masterson, Keith (2013-06-06).

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Williams RL, Hilton DJ, Pease S, Willson TA, Stewart CL, Gearing DP, Wagner EF, Metcalf D, Nicola NA, Gough NM (1988). "Myeloid leukaemia inhibitory factor maintains the developmental potential of embryonic stem cells".

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Zhang, X.; Neganova, I.; Przyborski, S.; Yang, C.; Cooke, M.; Atkinson, S. P.; Anyfantis, G.; Fenyk, S.; Keith, W. N.; Hoare, S. F.; Hughes, O.; Strachan, T.; Stojkovic, M.; Hinds, P. W.; Armstrong, L.; Lako, M. (2009).

866:. The announcement was met with excitement from the scientific community, but also with wariness from stem cell opposers. The treatment cells were, however, derived from the cell lines approved under George W. Bush's 368: 5240:

Kuroda, Yasumasa; Wakao, Shohei; Kitada, Masaaki; Murakami, Toru; Nojima, Makoto; Dezawa, Mari (2013). "Isolation, culture and evaluation of multilineage-differentiating stress-enduring (Muse) cells".

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Söderdahl, T; Küppers-Munther, B; Heins, N; Edsbagge, J; Björquist, P; Cotgreave, I; Jernström, B (2007). "Glutathione transferases in hepatocyte-like cells derived from human embryonic stem cells".

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Because ethical concerns regarding embryonic stem cells typically are about their derivation from terminated embryos, it is believed that reprogramming to these iPS cells may be less controversial.

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Besides becoming an important alternative to organ transplants, ESCs are also being used in the field of toxicology, and as cellular screens to uncover new chemical entities that can be developed as

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models to test drug responses and predict toxicity profiles. ESC derived cardiomyocytes have been shown to respond to pharmacological stimuli and hence can be used to assess cardiotoxicity such as

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Chung, Young Gie; Eum, Jin Hee; Lee, Jeoung Eun; Shim, Sung Han; Sepilian, Vicken; Hong, Seung Wook; Lee, Yumie; Treff, Nathan R.; Choi, Young Ho; Kimbrel, Erin A.; Dittman, Ralph E. (2014-06-05).

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published first in July, revealing a new technique for culturing the mouse embryos in the uterus to allow for an increase in cell number, allowing for the derivation of ES cell from these embryos.

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Research has focused on differentiating ESCs into a variety of cell types for eventual use as cell replacement therapies. Some of the cell types that have or are currently being developed include

1247:, bone marrow-mesenchymal stem cells and adipose-derived stem cells. Muse cells are able to generate cells representative of all three germ layers from a single cell both spontaneously and under 553:

mechanisms are needed in ES cells to repair DNA damages accurately, and if repair fails, to remove those cells with un-repaired DNA damages. Thus, mouse ES cells predominantly use high fidelity

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Researchers are currently focusing heavily on the therapeutic potential of embryonic stem cells, with clinical use being the goal for many laboratories. Potential uses include the treatment of

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Bernstein C, Bernstein H, Payne CM, Garewal H (June 2002). "DNA repair/pro-apoptotic dual-role proteins in five major DNA repair pathways: fail-safe protection against carcinogenesis".

903:. The inner cell mass (cells of interest), from the blastocyst stage of the embryo, is separated from the trophectoderm, the cells that would differentiate into extra-embryonic tissue. 372:, "Human embryonic stem cells have the potential to differentiate into various cell types, and, thus, may be useful as a source of cells for transplantation or tissue engineering." 3781: 3242: 444:

Studies involving ESCs are underway to provide an alternative treatment for diabetes. For example ESCs have been differentiated into insulin-producing cells, and researchers at

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Wakao, S.; Kitada, M.; Kuroda, Y.; Shigemoto, T.; Matsuse, D.; Akashi, H.; Tanimura, Y.; Tsuchiyama, K.; Kikuchi, T.; Goda, M.; Nakahata, T.; Fujiyoshi, Y.; Dezawa, M. (2011).

939:, changing the hormone environment, which causes the embryos to remain free in the uterus. After 4–6 days of this intrauterine culture, the embryos are harvested and grown in 461:

describes a translational process of generating human embryonic stem cell-derived cardiac progenitor cells to be used in clinical trials of patients with severe heart failure.

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Smith AG, Heath JK, Donaldson DD, Wong GG, Moreau J, Stahl M, Rogers D (1988). "Inhibition of pluripotential embryonic stem cell differentiation by purified polypeptides".

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Takahashi K, Tanabe K, Ohnuki M, Narita M, Ichisaka T, Tomoda K, Yamanaka S (2007). "Induction of pluripotent stem cells from adult human fibroblasts by defined factors".

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Thompson, James A.; Itskovitz-Eldor, Joseph; Shapiro, Sander S.; Waknitz, Michelle A.; Swiergiel, Jennifer J.; Marshall, Vivienne S.; Jones, Jeffrey M. (6 November 1998).

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Varlakhanova, Natalia V.; Cotterman, Rebecca F.; Devries, Wilhelmine N.; Morgan, Judy; Donahue, Leah Rae; Murray, Stephen; Knowles, Barbara B.; Knoepfler, Paul S. (2010).

881:(NIH) to provide funding for hESC research. The document also states that the NIH must provide revised federal funding guidelines within 120 days of the order's signing. 545:

Differentiated somatic cells and ES cells use different strategies for dealing with DNA damage. For instance, human foreskin fibroblasts, one type of somatic cell, use

1730:"Developmental activation of the Rb-E2F pathway and establishment of cell cycle-regulated cyclin-dependent kinase activity during embryonic stem cell differentiation" 323:

and tissue replacement after injury or disease. Pluripotent stem cells have shown promise in treating a number of varying conditions, including but not limited to:

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Yamanaka, Shinya; Takahashi, Kazutoshi (25 Aug 2006). "Induction of Pluripotent Stem Cells From Mouse Embryonic and Adult Fibroblast Cultures by Defined Factors".

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in a medium containing serum and conditioned by ES cells. After approximately one week, colonies of cells grew out. These cells grew in culture and demonstrated

4769: 3168: 584:(FDA), marking it the world's first human ESC human trial. The study leading to this scientific advancement was conducted by Hans Keirstead and colleagues at the 110:, including whether or not embryos at the pre-implantation stage have the same moral considerations as embryos in the post-implantation stage of development. 3718: 2750: 1283: 434:

Researchers have differentiated ESCs into dopamine-producing cells with the hope that these neurons could be used in the treatment of Parkinson's disease.

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reported a technique that delays embryo implantation, allowing the inner cell mass to increase. This process includes removing the donor mother's

877:, removing the restrictions put in place on federal funding for human stem cells by the previous presidential administration. This would allow the 4346: 3211: 3141: 1950:

Ying, Qi-Long; Wray, Jason; Nichols, Jennifer; Batlle-Morera, Laura; Doble, Bradley; Woodgett, James; Cohen, Philip; Smith, Austin (2008-05-22).

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However, as a first indication that the iPS cell technology can in rapid succession lead to new cures, it was used by a research team headed by

5625: 5321:"Awakened by Cellular Stress: Isolation and Characterization of a Novel Population of Pluripotent Stem Cells Derived from Human Adipose Tissue" 4478:

Takahashi, K; Yamanaka, S (2006). "Induction of pluripotent stem cells from mouse embryonic and adult fibroblast cultures by defined factors".

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Martin Evans revealed a new technique for culturing the mouse embryos in the uterus to allow for the derivation of ES cells from these embryos.

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Pluripotent: Embryonic stem cells are able to develop into any type of cell, excepting those of the placenta. Only embryonic stem cells of the

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induction. Expression of pluripotency genes and triploblastic differentiation are self-renewable over generations. Muse cells do not undergo

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that accumulated during the development of the teratocarcinoma. These genetic aberrations further emphasized the need to be able to culture

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are a huge discovery, as they are seemingly identical to embryonic stem cells and could be used without sparking the same moral controversy.

3101:"Human embryonic stem cell-derived oligodendrocyte progenitor cell transplants remyelinate and restore locomotion after spinal cord injury" 581: 17: 1072:

at the time limited to research using embryonic stem cell lines derived prior to August 2001. In March, 2009, the limitation was lifted.

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in the growth medium was found to compromise the potential uses of the embryonic stem cells in humans, according to scientists at the

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Takahashi, Kazutoshi; Tanabe, Koji; Ohnuki, Mari; Narita, Megumi; Ichisaka, Tomoko; Tomoda, Kiichiro; Yamanaka, Shinya (2007-11-30).

1377: 5630: 4851: 1588:"BMP Induction of Id Proteins Suppresses Differentiation and Sustains Embryonic Stem Cell Self-Renewal in Collaboration with STAT3" 351:

section below). Aside from these uses, ESCs can also be used for research on early human development, certain genetic disease, and

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Klimanskaya I, Chung Y, Meisner L, Johnson J, West MD, Lanza R (2005). "Human embryonic stem cells derived without feeder cells".

2824:"Mouse embryonic stem cells, but not somatic cells, predominantly use homologous recombination to repair double-strand DNA breaks" 3077: 772: 1139:(iPS cells). This observation was observed in mouse pluripotent stem cells, originally, but now can be performed in human adult 5319:

Heneidi, Saleh; Simerman, Ariel A.; Keller, Erica; Singh, Prapti; Li, Xinmin; Dumesic, Daniel A.; Chazenbalk, Gregorio (2013).

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culture until the inner cell mass forms “egg cylinder-like structures,” which are dissociated into single cells, and plated on

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Klimanskaya I, Chung Y, Becker S, Lu SJ, Lanza R (2006). "Human embryonic stem cell lines derived from single blastomeres".

3653:"Isolation of a pluripotent cell line from early mouse embryos cultured in medium conditioned by teratocarcinoma stem cells" 1537:

Thomson, J. A.; Itskovitz-Eldor, J; Shapiro, S. S.; Waknitz, M. A.; Swiergiel, J. J.; Marshall, V. S.; Jones, J. M. (1998).

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are created by growing up a single cell. Evans and Kaufman showed that the cells grown out from these cultures could form

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publish their research that details their isolation and genetic modifications of embryonic stem cells, creating the first "

5165:"Muse Cells Provide the Pluripotency of Mesenchymal Stem Cells: Direct Contribution of Muse Cells to Tissue Regeneration" 3764:

Klotzko, Arlene Judith; Klotzko, Visiting Scholar Royal Free and University College Medical School Arlene Judith (2006).

2932:"ES cells do not activate p53-dependent stress responses and undergo p53-independent apoptosis in response to DNA damage" 2629:

Menasché, Phillip; Vanneaux, Valérie; Fabreguettes, Jean-Roch; Bel, Alain; Tosca, Lucie; Garcia, Sylvie (21 March 2015).

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Martin MJ, Muotri A, Gage F, Varki A (2005). "Human embryonic stem cells express an immunogenic nonhuman sialic acid".

4873: 1467: 1221:. These properties were also successfully maintained (for more than 30 passages) with the established stem cell lines. 775:, published her paper in December and coined the term "Embryonic Stem Cell". She showed that embryos could be cultured 125:, and cellular/DNA repair. However, adverse effects in the research and clinical processes such as tumors and unwanted 843:

and Kazutoshi Takashi publish a paper describing the induction of pluripotent stem cells from cultures of adult mouse

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Mountford, JC (2008). "Human embryonic stem cells: origins, characteristics and potential for regenerative therapy".

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that in subsequently differentiate into the desired cell types. Pluripotency distinguishes embryonic stem cells from

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allows the cells to quickly grow in number, but not size, which is important for early embryo development. In ESCs,

2873:"Restoration of an absent G1 arrest and protection from apoptosis in embryonic stem cells after ionizing radiation" 821: 585: 328: 855: 3710: 2747: 2067:"A role for NANOG in G1 to S transition in human embryonic stem cells through direct binding of CDK6 and CDC25A" 639:

California public funds, CIRM is the largest funder of stem cell-related research and development in the world.

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has been accomplished with stem cells. This approach may very well prove valuable at studying disorders such as

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A blog focusing specifically on ES cells and iPS cells including research, biotech, and patient-oriented issues

1046: 5489: 4538: 2631:"Towards a clinical use of human embryonic stem cell derived-cardiac progenitors: a translational experience" 2609: 1401:"Morality and human embryo research. Introduction to the Talking Point on morality and human embryo research" 1136: 1088: 1076: 900: 878: 848: 794:". In creating knockout mice, this publication provided scientists with an entirely new way to study disease. 755:

1981: Embryonic stem cells (ES cells) were independently first derived from a mouse embryos by two groups.

4626:"Induction of Pluripotent Stem Cells from Mouse Embryonic and Adult Fibroblast Cultures by Defined Factors" 3299: 2385:

Siu, CW; Moore, JC; Li, RA (2007). "Human embryonic stem cell-derived cardiomyocytes for heart therapies".

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could convert adult cells into pluripotent stem cells. He was awarded the 2012 Nobel Prize along with Sir

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Jensen, J; Hyllner, J; Björquist, P (2009). "Human embryonic stem cell technologies and drug discovery".

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Ledermann B, Bürki K (1991). "Establishment of a germ-line competent C57BL/6 embryonic stem cell line".

3201: 5663: 4760:"Scientists Cure Mice Of Sickle Cell Using Stem Cell Technique: New Approach Is From Skin, Not Embryos" 3972:"Executive Order 13505—Removing Barriers To Responsible Scientific Research Involving Human Stem Cells" 2772:"DNA repair by nonhomologous end joining and homologous recombination during cell cycle in human cells" 1030: 546: 283:

that promote cell cycle progression are overactive, in part due to downregulation of their inhibitors.

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Thomson; Itskovitz-Eldor, J; Shapiro, SS; Waknitz, MA; Swiergiel, JJ; Marshall, VS; Jones, JM (1998).

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Thomson JA, Itskovitz-Eldor J, Shapiro SS, Waknitz MA, Swiergiel JJ, Marshall VS, Jones JM (1998).

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Ter Huurne, Menno; Chappell, James; Dalton, Stephen; Stunnenberg, Hendrik G. (5 October 2017).

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and the embryo was inserted into a third sheep which gave birth to the clone version of Dolly.

5725: 5610: 5565: 5555: 5539: 1294: 1239: 1106: 833: 699: 616: 530: 320: 107: 3510: 1451: 899:

cloned embryos created using a cell from a patient and a donated egg through the process of

5332: 5117: 5021: 4800: 4574: 4302: 4163: 4111: 4066: 3815: 3664: 3606: 3560: 3027: 2884: 2433: 2178: 1963: 1550: 1328: 1278: 863: 324: 288: 1653: 1636: 1369: 1123:, highly similar to embryonic stem cells, can be induced by the delivery of four factors ( 615:, estimated that it would take several months for the stem cells to replicate and for the 145: 8: 5715: 5705: 5649: 5437: 4764: 3066: 479: 471: 438: 206:

in the adult human body. When provided with the appropriate signals, ESCs initially form

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Waese, EY; Kandel, RA; Stanford, WL (2008). "Application of stem cells in bone repair".

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pluripotency of actively dividing stem cells. The problem was discovered when non-human

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model for early mouse development, EC cells harbor genetic mutations and often abnormal

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Due to their plasticity and potentially unlimited capacity for self-renewal, embryonic

228: 5381:

Understanding Stem Cells: A View of the Science and Issues from the National Academies

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formation when transplanted into a host environment in vivo, eradicating the risk of

1174: 859: 714:

1964: Lewis Kleinsmith and G. Barry Pierce Jr. isolated a single type of cell from a

612: 589: 417: 316: 122: 5269: 4986: 4943: 4330: 4039: 3264:"Great news: BioTime Subsidiary Asterias Acquires Geron Embryonic Stem Cell Program" 2965: 2699: 2595: 2551: 2508: 5570: 5484: 5350: 5340: 5293: 5249: 5214: 5176: 5135: 5125: 5039: 5029: 4966: 4923: 4885: 4808: 4744: 4714: 4704: 4677: 4647: 4637: 4610: 4582: 4517: 4495: 4487: 4442: 4401: 4383: 4310: 4279: 4257: 4249: 4214: 4191: 4171: 4139: 4119: 4074: 4019: 3943: 3916: 3894: 3886: 3823: 3682: 3672: 3634: 3614: 3568: 3481: 3471: 3430: 3389: 3381: 3340: 3332: 3123: 3119: 3115: 3045: 3035: 2988: 2943: 2902: 2892: 2843: 2835: 2791: 2783: 2722: 2679: 2642: 2575: 2531: 2488: 2451: 2441: 2394: 2357: 2296: 2255: 2237: 2196: 2186: 2137: 2127: 2086: 2078: 2036: 2028: 1987: 1971: 1922: 1906: 1865: 1849: 1808: 1790: 1749: 1741: 1700: 1692: 1681:"Concise Review: Control of Cell Fate Through Cell Cycle and Pluripotency Networks" 1648: 1621: 1599: 1558: 1506: 1496: 1422: 1414: 1336: 1059: 867: 810: 603:

In October 2010 researchers enrolled and administered ESCs to the first patient at

5380: 4828: 4079: 4054: 3828: 3803: 3321:"Deconstructing Stem Cell Tumorigenicity: A Roadmap to Safe Regenerative Medicine" 3073: 2226:"Embryonic Stem Cells and Tissue Engineering: Delivering Stem Cells to the Clinic" 1563: 1538: 1341: 1314: 5689: 5551: 5479: 5404: 5387: 5345: 5067: 4350: 3306: 3172: 2754: 2616: 1154: 1113:"for the discovery that mature cells can be reprogrammed to become pluripotent." 1094: 995: 983: 928: 916: 840: 787: 783: 760: 739: 715: 628: 604: 597: 573: 500: 272: 240: 211: 207: 126: 99: 72: 5677: 5620: 5615: 5582: 4709: 4692: 4642: 4625: 4491: 4447: 4430: 4388: 4253: 3890: 3435: 3418: 3417:

Wernig, Marius; Meissner, Alexander; Cassady, John P; Jaenisch, Rudolf (2008).

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functional ESC-derived hepatocytes with stable phase I and II enzyme activity.

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White, J.; Stead, E.; Faast, R.; Conn, S.; Cartwright, P.; Dalton, S. (2005).

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Cervantes RB, Stringer JR, Shao C, Tischfield JA, Stambrook PJ (March 2002).

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University of Oxford practical workshop on pluripotent stem cell technology

5364: 5305: 5261: 5253: 5226: 5190: 5149: 5053: 4978: 4935: 4820: 4736: 4669: 4602: 4509: 4464: 4415: 4355: 4322: 4271: 4031: 3957: 3908: 3526: 3495: 3444: 3403: 3354: 3137: 3059: 3040: 3000: 2916: 2857: 2805: 2734: 2691: 2656: 2587: 2543: 2500: 2465: 2406: 2371: 2316: 2269: 2210: 2191: 2151: 2132: 2100: 2050: 2001: 1936: 1895:"Distinct Cell-Cycle Control in Two Different States of Mouse Pluripotency" 1879: 1822: 1763: 1714: 1662: 1613: 1520: 1501: 1436: 1405: 1117: 1105:, who showed in 2006 that the introduction of four specific genes encoding 1064: 991: 936: 924: 756: 746: 723: 683: 522: 307:

play a role in transcriptionally regulating the embryonic stem cell cycle.

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for further purification. ESCs are predicted to be inherently safer than

474:. Studies have shown that cardiomyocytes derived from ESCs are validated 421: 413: 409: 215: 163: 53:: able to develop into any type of cell, including those of the placenta. 4586: 4314: 1975: 576:(a cell type of the brain and spinal cord) derived from human ESCs into 4652: 4500: 3899: 3711:"The 2007 Nobel Prize in Physiology or Medicine – Advanced Information" 2344:

Davila, JC; Cezar, GG; Thiede, M; Strom, S; Miki, T; Trosko, J (2004).

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to generate primitive ectoderm, which ultimately differentiates during

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or serum-free media supplements with two inhibitory drugs ("2i"), the

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that could arise with the creation of new lines under federal budget.

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Muse cells (Multi-lineage differentiating stress enduring cell) are

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On January 23, 2009, Phase I clinical trials for transplantation of

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Martin GR (1980). "Teratocarcinomas and mammalian embryogenesis".

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US scientists relieved as Obama lifts ban on stem cell research

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Aladjem MI, Spike BT, Rodewald LW, et al. (January 1998).

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Information & Alternatives to Embryonic Stem Cell Research

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Mao Z, Bozzella M, Seluanov A, Gorbunova V (September 2008).

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Fact sheet on ethical issues in embryonic stem cell research

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Ter Huurne, Menno; Stunnenberg, Hendrik G. (21 April 2021).

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Cardiovascular & Hematological Disorders Drug Targets

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characteristics, as demonstrated by the ability to form

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Lee, J.; Go, Y.; Kang, I.; Han, Y. M.; Kim, J. (2010).

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and that ES cells could be derived from these embryos.

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Takahashi, Kazutoshi; Yamanaka, Shinya (2006-08-25).

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Human embryonic stem cells have also been derived by

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and can only produce a limited number of cell types.

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However, a study published in the online edition of

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Techniques and conditions for derivation and culture

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were able to produce large quantities of pancreatic

106:

results in destruction of the blastocyst, a process

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Cappechi, 271:proteins involved in the 246: 129:have also been reported. 5601:Cellular differentiation 5182:10.3727/096368916X690881 2648:10.1093/eurheartj/ehu192 1069:Advanced Cell Technology 829:President George W. Bush 738:cells directly from the 277:Cyclin-dependent kinases 5561:Hematopoietic stem cell 5131:10.1073/pnas.1100816108 5035:10.1073/pnas.0911647107 4882:Nature Publishing Group 4813:10.1126/science.1152092 3678:10.1073/pnas.78.12.7634 3573:10.1126/science.6250214 2898:10.1073/pnas.0401346101 2447:10.1073/pnas.0404700101 2420:Perrier, A. L. (2004). 2289:Trends in Biotechnology 2021:The Biochemical Journal 1746:10.1091/mbc.e04-12-1056 951:(to prevent fibroblast 765:University of Cambridge 655:Concern and controversy 319:have been proposed for 310: 285:Retinoblastoma proteins 5254:10.1038/nprot.2013.076 3657:Proc Natl Acad Sci USA 3041:10.1073/pnas.062527199 2635:European Heart Journal 2350:Toxicological Sciences 2192:10.1073/pnas.032074999 2165:Levenberg, S. (2002). 1502:10.1186/1479-5876-9-29 1210:Lancet Medical Journal 1192:The online edition of 896:In vitro fertilization 891:Derivation from humans 875:President Barack Obama 820:1998: A team from the 803: 751: 458:European Heart Journal 385: 150: 142: 54: 38: 5721:1981 in biotechnology 5611:Stem cell controversy 5566:Mesenchymal stem cell 5556:Endothelial stem cell 5298:10.1089/scd.2013.0473 5163:Dezawa, Mari (2016). 3985:(46). 11 March 2009. 2840:10.1089/scd.2010.0058 2363:10.1093/toxsci/kfh100 2083:10.1083/jcb.200801009 1419:10.1038/embor.2009.37 1295:Stem cell controversy 1240:pluripotent stem cell 1107:transcription factors 1067:(medical director of 801: 749: 700:Stem cell controversy 531:chromosomal disorders 509:, a Russian-American 383: 321:regenerative medicine 148: 140: 79:, an early-stage pre- 44: 33: 5475:Embryonic stem cells 5169:Cell Transplantation 4890:10.1038/news050124-1 3767:A Clone of Your Own? 3076:. January 23, 2009. 2788:10.4161/cc.7.18.6679 2568:Nature Biotechnology 2133:10.1155/2016/6940283 1279:Fetal tissue implant 967:, and differentiate 935:and dosing her with 864:spinal cord injuries 541:Repair of DNA damage 472:small-molecule drugs 439:natural killer cells 325:spinal cord injuries 289:transcription factor 102:(embryoblast) using 94:stage 4–5 days post 58:Embryonic stem cells 5535:Embryonic stem cell 5337:2013PLoSO...864752H 5122:2011PNAS..108.9875W 5026:2010PNAS..107.8639K 4965:(9471): 1636–1641. 4805:2007Sci...318.1920H 4799:(5858): 1920–1923. 4765:The Washington Post 4587:10.1038/nature05944 4579:2007Natur.448..318W 4315:10.1038/nature05142 4307:2006Natur.444..481K 4168:1988Natur.336..684W 4116:1988Natur.336..688S 4071:1998Sci...282.1145T 4065:(5391): 1145–1147. 3820:1998Sci...282.1145T 3814:(5391): 1145–1147. 3669:1981PNAS...78.7634M 3611:1981Natur.292..154E 3565:1980Sci...209..768M 3270:. October 1, 2013. 3177:The Washington Post 3032:2002PNAS...99.3586C 2889:2004PNAS..10114443H 2883:(40): 14443–14448. 2715:Toxicology in Vitro 2438:2004PNAS..10112543P 2432:(34): 12543–12548. 2183:2002PNAS...99.4391L 1976:10.1038/nature06968 1968:2008Natur.453..519Y 1555:1998Sci...282.1145T 1399:Baldwing A (2009). 1333:1998Sci...282.1145T 1327:(5391): 1145–1147. 854:January, 2009: The 578:spinal cord-injured 513:who specialized in 480:torsades de pointes 341:Parkinson's disease 317:stem cell therapies 5403:2016-04-08 at the 5386:2010-04-09 at the 5094:Jul;28(7):907–923. 4349:2013-07-26 at the 3305:2017-10-24 at the 3171:2017-10-25 at the 2753:2009-08-08 at the 2615:2014-12-02 at the 2524:Skeletal Radiology 2033:10.1042/BJ20091439 1284:Induced stem cells 1171:sickle cell anemia 1169:, to cure mice of 804: 752: 527:prenatal diagnosis 511:medical researcher 497:Fragile-X syndrome 446:Harvard University 429:Clinical potential 390:tissue engineering 386: 376:Tissue engineering 151: 143: 55: 39: 5659: 5658: 5606:Stem cell therapy 5485:Cancer stem cells 5116:(24): 9875–9880. 5020:(19): 8639–8643. 4573:(7151): 318–324. 4301:(7118): 481–485. 4162:(6200): 684–687. 4110:(6200): 688–690. 3777:978-0-521-85294-4 3663:(12): 7634–7638. 3651:Martin G (1981). 3605:(5819): 154–156. 3559:(4458): 768–776. 3114:(19): 4694–4705. 2834:(11): 1699–1711. 2782:(18): 2902–2906. 2684:10.1002/jcp.21732 2574:(11): 1392–1401. 2493:10.1159/000101892 1962:(7194): 519–523. 1905:(4): 449–455.e4. 1789:(10): 4507–4519. 1697:10.1002/stem.2345 1454:Biomedical ethics 1374:stemcells.nih.gov 1173:, as reported by 994:. The extracted 860:Geron Corporation 834:ethical questions 627:, led by CEO Dr. 613:Geron Corporation 590:Geron Corporation 588:and supported by 287:that inhibit the 123:genetic disorders 71:derived from the 16:(Redirected from 5733: 5688: 5687: 5686: 5676: 5675: 5667: 5647: 5646: 5594:Related articles 5571:Neural stem cell 5480:Adult stem cells 5454: 5447: 5440: 5431: 5430: 5369: 5368: 5358: 5348: 5316: 5310: 5309: 5280: 5274: 5273: 5248:(7): 1391–1415. 5242:Nature Protocols 5237: 5231: 5230: 5213:(6): 1473–1481. 5201: 5195: 5194: 5184: 5160: 5154: 5153: 5143: 5133: 5101: 5095: 5091: 5085: 5084: 5082: 5081: 5064: 5058: 5057: 5047: 5037: 5004: 4991: 4990: 4954: 4948: 4947: 4911: 4905: 4904: 4902: 4901: 4892:. 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Evans 761:Matthew Kaufman 740:inner cell mass 716:teratocarcinoma 711: 702: 696: 662: 660:Adverse effects 657: 629:Michael D. West 605:Shepherd Center 598:Michael D. West 570: 564: 543: 501:Cystic fibrosis 492: 468: 431: 398: 378: 364: 313: 273:G1/S transition 249: 241:GSK-3 inhibitor 239:PD03259010 and 224: 208:precursor cells 172: 135: 100:inner cell mass 73:inner cell mass 28: 23: 22: 15: 12: 11: 5: 5739: 5729: 5728: 5723: 5718: 5713: 5708: 5693: 5692: 5680: 5657: 5656: 5654: 5653: 5640: 5637: 5636: 5634: 5633: 5628: 5623: 5621:Stem cell laws 5618: 5616:Stem cell line 5613: 5608: 5603: 5597: 5595: 5591: 5590: 5588: 5587: 5586: 5585: 5583:Precursor cell 5575: 5574: 5573: 5568: 5563: 5558: 5544: 5543: 5542: 5537: 5527: 5526: 5525: 5520: 5515: 5504: 5502: 5496: 5495: 5493: 5492: 5487: 5482: 5477: 5471: 5469: 5465: 5464: 5457: 5456: 5449: 5442: 5434: 5428: 5427: 5422: 5417: 5412: 5407: 5395: 5390: 5376: 5375:External links 5373: 5371: 5370: 5311: 5292:(7): 717–728. 5275: 5232: 5196: 5175:(5): 849–861. 5155: 5096: 5086: 5059: 4992: 4949: 4928:10.1038/nm1181 4922:(2): 228–232. 4906: 4864: 4834: 4782: 4750: 4703:(5): 861–872. 4683: 4636:(4): 663–676. 4616: 4556: 4530: 4486:(4): 663–676. 4470: 4441:(6): 777–780. 4435:Cell Stem Cell 4421: 4361: 4336: 4285: 4248:(5): 861–872. 4232: 4213:(2): 254–258. 4197: 4145: 4094: 4045: 4002: 3963: 3922: 3885:(4): 663–676. 3869: 3843: 3794: 3776: 3756: 3731: 3702: 3640: 3586: 3543: 3501: 3450: 3423:Cell Stem Cell 3409: 3360: 3311: 3285: 3255: 3225: 3192: 3180: 3157: 3091: 3065: 3006: 2987:(2): 145–178. 2971: 2942:(3): 145–155. 2922: 2863: 2828:Stem Cells Dev 2811: 2762: 2740: 2721:(5): 929–937. 2705: 2678:(3): 513–519. 2662: 2621: 2601: 2557: 2530:(7): 601–608. 2514: 2487:(4): 339–347. 2471: 2412: 2393:(2): 145–152. 2377: 2356:(2): 214–223. 2333: 2275: 2236:(8): 346–350. 2216: 2157: 2106: 2056: 2007: 1942: 1899:Cell Stem Cell 1885: 1848:(2): 141–149. 1842:Cell Stem Cell 1828: 1769: 1740:(4): 2018–27. 1720: 1691:(6): 1427–36. 1668: 1627: 1598:(3): 281–292. 1578: 1526: 1475: 1469:978-1601521576 1468: 1442: 1413:(4): 299–300. 1391: 1356: 1304: 1302: 1299: 1298: 1297: 1292: 1286: 1281: 1276: 1271: 1264: 1261: 1229:Main article: 1226: 1223: 1189: 1186: 1087:Main article: 1084: 1081: 1054: 1051: 986:from the late 912: 909: 892: 889: 887: 884: 883: 882: 871: 852: 837: 825: 818: 795: 780: 769:Gail R. Martin 744: 743: 710: 707: 698:Main article: 695: 694:Ethical debate 692: 690:transfection. 671:flow cytometry 661: 658: 656: 653: 594:Menlo Park, CA 566:Main article: 563: 562:Clinical trial 560: 542: 539: 507:Yury Verlinsky 491: 488: 467: 466:Drug discovery 464: 463: 462: 453: 442: 435: 430: 427: 402:cardiomyocytes 397: 394: 377: 374: 363: 360: 349:Ethical debate 312: 309: 248: 245: 223: 220: 171: 168: 134: 131: 26: 9: 6: 4: 3: 2: 5738: 5727: 5724: 5722: 5719: 5717: 5714: 5712: 5711:Biotechnology 5709: 5707: 5704: 5703: 5701: 5691: 5681: 5679: 5674: 5669: 5668: 5665: 5652: 5651: 5642: 5641: 5638: 5632: 5629: 5627: 5624: 5622: 5619: 5617: 5614: 5612: 5609: 5607: 5604: 5602: 5599: 5598: 5596: 5592: 5584: 5581: 5580: 5579: 5576: 5572: 5569: 5567: 5564: 5562: 5559: 5557: 5553: 5550: 5549: 5548: 5545: 5541: 5538: 5536: 5533: 5532: 5531: 5528: 5524: 5521: 5519: 5516: 5514: 5511: 5510: 5509: 5506: 5505: 5503: 5501: 5497: 5491: 5488: 5486: 5483: 5481: 5478: 5476: 5473: 5472: 5470: 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