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42 gesichtete, geschützte Fragmente: Plagiat

[1.] Pak/Fragment 023 01 - Diskussion
Bearbeitet: 26. April 2014, 09:43 Guckar
Erstellt: 6. April 2014, 09:52 (Hindemith)
BauernOpfer, Essential cell biology 2004, Fragment, Gesichtet, Pak, SMWFragment, Schutzlevel sysop

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Quelle: Essential cell biology 2004
Seite(n): -, Zeilen: Figure 8-10
23a diss Pak.png

Figure 1.3.4b: Above figure is a simple illustration of basal transcription regulation mediated by general transcription factors (GTFs) in a eukarayotic cell (Figure adapted from Essential cell biology E-2). To begin transcription, eukaryotic RNA polymerase II requires the general transcription factors. These transcription factors are called TFIIA, TFIIB, and so on. (A) The promoter contains a DNA sequence called the TATA box, which are located 25 nucleotides away from the site where transcription is initiated. (B) The TATA box is recognized and bound by transcription factor TFIID, which then enables the adjacent binding of TFIIB. (C) For simplicity the DNA distortion produced by the binding of TFIID is not shown. (D) The rest of the general transcription factors as well as the RNA polymerase itself assemble at the promoter. (E) TFIIH uses ATP to pry apart the double helix at the transcription start point, allowing transcription to begin. TFIIH also phosphorylates RNA polymerase II, releasing it from the general factors so it can begin the elongation phase of transcription. As shown, the site of phosphorylation is a long polypeptide tail that extends from the polymerase molecule.

23a source Pak.png

To begin transcription, eucaryotic RNA polymerase II requires the general transcription factors. These transcription factors are called TFIIA, TFIIB, and so on. (A) The promoter contains a DNA sequence called the TATA box, which is located 25 nucleotides away from the site where transcription is initiated. (B) The TATA box is recognized and bound by transcription factor TFIID, which then enables the adjacent binding of TFIIB. (C) For simplicity the DNA distortion produced by the binding of TFIID is not shown. (D) The rest of the general transcription factors as well as the RNA polymerase itself assemble at the promoter. (E) TFIIH uses ATP to pry apart the double helix at the transcription start point, allowing transcription to begin. TFIIH also phosphorylates RNA polymerase II, releasing it from the general factors so it can begin the elongation phase of transcription. As shown, the site of phosphorylation is a long polypeptide tail that extends from the polymerase molecule.

Anmerkungen

The figure is sufficiently referenced, but it is not clear to the reader that the extensive caption is also taken verbatim from the source.

Sichter
(Hindemith) Schumann

[2.] Pak/Fragment 028 12 - Diskussion
Bearbeitet: 6. April 2014, 21:12 Hindemith
Erstellt: 6. April 2014, 20:53 (Hindemith)
Abramovich et al 2002, Fragment, Gesichtet, KomplettPlagiat, Pak, SMWFragment, Schutzlevel sysop

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HPIP can bind to different members of the mammalian PBX family, inhibit the binding of PBX1/Hox complex to DNA and block the transcriptional activity of E2A-PBX1. HPIP can bind to the different members of the mammalian Pbx family, inhibit the binding of Pbx1/Hox complex to DNA and block the transcriptional activity of E2A-Pbx (Abramovich et al., 2000).
Anmerkungen

There is a reference to the source further down the page in the following paragraph for another sentence that is also taken from it.

Sichter
(Hindemith) Schumann

[3.] Pak/Fragment 028 07 - Diskussion
Bearbeitet: 6. April 2014, 20:55 Hindemith
Erstellt: 11. March 2014, 03:01 (Hindemith)
Abramovich et al 2000, BauernOpfer, Fragment, Gesichtet, Pak, SMWFragment, Schutzlevel sysop

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The new haematopoietic PBX-interacting protein (HPIP) was identified by a yeast two-hybrid screen of a fetal-liver haematopoietic cDNA library using PBX1 as bait. HPIP cDNA encodes a novel protein of 731 amino acid residues containing no homology to any known protein and has a calculated molecular mass of 80 kDa. HPIP is predicted to have a coiled-coil domain, suggesting that it interacts with other proteins. [...] The expression of PBX1 and HPIP was characterized observed by reverse transcription-PCR analysis of RNA obtained from bone marrow: HPIP expression was detected in the CD34+ fraction containing the haematopoietic progenitors and at lower levels in the CD34- matures cell population.

The same pattern was found for PBX1, indicating that HPIP and PBX1 are coexpressed in the same haematopoietic compartment.

A yeast two-hybrid screen of a fetal liver-hematopoietic cDNA library using PBX1a as bait led to the discovery of a novel non-homeodomaincontaining protein that interacts with PBX1 as well as PBX2 and PBX3.

[page 26174]

HPIP expression in primitive human hematopoietic cells was also demonstrated by reverse transcription-PCR analysis of RNA obtained from human bone marrow. Fig. 2B shows that HPIP is strongly expressed in the CD34+ fraction containing the hematopoietic progenitors and at lower levels in the CD34- mature cell population. The same pattern of expression was found for PBX1, indicating that HPIP and PBX1 are co-expressed in the same hematopoietic compartment.

[page 26176]

HPIP cDNA encodes a novel protein of 731 amino acid residues containing no homology to any known protein. The predicted HPIP protein has a calculated molecular mass of 80 kDa [...]. HPIP is predicted to have a coil-coil domain, suggesting that it interacts with other proteins.

Anmerkungen

The source is given just before the documented passage and it can be understood from the text that what follows is describing the work of the authors of the source. It is not clear, however, that the text is taken from the source.

Note: the sentence not taken from this source (line 13), can be found here: Pak/Fragment 028 12

Sichter
(Hindemith) Schumann

[4.] Pak/Fragment 012 21 - Diskussion
Bearbeitet: 6. April 2014, 18:34 Hindemith
Erstellt: 6. April 2014, 16:30 (Hindemith)
Fragment, Gesichtet, Pak, SMWFragment, Schutzlevel sysop, Stem Cell Information 2001, Verschleierung

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Quelle: Stem_Cell_Information_2001
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Bone Marrow: The classic source of HSCs is the bone marrow. About 1 in every 100,000 cells in the marrow is a long-term, blood-forming stem cell; other cells present include stromal cells, stromal stem cells, blood progenitor cells, and mature and maturing white and red blood cells.

Peripheral Blood: For clinical transplantation of human HSCs, cells are preferably harvested from peripheral circulating blood.

Bone Marrow

The classic source of hematopoietic stem cells (HSCs) is bone marrow. [...] About 1 in every 100,000 cells in the marrow is a long-term, blood-forming stem cell; other cells present include stromal cells, stromal stem cells, blood progenitor cells, and mature and maturing white and red blood cells.

Peripheral Blood

[...] For clinical transplantation of human HSCs, doctors now prefer to harvest donor cells from peripheral, circulating blood.

Anmerkungen

The source is not mentioned.

Sichter
(Hindemith) Schumann

[5.] Pak/Fragment 008 05 - Diskussion
Bearbeitet: 6. April 2014, 18:27 Hindemith
Erstellt: 6. April 2014, 15:18 (Hindemith)
Fragment, Gesichtet, Metcalf 2007, Pak, SMWFragment, Schutzlevel sysop, Verschleierung

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Human haematopoietic stem cells (hHSCs) are rare small mononuclear cells

that tend to be noncycling or to have long cell cycles. They divide to form more hHSCs (self-generation or symmetric division) [...] These immature progenitors are committing to lymphocyte progenitor formation (common lymphoid progenitors, CLPs) or to the formation of myeloid progenitors (common myeloid progenitors, CMPs). CMPs are large blast cells that can then form megakaryocyteerythroid progenitors (MEPs) committed to the formation of erythroid and megakaryocytic cell lineages. CMPs can also form more restricted granulocytemacrophage progenitors (GMPs) that are able to generate granulocytic, macrophage, and eosinophil progenitors. These lineage-restricted cells can generate respective mature lineage populations. 17-20


17.Kondo M, Scherer DC, Miyamoto T, et al. Cell-fate conversion of lymphoid-committed progenitors by instructive actions of cytokines. Nature. 2000;407:383-386.

18.Kondo M, Wagers AJ, Manz MG, et al. Biology of hematopoietic stem cells and progenitors: implications for clinical application. Annu Rev Immunol. 2003;21:759-806.

19.K A. Transcriptional accessibility for genes of multiple tissues and hematopoietic lineages is hierarchically controlled during early hematopoiesis. . Bllod. 2003;101:383-389.

20.Shizuru JA, Negrin RS, Weissman IL. Hematopoietic stem and progenitor cells: clinical and preclinical regeneration of the hematolymphoid system. Annu Rev Med. 2005;56:509-538.

Hematopoietic stem cells (HSCs) are rare (1 per 105 bone-marrow cells), small mononuclear cells that tend to be noncycling or to have long cell cycles. HSCs divide to form more HSC (self-generation) or to form cells committed either to lymphocyte formation (common lymphoid progenitors, CLPs) or to the formation of myeloid cells (common myeloid progenitors, CMPs) (Figure 1). CMPs are large blast cells that can then form megakaryocyteerythroid progenitors (MEPs) committed to the formation of erythroid and megakaryocytic progeny. CMPs can also form more restricted granulocyte- macrophage progenitors (GMPs) able to generate granulocytic, macrophage, and eosinophil progenitors and, through these lineage-restricted cells, generate respective mature populations (Kondo et al., 1997; Akashi et al., 2000).

Akashi, K., Traver, D., Miyamoto, T., and Weissman, I.L. (2000). A clonogenic common myeloid progenitor that gives rise to all myeloid lineages. Nature 404, 193–197.

Kondo, M., Weissman, I.L., and Akashi, K. (1997). Identification of clonogenic common lymphoid progenitors in mouse bone marrow. Cell 91, 661–672.

Anmerkungen

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(Hindemith) Schumann

[6.] Pak/Fragment 017 06 - Diskussion
Bearbeitet: 6. April 2014, 18:27 Hindemith
Erstellt: 6. April 2014, 15:32 (Hindemith)
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Individual cytokines can be lineage specific or can regulate cells in multiple lineages, and for some cell types, such as stem cells or megakaryocyte progenitors, the simultaneous action of multiple cytokines is required for proliferative responses. Individual cytokines can be lineage specific or can regulate cells in multiple lineages, and for some cell types, such as stem cells or megakaryocyte progenitors, the simultaneous action of multiple cytokines is required for proliferative responses.
Anmerkungen

The source is not mentioned here.

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(Hindemith) Schumann

[7.] Pak/Fragment 023 14 - Diskussion
Bearbeitet: 6. April 2014, 09:46 Hindemith
Erstellt: 10. March 2014, 04:14 (Hindemith)
Fragment, Gesichtet, KomplettPlagiat, Pak, SMWFragment, Schutzlevel sysop, Wikipedia cell signaling 2008

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[In the figure, the ligand called epidermal growth factor (EGF) binds to] the receptor (called EGFR). This activates the receptor to phosphorylate itself. The phosphorylated receptor binds to an adaptor protein (GRB2), which couples the signal to further downstream signaling processes. In Figure 3, the ligand (called epidermal growth factor (EGF)) binds to the receptor (called EGFR). This activates the receptor to phosphorylate itself. The phosphorylated receptor binds to an adaptor protein (GRB2) which couples the signal to further downstream signaling processes.
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Copied text starts on the previous page: Pak/Fragment_022_26 and continues on the following page.

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(Hindemith) Agrippina1

[8.] Pak/Fragment 019 04 - Diskussion
Bearbeitet: 6. April 2014, 09:11 Hindemith
Erstellt: 10. March 2014, 01:07 (Graf Isolan)
Fragment, Gesichtet, Hapel and Stanley 2006, Pak, SMWFragment, Schutzlevel sysop, Verschleierung

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1.3.3. Tyrosine kinase receptor and cell signalling

Many cytokines such as CSF-1, SCF and FL are homodimeric that share some sequence homology and are structurally similar to one another. These three cytokines/ growth factors have complex patterns of expression due to alternative mRNA splicing. This allows them to be expressed as either membrane-spanning, cell-surface or secreted glycoproteins. All three growth factors are widely expressed in all tissues and two of them, SCF and CSF-1, also affect non-haematopoietic as well as haematopoietic cells.

The receptors for the above cytokines are members of the platelet derived growth factor (PDGF) receptor family. Each receptor possesses an extra-cellular domain, consisting of a five immunoglobulin (Ig) like repeat. These repeats are: a heavily glycosylated with N-linked sugars, a trans-membrane domain, intra-cellular domains containing a juxta-membrane region, a src-related tyrosine kinase domain that is interrupted by a kinase insert domain, and a carboxy-terminal tail. The three amino terminal Ig-like domains incorporate the ligand binding domains of the SCF and CSF-1 receptors. Binding of this type of dimeric receptor to its cognate ligand stabilizes the non-covalent association between the two chains of the receptor at the cell surface and permits the trans-phosphorylation of the intra-cellular domain of one chain by the other.

Tyrosine phosphorylation in response to cytokine binding is not restricted to those proteins that are stably associated with the receptor. Regions containing tyrosines that are phosphorylated as a consequence of receptor activation act as docking sites for src-homology region 2 (SH2) domains of signaling and adaptor proteins. These proteins in turn may interact with plasma membrane associated proteins. An example is the association of recruited Grb2/Sos with Ras, which leads to their activation. The associated proteins may themselves become tyrosine phosphorylated.

Tyrosine Kinase Receptors

CSF-1, SCF and FL are homodimeric cytokines that share some sequence homology and structurally are similar to one another.36-40 All three cytokines have complex patterns of expression due to alternative mRNA splicing. This allows them to be expressed as either membrane-spanning, cell-surface or secreted glycoproteins. All three growth factors are widely expressed in all tissues and at least two of them, SCF and CSF-1, affect non-haematopoietic as well as haematopoietic cells.

The receptors for the above cytokines are members of the PDGF receptor family.39-42 Each receptor possesses an extra-cellular domain comprising five Ig-like repeats which are heavily glycosylated with N-linked sugars, a trans-membrane domain, and intra-cellular domains containing a juxta-membrane region, a src-related tyrosine kinase domain that is interrupted by a kinase insert domain, and a carboxy-terminal tail. The three amino terminal Ig-like domains incorporate the ligand binding domains of the SCF and CSF-1 receptors.

Binding of this type of dimeric receptor to its cognate ligand stabilises the non-covalent association between the two chains of the receptor at the cell surface and permits the trans-phosphorylation of the intra-cellular domain of one chain by the other.

Tyrosine phosphorylation in response to cytokine binding is not restricted to those proteins that are stably associated with the receptor. Regions containing tyrosines that are phosphorylated as a consequence of receptor activation act as docking sites for src homology region 2 (SH2) domains of signalling and adaptor proteins. These proteins in turn may interact with plasma membrane associated proteins. An example is the association of recruited Grb2/Sos with Ras, which leads to their activation. Or the associated proteins may themselves become tyrosine phosphorylated.


36. Anderson DM, Williams DE, Tushinski R et al. Alternate splicing of mRNA encoding human mast cell growth factor and localisation of the gene to chromosome 12q22-24. Cell Growth Differ 1991; 2:373-378.

37. Bazan JF. Genetic and structural homology of stem cell factor and macrophage colony stimulating factor. Cell 1991; 65:9-10.

38. Flanagan JG, Chan DC, Leder P. Transmembrane form of the kit ligand growth factor is determined by alternative splicing and is missing in the Sld mutant. Cell 1991; 64:1025-1035.

39. Lyman SD, Jacobsen SEW. c-kit ligand and flt-3 ligand : stem cell factors with overlapping yet distinct activities. Blood 1998; 91:1101–1134.

40. Stanley ER. CSF-1. In: Oppenheim JJ, Feldmann M, eds. Cytokine Database. London: Academic 2000:913–934.

41. Qui F, Ray P, Brown K et al. Primary structure of c-kit: relationship with the CSF1/PDGF receptor kinase family - oncogenic activation of v-kit involves deletion of extracellular domain and C terminus. EMBO J 1988; 7:1003-1011.

42. Coussens L, van Beveren C, Smith D et al. Structural alteration of viral homologue of receptor proto oncogene fms at carboxy-terminus. Nature 1986; 32:277-280.

Anmerkungen

A literal copy except for minute details, without any part of it marked as a citation. No source given.

Sichter
(Graf Isolan) Agrippina1

[9.] Pak/Fragment 053 05 - Diskussion
Bearbeitet: 6. April 2014, 09:00 Hindemith
Erstellt: 9. March 2014, 07:40 (SleepyHollow02)
Ahmed 2007, Fragment, Gesichtet, KomplettPlagiat, Pak, SMWFragment, Schutzlevel sysop

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3.2.11. Colony forming cell assay

Haematopoietic colony-forming cells (CFC) were assayed using methylcellulose-based medium (MethoCult H 4434) (fig. 3.2.11). Required number of pre-aliquoted tubes of MethoCult medium was thawed overnight under refrigeration (2-8°C) or at room temperature. The cells were diluted with IMDM + 2% FBS to 10X the final concentration(s) required for plating. For a duplicate assay 0.3 ml of diluted cells was added to 3 ml MethoCult tube and the contents vortexed vigorously. After about 5 minutes, 1.1 ml of cell: methylcellulose mixture was dispensed into 35 mm culture dishes using sterile a 3 ml syringe and16-gauge bunt-end needle. The 35 mm culture dishes were placed into 10 cm petri dish along with an extra 35 mm dish containing sterile water to maintain humidity and placed in a CO2 incubator at 37°C and >95% humidity.

CFC numbers were evaluated after an incubation period of 12-14 days and distinguished into following classes: Colony-forming unit-erythroid (CFU-E): Produces 1-2 cell clusters containing a total of 8-200 erythroblasts. A CFU-E consists of mature erythroid progenitors that require erythropoietin (EPO) for differentiation. Burst-forming unit-erythroid (BFU-E): Produces a colony containing >200 erythroblasts in a single or multiple clusters. A BFU-E consists of more immature progenitors than CFU-E and requires EPO and cytokines with burst-promoting activity such as Interleukin-3 (IL-3) and Stem Cell Factor (SCF) for optimal colony growth.

Colony-forming unit-granulocyte, macrophage (CFU-GM): Produces a colony containing at least 20 granulocyte cells (CFU-G), macrophages (CFU-M) or cells of both lineages (CFU-GM). CFU-GM colonies arising from primitive progenitors may contain thousands of cells in single or multiple clusters. Colony-forming unit-granulocyte, erythroid, macrophage, megakaryocyte (CFU-GEMM):

3.2.8 Human CFC Assay

Haematopoietic colony-forming cells (CFC) were assayed using methylcellulose-based medium (MethoCult; H 4434) (fig 3.3). Required number of pre-aliquoted tubes of MethoCult medium was thawed overnight under refrigeration (2-8°C) or at room temperature. The cells were diluted with IMDM + 2% FBS to 10X the final concentration(s) required for plating. For a duplicate assay 0.3 ml of diluted cells was added to 3 ml MethoCult tube and the contents vortexed vigorously. After about 5 minutes, 1.1 ml of cell: methylcellulose mixture was dispensed into 35 mm culture dishes using sterile a 3 ml syringe and16-gauge bunt-end needle. The 35 mm culture dishes were placed into 10 cm petri dish along with an extra 35 mm dish containing sterile water to maintain humidity and placed in a CO2 incubator at 37°C and >95%

[Page 43]

humidity. CFC numbers were evaluated after an incubation period of 12-14 days and distinguished into following classes:

Colony-forming unit-erythroid (CFU-E): Produces 1-2 cell clusters containing a total of 8-200 erythroblasts. A CFU-E consists of mature erythroid progenitors that require erythropoietin (EPO) for differentiation.

Burst-forming unit-erythroid (BFU-E): Produces a colony containing >200 erythroblasts in a single or multiple clusters. A BFU-E consists of more immature progenitors than CFU-E and require [sic] EPO and cytokines with burst-promoting activity such as Interleukin-3 (IL-3) and Stem Cell Factor (SCF) for optimal colony growth.

Colony-forming unit-granulocyte, macrophage (CFU-GM): Produces a colony containing at least 20 granulocyte cells (CFU-G), macrophages (CFU-M) or cells of both lineages (CFU-GM). CFU-GM colonies arising from primitive progenitors may contain thousands of cells in single or multiple clusters.

Colony-forming unit-granulocyte, erythroid, macrophage, megakaryocyte (CFU-GEMM):

Anmerkungen

A literal, unattributed copy that continues on teh next page.

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(SleepyHollow02) Agrippina1

[10.] Pak/Fragment 085 02 - Diskussion
Bearbeitet: 6. April 2014, 08:36 Hindemith
Erstellt: 10. March 2014, 22:54 (Hindemith)
Fragment, Gesichtet, KomplettPlagiat, Pak, SMWFragment, Schutzlevel sysop, Wikipedia hematopoietic stem cell 2008

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Balanced HSCs repopulate peripheral white blood cells in the same ratio of myeloid to lymphoid cells as seen in unmanipulated mice (on average about 15% myeloid and 85% lymphoid cells, or 3≤ñ≤10 [sic]). Balanced HSCs repopulate peripheral white blood cells in the same ratio of myeloid to lymphoid cells as seen in unmanipulated mice (on average about 15% myeloid and 85% lymphoid cells, or 3≤ρ≤10).
Anmerkungen

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(Hindemith) Singulus

[11.] Pak/Fragment 013 17 - Diskussion
Bearbeitet: 6. April 2014, 08:36 Hindemith
Erstellt: 11. March 2014, 00:05 (Hindemith)
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The long-term reconstitution potential of human HSCs has been described in SCID mouse repopulating assays. In the most advanced xenotransplant models by utilizing the Rag2 and IL-2Rγ double-deficient mouse40 or the NOD/SCID IL2Rγ deficient mouse.41 The CD34+CD38- or the CD34+CD90+ fractions of human bone marrow and cord blood have been shown to reconstitute all human hematolymphoid lineage cells for a long term, indicating that these CD34+ fractions contains normal human HSCs. An injection of at least 1000 cells of the human CD34+CD38- bone marrow cells is required to obtain multi-lineage, longterm reconstitution in xenotransplant models. The percent of hCD34+CD38- or hCD34+CD90+ cells that are multipotent, long-term HSCs is still difficult to determine in a engraft model.

40. Traggiai E, Chicha L, Mazzucchelli L, et al. Development of a human adaptive immune system in cord blood cell-transplanted mice. Science. 2004;304:104-107.

41. Ishikawa F, Yasukawa M, Lyons B, et al. Development of functional human blood and immune systems in NOD/SCID/IL2 receptor {gamma} chain(null) mice. Blood. 2005;106:1565-1573.

The long-term reconstitution potential of human HSCs has been directly shown in SCID mouse repopulating assays. In the most advanced xenotransplant models by utilizing the Rag2 and IL-2Rγ double-deficient mouse (Traggiai et al., 2004) or the NOD–SCID IL2Rγ deficient mouse (Ishikawa et al., 2005), hCD34+hCD38- or the hCD34+hCD90+ fractions of human bone marrow and cord blood can reconstitute all human hematolymphoid lineage cells for a long term, indicating that these hCD34+ fractions contains normal human HSCs. [...] injection of at least 1000 cells of the human hCD34+hCD38- bone marrow cells is required to obtain multi-lineage, long-term reconstitution in xenotransplant models. Thus, it is unknown whether the HSC that is capable of reconstitution of all lineages at the single cell level exists in human hematopoiesis, and if so, what percent of hCD34+hCD38- or hCD34+hCD90+ cells are multipotent, longterm HSCs.

Traggiai E, Chicha L, Mazzucchelli L, Bronz L, Piffaretti JC, Lanzavecchia A et al. (2004). Development of a human adaptive immune system in cord blood cell-transplanted mice. Science 304: 104–107.

Ishikawa F, Yasukawa M, Lyons B, Yoshida S, Miyamoto T, Yoshimoto G et al. (2005). Development of functional human blood and immune systems in NOD/SCID/IL2 receptor {gamma} chain(null) mice. Blood 106: 1565–1573.

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(Hindemith) Schumann

[12.] Pak/Fragment 014 01 - Diskussion
Bearbeitet: 6. April 2014, 08:36 Hindemith
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1.2.2. The subsets of human lymphoid and myeloid progenitor.

The existence of lymphoid-committed progenitors in the human bone marrow with the hCD34+CD38+CD45RA+CD10+ phenotype has been reported.42 In human cord blood, common lymphoid progenitors (CLP) activity is found in the CD7+ fraction of CD34+CD38-CD45RA+ population. The hCD34+CD38- CD45RA+CD7+ population do not exist in the adult bone marrow.43 IL-7Rα, a critical marker for the murine CLP, is expressed in human hCD10+ CLPs in the bone marrow, but not in hCD7+ CLPs in the cord blood. These data suggest that the CLP phenotype and the requirement of IL-7 signaling might have changed during the human ontogeny. In the myelo-erythroid pathway, CMP, GMP and MEP subsets can be isolated from the hCD34+CD38+ fraction in both the bone marrow and cord blood (Manz et al., 2002). All these fractions of cells are negative for the early lymphoid markers hCD10, hCD7 or hIL-7Rα. These myelo-erythroid progenitors can be prospectively isolated according to the expression of hCD45RA and hIL-3Rα. CD45RA-hIL-3Rαlo (CMPs), hCD45RA+hIL-3Rαlo (GMPs) and hCD45RA-hIL-3Rα(MEPs) efficiently form distinct myelo-erythroid colony types according to their definitions (Fig.1.1.2). CMPs give rise to MEPs and GMPs in vitro, and a significant proportion of CMPs possess clonal GM and MegE potentials.44 Thus, the hierarchical myeloid progenitor relationships demonstrated in mice is well preserved in human haematopoiesis.

Phenotypic comparisons between mouse and human subsets show that CD34, a marker positive only for murine CMPs and GMPs, is uniformly expressed on all three human subsets, and that the FcγRII/III (CD16/CD32), marking murine CMPs and GMPs, was not detectable in any of the human myeloid progenitors44 All haemato/lymphoid progenitors develop from CD34+CD38- HSC population 41but themselves have no self-renewal activity in xenogenic transplantation models 45, indicating that they are downstream of human LTHSCs in both the bone marrow and the cord blood.

Flt3 shows a significant difference in its distribution in human and mouse hematopoiesis. This suggests a critical role of Flt3 signaling in hematopoietic [development in humans.]


41. Ishikawa F, Yasukawa M, Lyons B, et al. Development of functional human blood and immune systems in NOD/SCID/IL2 receptor {gamma} chain(null) mice. Blood. 2005;106:1565-1573.

42. Galy AH, Cen D, Travis M, Chen S, Chen BP. Delineation of T-progenitor cell activity within the CD34+ compartment of adult bone marrow. Blood. 1995;85:2770-2778.

43. Hao Z, Rajewsky K. Homeostasis of peripheral B cells in the absence of B cell influx from the bone marrow. J Exp Med. 2001;194:1151-1164.

44. Manz MG, Miyamoto T, Akashi K, Weissman IL. Prospective isolation of human clonogenic common myeloid progenitors. Proc Natl Acad Sci U S A. 2002;99:11872-11877.

45. Ishikawa F, Niiro H, Iino T, et al. The developmental program of human dendritic cells is operated independently of conventional myeloid and lymphoid pathways. Blood. 2007;110:3591-3660.

Human lymphoid and myeloid progenitor subsets within the hCD34+hCD38+ MPP fraction

Galy et al. (1995) first reported the existence of lymphoid-committed progenitors in the human bone marrow with the hCD34+hCD38+hCD45RA+hCD10+ phenotype. [...] In human cord blood, CLP activity is found in the hCD7+ fraction of hCD34+hCD38-hCD45RA+ population (Hao et al., 2001). Interestingly, the hCD7+ hCD34+hCD38-hCD45RA+ population does not exist in the adult bone marrow (Hao et al., 2001). IL-7Rα, a critical marker for the murine CLP, is expressed in human hCD10+ CLPs in the bone marrow, but not in hCD7+ CLPs in the cord blood. These data suggest that the CLP phenotype and the requirement of IL-7 signaling may change during human ontogeny.

In the myelo-erythroid pathway, CMP, GMP and MEP subsets are isolatable within the hCD34+hCD38+ fraction in both the bone marrow and cord blood (Manz et al., 2002). All are negative for the early lymphoid markers hCD10, hCD7 or hIL-7Rα. These myeloerythroid progenitors are prospectively isolatable according to the expression of hCD45RA and hIL-3Rα. hCD45RA-hIL-3Rαlo (CMPs), hCD45RA+hIL-3Rαlo (GMPs) and hCD45RA-hIL-3Rα- (MEPs) efficiently formed distinct myelo-erythroid colony types according to their definitions. CMPs give rise to MEPs and GMPs in vitro, and a significant proportion of CMPs possess clonal GM and MegE potentials (Manz et al., 2002). Thus, the hierarchical myeloid progenitor relationships demonstrated in mice is well preserved in human hematopoiesis. Phenotypic comparisons between mouse and human subsets show that CD34, a marker positive for only murine CMPs and GMPs, is uniformly expressed on all three human subsets, and that the FcγRII/III (CD16/CD32), marking murine CMPs and GMPs, was not detectable in none of human myeloid progenitors (Manz et al., 2002).

All of these hematolymphoid progenitors develop from hCD34+hCD38- HSC population (Ishikawa et al., 2005), but themselves have no self-renewal activity in xenogeneic transplantation models (Ishikawa et al., 2007), indicating that they are downstream of human LT-HSCs in both the bone marrow and the cord blood.

[page 6693]

The significant difference of Flt3 distribution in human and mouse hematopoiesis suggests that the critical role of Flt3 signaling in hematopoietic development could also be different.


Galy A, Travis M, Cen D, Chen B. (1995). Human T, B, natural killer, and dendritic cells arise from a common bone marrow progenitor cell subset. Immunity 3: 459–473.

Hao QL, Zhu J, Price MA, Payne KJ, Barsky LW, Crooks GM. (2001). Identification of a novel, human multilymphoid progenitor in cord blood. Blood 97: 3683–3690.

Ishikawa F, Niiro H, Iino T, Yoshida S, Saito N, Onohara S et al. (2007). The developmental program of human dendritic cells is operated independently of conventional myeloid and lymphoid pathways. Blood (in press).

Ishikawa F, Yasukawa M, Lyons B, Yoshida S, Miyamoto T, Yoshimoto G et al. (2005). Development of functional human blood and immune systems in NOD/SCID/IL2 receptor {gamma} chain(null) mice. Blood 106: 1565–1573.

Manz MG, Miyamoto T, Akashi K, Weissman IL. (2002). Prospective isolation of human clonogenic common myeloid progenitors. Proc Natl Acad Sci USA 99: 11872–11877.

Anmerkungen

The source is not mentioned at all.

Note that the reference "(Manz et al., 2002)" is atypical here as typically a reference would refer by a number to the bibliography, without giving the author's name in the main body of text.

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(Hindemith) Schumann

[13.] Pak/Fragment 015 01 - Diskussion
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hFlt3 is expressed in leukemic blasts in most cases with acute myelogenous leukemia (AML)46,47 (Carow et al., 1996; Rosnet et al., 1996), and FLT3 is one of the most frequently mutated genes in AML.48,49 The signal from FLT3 mutations should be controlled under the regulation of normal Flt3

[FIGURE 1.2.2, see: Pak/Fragment 015 05]

expression machinery, signaling from FLT3 mutations should involve HSCs and GMPs, both of which are critical targets for leukemic transformation in mouse AML models.51-54 Thus, special considerations are required in utilizing mouse models to understand the role of FLT3 mutations in human leukemogenesis.


46.Carow CE, Levenstein M, Kaufmann SH, et al. Expression of the hematopoietic growth factor receptor FLT3 (STK-1/Flk2) in human leukemias. Blood. 1996;87:1089-1096.

47.Rosnet O, Buhring HJ, Marchetto S, et al. Human FLT3/FLK2 receptor tyrosine kinase is expressed at the surface of normal and malignant hematopoietic cells. Leukemia. 1996;10:238-248.

48.Gilliland DG, Griffin JD. The roles of FLT3 in hematopoiesis and leukemia. Blood. 2002;100:1532- 1542.

49.Stirewalt DL, Meshinchi S, Kussick SJ, et al. Novel FLT3 point mutations within exon 14 found in patients with acute myeloid leukaemia. Br J Haematol. 2004;124:481-484.

51.Cozzio A, Passegue E, Ayton PM, Karsunky H, Cleary ML, Weissman IL. Similar MLL-associated leukemias arising from self-renewing stem cells and short-lived myeloid progenitors. Genes Dev. 2003;17:3029-3035.

52.So CW, Karsunky H, Passegue E, Cozzio A, Weissman IL, Cleary ML. MLL-GAS7 transforms multipotent hematopoietic progenitors and induces mixed lineage leukemias in mice. Cancer Cell. 2003;3:161-171.

53.Huntly BJ, Shigematsu H, Deguchi K, et al. MOZ-TIF2, but not BCR-ABL, confers properties of leukemic stem cells to committed murine hematopoietic progenitors. Cancer Cell. 2004;6:587-596.

54.Wang GG, Pasillas MP, Kamps MP. Meis1 programs transcription of FLT3 and cancer stem cell character, using a mechanism that requires interaction with Pbx and a novel function of the Meis1 Cterminus. Blood. 2005;106:254-264.

hFlt3 is expressed in leukemic blasts in most cases with acute myelogenous leukemia (AML) (Carow et al., 1996; Rosnet et al., 1996), and FLT3 is one of the most frequently mutated genes in AML (Gilliland, 2002; Stirewalt and Radich, 2003). [...] Because the signal from FLT3 mutations should be controlled under the regulation of normal Flt3 expression machinery, signaling from FLT3 mutations should involve HSCs and GMPs, both of which are critical targets for leukemic transformation in mouse AML models (see the next section) (Cozzio et al., 2003; So et al., 2003; Huntly et al., 2004; Wang et al., 2005). Thus, special considerations are required in utilizing mouse models to understand the role of FLT3 mutations in human leukemogenesis.

Carow CE, Levenstein M, Kaufmann SH, Chen J, Amin S, Rockwell P et al. (1996). Expression of the hematopoietic growth factor receptor FLT3 (STK-1/Flk2) in human leukemias. Blood 87: 1089–1096.

Cozzio A, Passegue E, Ayton PM, Karsunky H, Cleary ML, Weissman IL. (2003). Similar MLL-associated leukemias arising from self-renewing stem cells and short-lived myeloid progenitors. Genes Dev 17: 3029–3035.

Gilliland DG. (2002). Molecular genetics of human leukemias: new insights into therapy. Semin Hematol 39: 6–11.

Huntly BJ, Shigematsu H, Deguchi K, Lee BH, Mizuno S, Duclos N et al. (2004). MOZ-TIF2, but not BCR-ABL, confers properties of leukemic stem cells to committed murine hematopoietic progenitors. Cancer Cell 6: 587–596.

Rosnet O, Buhring HJ, Marchetto S, Rappold I, Lavagna C, Sainty D et al. (1996). Human FLT3/FLK2 receptor tyrosine kinase is expressed at the surface of normal and malignant hematopoietic cells. Leukemia 10: 238–248.

So CW, Karsunky H, Passegue E, Cozzio A, Weissman IL, Cleary ML. (2003). MLL-GAS7 transforms multipotent hematopoietic progenitors and induces mixed lineage leukemias in mice. Cancer Cell 3: 161–171.

Stirewalt DL, Radich JP. (2003). The role of FLT3 in haematopoietic malignancies. Nat Rev Cancer 3: 650–665.

Wang J, Iwasaki H, Krivtsov A, Febbo PG, Thorner AR, Ernst P et al. (2005). Conditional MLL-CBP targets GMP and models therapyrelated myeloproliferative disease. EMBO J 24: 368–381.

Anmerkungen

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Note that against the usual convention in the thesis the references "(Carow et al., 1996; Rosnet et al., 1996)" are given in the main body of text.

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[14.] Pak/Fragment 016 01 - Diskussion
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Further phenotypic and functional characterization of human hematopoietic cells should be performed by utilizing improved efficient xenotransplant models. These approaches will eventually help to identify LSCs that should be a critical cellular target in leukemia treatment.55

55.Iwasaki H, Mizuno S, Mayfield R, et al. Identification of eosinophil lineage-committed progenitors in the murine bone marrow. J Exp Med. 2005;201:1891-1897.

Further phenotypic and functional characterization of human hematopoietic cells should be performed by utilizing improved efficient xenotransplant models. These approaches will eventually help to identify LSCs that should be a critical cellular target in leukemia treatment.
Anmerkungen

The source is not given.

The text cannot be found in the referenced publication.

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[15.] Pak/Fragment 027 01 - Diskussion
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PBX1 is a homeodomain protein that functions in complexes with other homeodomain-containing proteins to regulate gene expression during development and/or differentiation processes. PBX is a member of the PBC protein family. The human PBX1 protein was initially identified as the chromosome 1 participant of the t(1;19) translocation, which occurs in 25 % of paediatric pre-B cell acute lymphocytic leukaemia which creates a chimeric gene designated E2A-PBX1.102

The mechanism by which E2A-PBX1 causes leukaemia is still unclear. However, the structure of the fusion protein, in which the majority of PBX1, including the homeodomain, is fused to the transcriptional activation domain of suggests that the oncogenic properties of E2A-PBX1 result from inappropriate regulation of target genes, of which the expression during haematopoiesis is normally regulated by wild-type PBX proteins.103,104 In vitro and in vivo data suggest that PBX functions in combination with heterologous homeodomain proteins, including class I HOX proteins. As HOX cofactors, PBC proteins improve HOX specificity due to the increased size of the cooperative binding site and the strength of DNA binding sites by different groups of HOX proteins.89,93

In addition, cooperative DNA binding with PBC proteins may act to change the regulatory signal of HOX proteins, from repressors to activators.78 PBX proteins appear to function as part of large nucleoprotein complexes. The interactions within these complexes are probably decisive factors that allow the DNA binding proteins to discriminate among target regulatory elements. How these complexes are regulated during either early embryonic development or cellular differentiation of somatic cells to control gene expression is still unclear. Abramovich et al. (2000) speculated that the characterization of additional PBX-interacting proteins might shed light on the mechanism of PBX function, and specifically sought to identify novel cofactors or modifiers of PBX1.105 Although the PBX homeodomain protein is thought to function as a transcription factor, its mechanism of action is still unknown.


78.Pineault N, Helgason CD, Lawrence HJ, Humphries RK. Differential expression of Hox, Meis1, and Pbx1 genes in primitive cells throughout murine hematopoietic ontogeny. Exp Hematol. 2002;30:49-57.

89.Mann RS, Chan SK. Extra specificity from extradenticle: the partnership between HOX and PBX/EXD homeodomain proteins. Trends Genet. 1996;12:258-262.

93.Shen WF, Montgomery JC, Rozenfeld S, et al. AbdB-like Hox proteins stabilize DNA binding by the Meis1 homeodomain proteins. Mol Cell Biol. 1997;17:6448-6458.

102.Kamps MP, Look AT, Baltimore D. The human t(1;19) translocation in pre-B ALL produces multiple nuclear E2A-Pbx1 fusion proteins with differing transforming potentials. Genes Dev. 1991;5:358-368.

103.Thorsteinsdottir U, Krosl J, Kroon E, Haman A, Hoang T, Sauvageau G. The oncoprotein E2APbx1a collaborates with Hoxa9 to acutely transform primary bone marrow cells. Mol Cell Biol. 1999;19:6355-6366.

104.LeBrun DP, Cleary ML. Fusion with E2A alters the transcriptional properties of the homeodomain protein PBX1 in t(1;19) leukemias. Oncogene. 1994;9:1641-1647.

105.Abramovich C, Shen WF, Pineault N, et al. Functional cloning and characterization of a novel nonhomeodomain protein that inhibits the binding of PBX1-HOX complexes to DNA. J Biol Chem. 2000;275:26172-26177.

PBX1 is a homeodomain protein that functions in complexes with other homeodomain-containing proteins to regulate gene expression during developmental and/or differentiation processes. [...]

[...] PBX1, a member of the PBX family along with PBX2 and PBX3 (4), was initially identified as the chromosome 1 participant of the t(1;19) translocation, which occurs in 25% of pediatric pre-B cell acute lymphocytic leukemia and that creates a chimeric gene designated E2A-PBX1 (5, 6). The mechanism by which E2A-PBX1 causes leukemia is still unclear. However, the structure of the protein, in which the majority of PBX1, including the homeodomain, is fused to the transcriptional activation domain of E2A (6, 7), suggests that the oncogenic properties of E2A-PBX1 result from inappropriate regulation of target genes whose expression during hematopoiesis is normally regulated by wild type PBX proteins (8–11).

In vitro and in vivo data strongly suggest that PBX functions in combination with heterologous homeodomain proteins, including class I HOX proteins. As HOX cofactors, PBC proteins improve HOX specificity due to the increased size of the cooperative binding site and the strength of DNA binding, as well as by modulating recognition of cooperative binding sites by different groups of HOX proteins (12–14). In addition, cooperative DNA binding with PBC proteins may act to change the regulatory signal of HOX proteins, from repressors to activators (15). [...]

[...]

PBX proteins thus appear to function as part of large nucleoprotein complexes. The interactions within these complexes are probably decisive factors that allow the DNA binding proteins to discriminate among target regulatory elements. How these complexes are regulated during either early embryonic development or cellular differentiation of somatic cells to control gene expression is still unclear. We speculated that characterization of additional PBX-interacting proteins might shed

[page 26173]

light on the mechanism of PBX function, and specifically, we sought to identify novel cofactors or modifiers of PBX1 using the yeast two-hybrid system.

[page 26176]

Although PBX homeodomain protein is thought to function as a transcription factor, its mechanism of action remains unknown.


4. Monica, K., Galili, N., Nourse, J., Saltman, D., and Cleary, M. L. (1991) Mol. Cell. Biol. 11, 6149–6157

5. Kamps, M. P., Look, A. T., and Baltimore, D. (1991) Genes Dev. 5, 358–368

6. Nourse, J., Mellentin, J. D., Galili, N., Wilkinson, J., Stanbridge, E., Smith S. D., and Cleary, M. L. (1990) Cell 60, 535–545

7. Kamps, M. P., Murre, C., Sun, X. H., and Baltimore, D. (1990) Cell 60, 547–555

8. LeBrun, D. P., and Cleary, M. L. (1994) Oncogene 9, 1641–1647

9. Lu, Q., Knoepfler, P. S., Scheele, J., Wright, D. D., and Kamps, M. P. (1995) Mol. Cell. Biol. 15, 3786–3795

10. Van Dijk, M. A., Voorhoeve, P. M., and Murre, C. (1993) Proc. Natl. Acad. Sci. U. S. A. 90, 6061–6065

11. Thorsteinsdottir, U., Krosl, J., Kroon, E., Haman, A., Hoang, T., and Sauvageau, G. (1999) Mol. Cell. Biol. 19, 6355–6366

12. Mann, R. S., and Chan, S. K. (1996) Trends Genet. 12, 258–262

13. Shen, W. F., Chang, C. P., Rozenfeld, S., Sauvageau, G., Humphries, R. K., Lu, M., Lawrence, H. J., Cleary, M. L., and Largman, C. (1996) Nucleic Acids Res. 24, 898–906

14. Shen, W. F., Rozenfeld, S., Lawrence, H. J., and Largman, C. (1997) J. Biol. Chem. 272, 8198–8206

15. Pinsonneault, J., Florence, B., Vaessin, H., and McGinnis, W. (1997) EMBO J. 16, 2032–2042

Anmerkungen

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Note that also most of the references are parallel in the thesis and the source.

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[16.] Pak/Fragment 103 21 - Diskussion
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HPIP-WT is strongly expressed in the hCB CD34+ fraction containing the haematopoietic progenitors and at lower levels in the hCB CD34- fraction comprising the mature cell population. The same pattern of expression was found for PBX1, indicating that HPIP and PBX1 are co-expressed in the same human haematopoietic cell compartment. Fig. 2B shows that HPIP is strongly expressed in the CD34+ fraction containing the hematopoietic progenitors and at lower levels in the CD34- mature cell population. The same pattern of expression was found for PBX1, indicating that HPIP and PBX1 are co-expressed in the same hematopoietic compartment.
Anmerkungen

The source is given two sentences further down. Those two sentences have not been taken from the source, at least not literally.

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[17.] Pak/Fragment 016 06 - Diskussion
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Using limiting dilution strategies combined with other streamlined experimental and statistical methods for examining HSCs at the clonal level, HSCs fall into three distinct lineage-biased clusters.56-58 These are quantitatively defined by the ratio “ρ” of lymphoid to myeloid cells that the HSC generates upon differentiation (which makes ñ [sic] a peripheral predictor for the clonal association of a reconstituted haematopoietic system). Balanced HSCs repopulate peripheral white blood cells in the same ratio of myeloid to lymphoid cells as seen in unmanipulated mice (on average about 15% myeloid and 85% lymphoid cells, or 3≤ñ≤10 [sic]), in the NOD/SCID mice. Myeloid-biased (My-bi) HSC give rise to too few lymphocytes resulting in ratios 0<ρ<3, whereas lymphoid-biased (Ly-bi) HSC generate too few myeloid cells, which results in lymphoid-to-myeloid ratios of 10<ρ<00 [sic]. All three types are normal HSCs in the sense that they have self-renewal capacity and can regenerate all haematopoietic lineages (pluripotency).

56. Muller-Sieburg [sic] CE, Cho RH, Thoman M, Adkins B, Sieburg HB. Deterministic regulation of hematopoietic stem cell self-renewal and differentiation. Blood. 2002;100:1302-1309.

57. Muller-Sieburg [sic] CE, Cho RH, Karlsson L, Huang JF, Sieburg HB. Myeloid-biased hematopoietic stem cells have extensive self-renewal capacity but generate diminished lymphoid progeny with impaired IL-7 responsiveness. Blood. 2004;103:4111-4118.

58. Sieburg HB, Cho RH, Dykstra B, Uchida N, Eaves CJ, Muller-Sieburg [sic] CE. The hematopoietic stem compartment consists of a limited number of discrete stem cell subsets. Blood. 2006;107:2311-2316.

Using limiting dilution strategies combined with other streamlined experimental and statistical methods for examining HSCs at the clonal level, it was shown that HSCs fall into three distinct lineage-bias[1] [2] [3] clusters. These are quantitatively defined by the ratio ρ of lymphoid to myeloid cells that HSC generate upon differentiation (which makes ρ a peripheral predictor for the clonal association of a reconstituted hematopoietic system). Balanced HSCs repopulate peripheral white blood cells in the same ratio of myeloid to lymphoid cells as seen in unmanipulated mice (on average about 15% myeloid and 85% lymphoid cells, or 3≤ρ≤10). Myeloid-biased (My-bi) HSC give rise to too few lymphocytes resulting in ratios 0<ρ<3, whereas lymphoid-biased (Ly-bi) HSC generate too few myeloid cells, which results in lymphoid-to-myeloid ratios of 10 < ρ < oo. All three types are normal HSC in that they have self-renewal capacity and can regenerate all hematopietic [sic] lineages (pluripotency).

1. Muller-Sieburg [sic] CE, Cho RH, Thoman M, Adkins B, Sieburg HB, Deterministc [sic] regulation of hematopoietic stem cell self-renewal and differentiation. Blood. 2002; 100; 1302-9

2. Muller-Sieburg [sic] CE, Cho RH, Karlson [sic] L, Huang JF, Sieburg HB. Myeloid-biased hematopoietic stem cells have extensive self-renewal capacity but generate diminished progeny with impaired IL-7 responsiveness. Blood. 2004; 103:4111-8.

3. Sieburg HB, Cho RH, Dykstra B, [sic] Eaves, CJ, Muller-Sieburg [sic], CE. The hematopoietic stem cell [sic] compartment consists of a limited number of discrete stem cell subsets. Blood. 2006; 107:2311-6. Epub 2005 Nov 15.

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[18.] Pak/Fragment 013 02 - Diskussion
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Embryonic Stem (ES) cells are pluripotent stem cells. They are able to differentiate into all derivatives of the three primary germ layers: ectoderm, endoderm, and mesoderm. Embryonic Stem (ES) cells are pluripotent. This means they are able to differentiate into all derivatives of the three primary germ layers: ectoderm, endoderm, and mesoderm.
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[19.] Pak/Fragment 018 14 - Diskussion
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The action of cytokines may be autocrine, paracrine, and endocrine. Cytokines are critical to the development and functioning of both the innate and adaptive immune response, although not limited to just the immune system. They are often secreted by immune cells that have encountered a pathogen, thereby activating [and recruiting further immune cells to increase the system's response to the pathogen.] The action of cytokines may be autocrine, paracrine, and endocrine. Cytokines are critical to the development and functioning of both the innate and adaptive immune response, although not limited to just the immune system. They are often secreted by immune cells that have encountered a pathogen, thereby activating and recruiting further immune cells to increase the system's response to the pathogen.
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[20.] Pak/Fragment 019 01 - Diskussion
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[They are often secreted by immune cells that have encountered a pathogen, thereby activating] and recruiting further immune cells to increase the system's response to the pathogen. Cytokines are also involved in several developmental processes during embryogenesis. They are often secreted by immune cells that have encountered a pathogen, thereby activating and recruiting further immune cells to increase the system's response to the pathogen. Cytokines are also involved in several developmental processes during embryogenesis.
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The blood within the umbilical cord, known as cord blood, is a rich and readily available source of primitive, undifferentiated stem [cells (of type CD34+ and CD38-).] Recently, it has been discovered that the blood within the umbilical cord, known as cord blood, is a rich and readily available source of primitive, undifferentiated stem cells (i.e. CD34-positive and CD38-negative).
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[The blood within the umbilical cord, known as cord blood, is a rich and readily available source of primitive, undifferentiated stem] cells (of type CD34+ and CD38-). These cord blood cells can be used for bone marrow transplant. Recently, it has been discovered that the blood within the umbilical cord, known as cord blood, is a rich and readily available source of primitive, undifferentiated stem cells (i.e. CD34-positive and CD38-negative). These cord blood cells can be used for bone marrow transplant.
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[A multi-] potential progenitor that produces a colony containing erythroblasts and cells of at least two other recognizable lineages. Due to their primitive nature, CFU-GEMM tend to produce large colonies of >500 cells. A multi-potential progenitor that produces a colony containing erythroblasts and cells of at least two other recognizable lineages. Due to their primitive nature, CFU-GEMM tend to produce large colonies of >500 cells.
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The CFU that can reside in the spleen and form a colony are called CFU-S. The colony-forming unit–spleen (CFU–S) depends on the ability of infused bone marrow cells to give rise to clones of maturing haematopoietic cells in the spleens of irradiated mice after 8 to 12 days. It is used to measure more mature progenitor or Transit Amplifying Cells rather than stem cells.26

26.Coulombel L. Identification of hematopoietic stem/progenitor cells: strength and drawbacks of functional assays. Oncogene. 2004;23:7210-7222.

Another CFU, the colony-forming unit–spleen (CFU–S) was the basis of an in vivo clonal colony formation, which depends on the ability of infused bone marrow cells to give rise to clones of maturing hematopoietic cells in the spleens of irradiated mice after 8 to 12 days. It was used extensively in early studies, but is now considered to measure more mature progenitor or Transit Amplifying Cells rather than stem cells.
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The colony-forming unit–spleen (CFU–S) was the basis of an in vivo clonal colony formation, which depends on the ability of infused bone marrow cells to give rise to clones of maturing haematopoietic cells in the spleens of irradiated mice after 8 to 12 days. It is considered to be a measure of more mature progenitor or Transit Amplifying Cells rather than stem cells. Another CFU, the colony-forming unit–spleen (CFU–S) was the basis of an in vivo clonal colony formation, which depends on the ability of infused bone marrow cells to give rise to clones of maturing hematopoietic cells in the spleens of irradiated mice after 8 to 12 days. It was used extensively in early studies, but is now considered to measure more mature progenitor or Transit Amplifying Cells rather than stem cells.
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[Wells were rinsed once with 1 ml PBS and added to tube. 1 ml Trypsin-EDTA was added to each well and incubated for 3 to 5 minutes] and examined for detached cells. Once the adherent cells are detached, the wells are washed with more PBS and the medium collected in the appropriated tube.

The wells are finally washed with 1 ml IMDM containing 2 % FBS and transferred to the specific tube. The tubes were centrifuged at 1200 rpm for 10 minutes and the supernatant was removed without disturbing the cell pellet. Approximately 200ìL of medium was left along with the cell pellet and vortexed. To this 3ml of Methocult (H4434) methylcellulose medium was added and vortexed again. Each tube (contents of one well) was plated individually into 2 different (1.5 ml/dish) 35 mm petri dish with 1 ml syringe (without needles attached). Different dishes (6-8) were placed in a 15 cm petri-dish along with an additional 60 mm open dish containing 5 ml sterile water to maintain humidity. The dishes are incubated at 37°C in humidified incubator (>95 %) with 5% CO2 in air for 16-20 days. Colonies were scored as positive if one or more BFU-E, CFU-GM or CFU-GEMM were detected or scored as negative if no colonies were present.

[The number of LTC-IC for the test cell population was calculated by dividing the total number of CFC detected in the culture by the average number of clonogenic progenitors per LTC-IC for the standard conditions used. Alternatively the values were expressed as LTC-IC derived CFC per number of test cells.115]


[115.Hogge DE, Lansdorp PM, Reid D, Gerhard B, Eaves CJ. Enhanced detection, maintenance, and differentiation of primitive human hematopoietic cells in cultures containing murine fibroblasts engineered to produce human steel factor, interleukin-3, and granulocyte colony-stimulating factor. Blood. 1996;88:3765-3773.]

Wells were rinsed once with 0.2 ml PBS and added to tube. 0.1 ml Trypsin-EDTA was added to each well and incubated for 3 to 5 minutes and examined for detached cells. Once the adherent cells are detached, the wells are washed with more PBS and the medium collected in the appropriated tube. The wells are finally washed with 0.2 ml IMDM containing 2% FBS and transferred to the appropriate tube. The tubes were centrifuged at 1200 rpm for 10 minutes and the supernatant removed without disturbing cell pellet. Approximately 0.1 ml of medium was left along with the cell pellet and vortexed. To this 1 ml of Methocult (H4435) methylcellulose medium was added and vortexed again. Each tube (contents of one well) was plated individually into 35 mm petri dish with 1 ml syringe (without needles attached). Several dishes (6-8) were placed in a 15 cm petri-dish along with an additional 60 mm open dish containing 5 ml sterile water to maintain humidity. The dishes are incubated at 37°C in humidified incubator (>95%) with 5% CO2 in air for 12 to 16 days. Colonies were counted and a well scored as positive if one or more BFU-E, CFU-GM or CFU-GEMM were detected or scored as negative if no colonies were present. [The LTC-IC frequency in the test cell population was calculated from the proportion of negative wells (no CFC present) and the method of maximum likelihood. Statistical analysis was performed using L-Calc™ software for limiting dilution analyses.]
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[Many of the signaling pathways activated by SCF, CSF] 1 and flat-3 ligand (FL) receptors, including the Ras/Raf-mitogen activated protein kinase cascade, the janus kinase (JAK)/ signal transducers and activators of transcription (STATs) pathway, Src family members and phosphatidylinositol-3-kinase (PI3K), are shared.

All three receptors are likely to exhibit ligand induced, Cbl-mediated decreases in receptor expression. The SCF and CSF-1 receptors are encoded by the proto-oncogenes c-kit and c-fms, respectively. The oncogenes derived from these two proto-oncogenes are present in mutated forms in retroviruses that cause sarcoma in cats. The mutations in the receptor genes cause constitutive activation of the receptors in the absence of cytokines, thus contributing to unregulated cell proliferation.

1.3.4. The role of transcription factors

Nuclear transcription factors are essential for stem cell lineage commitment.

[Page 3]

The Role of Transcription Factors

Experiments using gene manipulation have shown that nuclear transcription factors are essential for stem cell lineage commitment.

[Page 4]

Many of the signalling pathways activated by SCF, CSF-1 and FL receptors, including the Ras/Raf-mitogen activated protein kinase cascade, the Janus kinase (JAK) signal transducers and activation of transcription (STAT) pathway, Src family members and phosphatidylinositol-3-kinase (PI3K), are shared.43 All three receptors are likely to exhibit ligand induced, Cbl-mediated decreases in receptor expression.44

The SCF and CSF-1 receptors are encoded by the proto-oncogenes c-kit and c-fms respectively.45 The oncogenes derived from these two proto-oncogenes are present in mutated forms in retroviruses that cause sarcoma in cats. The mutations in the receptor genes cause constitutive activation of the receptors in the absence of cytokines.45 thus contributing to unregulated cell proliferation.


43. Linnekin D. Early signalling pathways activated by c-Kit in haemopoietic cells. Int J Biochem Cel Biol 1999; 31:1053-1074.

44. Lee PS, Wang Y, Dominguez MG et al. The Cbl protooncoprotein stimulates CSF-1 receptor multiubiquitination and endocytosis, and attenuates macrophage proliferation. EMBO J 1999; 18:3616-3628.

45. Sherr CJ. Colony Stimulating Factor 1 Receptor. Blood 1990; 75:1-1.

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[One of the signal transduction] pathways that are activated is called the mitogen-activated protein kinase (MAPK) pathway.

The signal transduction component labelled as "MAPK" in the pathway was originally called "ERK," so the pathway is called the MAPK/ERK pathway. The MAPK protein is an enzyme, a protein kinase that can attach phosphate to target proteins such as the transcription factor MYC and, thus, alter gene transcription and, ultimately, cell cycle progression. Many cellular proteins are activated downstream of the growth factor receptors (such as EGFR) that initiate this signal transduction pathway (Fig.1.3.4b). Some signaling transduction pathways respond differently depending on the amount of signaling received by the cell. For instance, the hedgehog protein activates different genes, depending on the amount of hedgehog protein present. Complex multi-component signal transduction pathways provide opportunities for feedback, signal amplification, and interactions inside one cell between multiple signals and signaling pathways.

For example, one of the signal transduction pathways that is activated is called the mitogen-activated protein kinase (MAPK) pathway. The signal transduction component labeled as "MAPK" in the pathway was originally called "ERK" so the pathway is called the MAPK/ERK pathway. The MAPK protein is an enzyme, a protein kinase that can attach phosphate to target proteins such as the transcription factor MYC and thus alter gene transcription and, ultimately, cell cycle progression. Many cellular proteins are activated downstream of the growth factor receptors (such as EGFR) that initiate this signal transduction pathway.

Some signaling transduction pathways respond differently depending on the amount of signaling received by the cell. For instance the hedgehog protein activates different genes depending on the amount of hedgehog protein present.

Complex multi-component signal transduction pathways provide opportunities for feedback, signal amplification, and interactions inside one cell between multiple signals and signaling pathways.

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Many growth factors (CSF, GCSF, GMCSF, FLT3, TPO,IL-6, IL-3, IL-11 etc.) bind to receptors at the cell surface and stimulate cells to progress through the cell cycle and divide. Several of these receptors are kinases that start to phosphorylate themselves and other proteins when binding to a ligand. This phosphorylation can generate a binding site for a different protein and thus induce protein-protein interaction. Many growth factors bind to receptors at the cell surface and stimulate cells to progress through the cell cycle and divide. Several of these receptors are kinases that start to phosphorylate themselves and other proteins when binding to a ligand. This phosphorylation can generate a binding site for a different protein and thus induce protein-protein interaction.
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[30.] Pak/Fragment 020 14 - Diskussion
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HSCs express a great diversity of transcripts, including a wide range of genes originally believed to be restricted to more mature and lineage-committed cell types. The single-cell polymerase chain reaction strategies had suggested that such loose transcription of lineage-associated transcripts was necessary to prime primitive progenitor cells for differentiation toward downstream fates.63

An alternative interpretation is that stem cells possess global transcriptional accessibility and that it is the step-wise restriction of locus accessibility that underlies lineage specification.64,65 Cell fate can also be switched from one committed cell type to another on over expression of IL2R or GM-CSF receptors 17,18 GATA-166 or C/EBP /ß [sic] .67 An additional mechanism of lineage specification has been revealed which attributes to the ablation of the transcription factors Pax5 or GATA-1 in lineage-restricted progenitors. This was sufficient to despecify their B-cell and erythroid fate, respectively, and allow a multilineage developmental potential.


17. Kondo M, Scherer DC, Miyamoto T, et al. Cell-fate conversion of lymphoid-committed progenitors by instructive actions of cytokines. Nature. 2000;407:383-386.

18. Kondo M, Wagers AJ, Manz MG, et al. Biology of hematopoietic stem cells and progenitors: implications for clinical application. Annu Rev Immunol. 2003;21:759-806.

63. Mikkola HK, Fujiwara Y, Schlaeger TM, Traver D, Orkin SH. Expression of CD41 marks the initiation of definitive hematopoiesis in the mouse embryo. Blood. 2003;101:508-516.

64. Akashi K, Reya T, Dalma-Weiszhausz D, Weissman IL. Lymphoid precursors. Curr Opin Immunol. 2000;12:144-150.

65. Akashi K, He X, Chen J, et al. Transcriptional accessibility for genes of multiple tissues and hematopoietic lineages is hierarchically controlled during early hematopoiesis. Blood. 2003;101:383- 389.

66. Iwasaki H, Mizuno S, Wells RA, Cantor AB, Watanabe S, Akashi K. GATA-1 converts lymphoid and myelomonocytic progenitors into the megakaryocyte/erythrocyte lineages. Immunity. 2003;19:451-462.

67. Xie H, Ye M, Feng R, Graf T. Stepwise reprogramming of B cells into macrophages. Cell. 2004;117:663-676.

One surprising finding of these studies is the discovery that HSCs express a great diversity of transcripts, including a wide range of genes originally believed to be restricted to more mature and lineage-committed cell types. The findings of these genome-wide expression studies had in fact been foreshadowed by earlier studies using single-cell polymerase chain reaction strategies that had suggested that such promiscuous transcription of lineage-associated transcripts was necessary to prime primitive progenitor cells for differentiation toward downstream fates.39,40 An alternative interpretation posits that stem cells possess global transcriptional accessibility and that it is the step-wise restriction of locus accessibility that underlies lineage specification.37 [...] This has been exemplified in experiments in which cell fate was switched from one committed cell type to another on overexpression of IL2R or GM-CSF receptors,41 GATA-1,42 or C/EBPα/β.43 An additional mechanism of lineage specification was revealed in experiments in which ablation of the transcription factors Pax5 or GATA-1 in lineage-restricted progenitors was found to be sufficient to despecify their B-cell and erythroid fate, respectively, and allow a multilineage developmental potential.44,45

37. Akashi K, He X, Chen J, Iwasaki H, Niu C, Steenhard B, Zhang J, Haug J, Li L: Transcriptional accessibility for genes of multiple tissues and hematopoietic lineages is hierarchically controlled during early hematopoiesis. Blood 2003, 101:383–389

39. Miyamoto T, Iwasaki H, Reizis B, Ye M, Graf T, Weissman IL, Akashi K: Myeloid or lymphoid promiscuity as a critical step in hematopoietic lineage commitment. Dev Cell 2002, 3:137–147

40. Hu M, Krause D, Greaves M, Sharkis S, Dexter M, Heyworth C, Enver T: Multilineage gene expression precedes commitment in the hemopoietic system. Genes Dev 1997, 11:774–785

41. Kondo M, Scherer DC, Miyamoto T, King AG, Akashi K, Sugamura K, Weissman IL: Cell-fate conversion of lymphoid-committed progenitors by instructive actions of cytokines. Nature 2000, 407:383–386

42. Iwasaki H, Mizuno S, Wells RA, Cantor AB, Watanabe S, Akashi K: GATA-1 converts lymphoid and myelomonocytic progenitors into the megakaryocyte/erythrocyte lineages. Immunity 2003, 19:451–462

43. Xie H, Ye M, Feng R, Graf T: Stepwise reprogramming of B cells into macrophages. Cell 2004, 117:663–676

44. Mikkola I, Heavey B, Horcher M, Busslinger M: Reversion of B cell commitment upon loss of Pax5 expression. Science 2002, 297:110–113

45. Kitajima K, Zheng J, Yen H, Sugiyama D, Nakano T: Multipotential differentiation ability of GATA-1-null erythroid-committed cells. Genes Dev 2006, 20:654–659

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Figure 1.2.2: A model of the haematopoietic developmental hierarchy. Self-renewing HSCs reside at the top of the hierarchy, thereby giving rise to a number of multipotent progenitors. Multipotent progenitors give rise to oligo-potent progenitors including the CLP, which gives rise to mature B lymphocytes, T lymphocytes, and natural killer (NK) cells. The common myeloid progenitor (CMP) gives rise to granulocyte-macrophage progenitors, which differentiate into monocytes/macrophages and granulocytes, and megakaryocyte/erythrocyte progenitors, which differentiate into megakaryocytes/platelets and erythrocytes. The cell surface phenotype of many of these cell types is shown for the murine and human systems 50(Figure adapted from Bryder D et al., 2006).


50. Bryder D, Rossi DJ, Weissman IL. Hematopoietic stem cells: the paradigmatic tissue-specific stem cell. Am J Pathol. 2006;169:338-346.

15a source Pak.png

Figure 1. Model of the hematopoietic developmental hierarchy. Self-renewing HSCs reside at the top of the hierarchy, giving rise to a number of multipotent progenitors. Multipotent progenitors give rise to oligo-potent progenitors including the CLP, which gives rise to mature B lymphocytes, T lymphocytes, and natural killer (NK) cells. The common myeloid progenitor (CMP) gives rise to granulocyte-macrophage progenitors, which differentiate into monocytes/macrophages and granulocytes, and megakaryocyte/erythrocyte progenitors, which differentiate into megakaryocytes/platelets and erythrocytes. Both CMPs and CLPs have been proposed to give rise to dendritic cells. The cell surface phenotype of many of these cell types is shown for the murine and human systems.

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[As these immature cells develop, they lose receptors for] some cytokines e.g. SCF and IL-3, while retaining receptors for later acting cytokines such as CSF-1.

Eventually immature cells reach the stage of the committed progenitor cell, where their proliferation and differentiation are along one particular lineage, guided by the lineage-restricted cytokines. Synergy occurs between some late-acting lineages restricted cytokines such as CSF-1, EPO and G-CSF, with early acting cytokines such as SCF and IL-3, in stimulating the proliferation and differentiation of multi-potent cells. This provides a mechanism by which the tightly regulated changes in the level of a late acting cytokine can be coupled to the channelling of multi-potent progenitor cells into a lineage to satisfy the demand for differentiated cells. The underlying mechanism of synergy may lie at the level of either the receptors or at the level of post receptor signaling pathways (Molecular cell biology. Lodish, Harvey F. 5. ed. )

[Page 2]

As these immature cells develop, they lose receptors for some cytokines e.g., SCF and IL-3, while retaining receptors for later acting cytokines such as CSF-1. Eventually at least some of the immature cells reach the stage of committed progenitor cell, where their further proliferation and differentiation are along one particular lineage, dictated by the relevant lineage-restricted cytokine.

[Page 3]

Synergy occurs between some late-acting lineage restricted cytokines such as CSF-1, EPO and G-CSF, with early acting cytokines such as SCF and IL-3, in stimulating the proliferation and differentiation of multi-potent cells. This provides a mechanism by which the tightly regulated changes in the level of a late acting cytokine, can be coupled to the channelling of multi-potent progenitor cells into a lineage to satisfy the demand for differentiated cells. The underlying mechanism of synergy may lie at the level of either the receptors or at the level of post receptor signalling pathways.

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There is an enormous proliferative and developmental capacity of the more committed multipotent, oligo-potent and lineage-restricted progenitor cells within the haematopoietic hierarchy. A significant degree of homeostatic control of mature blood cells is also mediated at the level of these progenitors.25

25.Dzierzak E, Speck NA. Of lineage and legacy: the development of mammalian hematopoietic stem cells. Nat Immunol. 2008;9:129-136.

However, because of the enormous proliferative and developmental capacity of the more committed multipotent, oligo-potent, and lineage-restricted progenitor cells within the hematopoietic hierarchy, a significant degree of homeostatic control of mature blood cells is also mediated at the level of these progenitors.
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Clonogenic haematopoietic assays detect progenitor cells committed to a specific lineage by seeding candidate populations into semi-solid methylcellulose media. Progenitors identified by this assay are retrospectively classified as colony forming cells (CFC) and can be quantitatively subdivided into lineage restricted subtypes by examining the composition of the resulting progeny (methods, p-51)(Fig. 4.5.1a). 4.4 Constitutive expression of ABCG2 increases the production of CFC in vitro

Clonogenic haematopoietic assays detect progenitor cells committed to a specific lineage by seeding candidate populations into semi-solid methylcellulose media. Progenitors identified by this assay are retrospectively classified as colony forming cells (CFC) and can be quantitatively subdivided into lineage restricted subtypes by examining the composition of the resulting progeny (described in the methods section).

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3.2.6. Purification of umbilical cord blood CD34+ cells (CB CD34+) from mononuclear umbilical cord blood (UCB).

Umbilical cord blood was collected in heparinised syringes according to institutional guidelines following normal full-term deliveries. Informed consent was obtained in all cases. Mononuclear cells (MNC) were separated using density gradient centrifugation. Fresh umbilical cord blood, not older than 12 hours, was diluted with 2 volumes of PBS and layered over Pancoll. Usually 35 ml of diluted blood was layered over 15 ml Pancoll in a 50 ml conical tube. This was centrifuged at 400x g for 30 minutes at 20°C in a swinging-bucket rotor without brakes. The upper layer was aspirated and discarded, leaving the interphase undisturbed. The interphase containing MNC such as lymphocytes, monocytes and thrombocytes was then transferred to a new 50 ml tube, washed twice with large volumes of PBS, and then counted before labelling with magnetic bead or fluorochrome conjugated antibodies.

hCB CD34+ cell purification was conducted using MACS CD34+ Cell Isolation Kit that uses positive selection method. Cells were resuspended in a volume of 300 ml per 1x108 MNCs These were blocked with 100 ml of FcR Blocking Reagent and labelled with 100 ml of CD34 Microbeads. When working with higher cell number, all the reagent volumes & the total volume was scaled up accordingly. This was followed by incubation for 30 minutes at 4-8°C. Cells were then washed twice by adding 10x the labelling volume of buffer and centrifuged at 300 x g for 15 minutes. The resultant cell pellet was then resuspended in 500 ml of MACS buffer and loaded into MS Column mounted on magnetic separator. The negative cells were allowed to pass through and the column was washed at least three times with 2 ml buffer.

3.2.6 Purification of human UCB CD133+ and CD34+ cells

Density Gradient Centrifugation:

Umbilical cord blood was collected in heparinised syringes according to institutional guidelines following normal full-term deliveries. Informed consent was obtained in all cases. Mononuclear cells (MNC) were separated using density gradient centrifugation. Fresh umbilical cord blood, not older than 12 hours, was diluted with 2 volumes of PBS and layered over Pancoll. Usually 35 ml of diluted blood was layered over 15 ml Pancoll in a 50 ml conical tube. This was centrifuged at 400x g for 30 minutes at 20°C in a swinging-bucket rotor without brakes. The upper layer was aspirated and discarded, leaving the interphase undisturbed. The interphase containing MNC such as lymphocytes, monocytes and thrombocytes was then transferred to a new 50 ml tube, washed twice with large volumes of PBS, and then counted before labelling with magnetic bead or fluorochrome conjugated antibodies.

Magnetic Separation:

CD133+ cell purification was conducted using MACS CD133 Cell Isolation Kit that uses positive selection method. Cells were resuspended in a volume of 300 μl per 108 cells, blocked with 100 μl of FcR Blocking Reagent and labelled with 100 μl of

[page 41]

Materials & Methods

CD133 Microbeads. When working with higher cell number, all the reagent volumes & the total volume was scaled up accordingly. This was followed by incubation for 30 minutes at 4-8°C. Cells were then washed twice by adding 10x the labelling volume of buffer and centrifuged at 300 x g for 15 minutes. The resultant cell pellet was then resuspended in 500 μl of MACS buffer and loaded into MS Column mounted on magnetic separator. The negative cells were allowed to pass through and the column was washed at least three times with 2 ml buffer.

Anmerkungen

The source is not given.

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(SleepyHollow02) Schumann

[36.] Pak/Fragment 049 01 - Diskussion
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[The column was then removed from the separator, placed on a collection tube, loaded with fresh buffer, and the magnetically labelled cells flushed out using the] plunger. The magnetic separation was usually repeated to get a purity of more than 95%. Purified cells were then frozen in FBS with 10% DMSO and thawed when needed for pre-stimulation and transduction. hCB CD34+ cell enrichment was done by FACS. For separation by FACS, MNCs were thawed from frozen stocks or prepared freshly from UCB and labelled using anti CD34-PE antibody (100 ml per 108 cells), for 30 minutes on ice. Labelled cells were then washed twice with PBS, resuspended in FACS buffer and sorted. The sorted cells with purity above 95 % were used for 48 hour pre-stimulation followed by transduction. The column was then removed from the separator, placed on a collection tube, loaded with fresh buffer, and the magnetically labelled cells flushed out using the plunger. The magnetic separation was usually repeated to get a purity of more than 95%. Purified cells were then frozen in FBS with 10% DMSO and thawed when needed for pre-stimulation and transduction.

FACS sorting:

CD34+ cell enrichment was done either by MACS as done for CD133+ cells or by FACS. For separation by FACS, MNCs were thawed from frozen stocks or prepared freshly from UCB and labelled using anti CD34-PE antibody (100 μL per 108 cells), for 30 minutes on ice. Labelled cells were then washed twice with PBS, resuspended in FACS buffer and sorted. The sorted cells with purity above 95% were used for 48 hour pre-stimulation followed by transduction.

Anmerkungen

The source is not given.

Sichter
(SleepyHollow02) Schumann

[37.] Pak/Fragment 017 09 - Diskussion
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Cytokines exert their action through high-affinity receptors on the cell surface that are linked to pathways of cellular activation, commitment, differentiation, survival, proliferation and differentiation.

Cross-linking of receptor subunits on the outside of the cell wall leads to adjoining of kinases associated with the intracellular receptor tails, either as intrinsic activities, or because of pre-association of secondary kinase molecules. This intracellular association of signaling molecules results in phosphorylation of tyrosine residues in the receptor tail and binding of further signaling molecules that have phospho-tyrosine-binding domains. Several aspects of the downstream intracellular pathways of cytokines are similar, because different activated receptor cytoplasmic domains often bind a common signaling molecule or family of signaling molecules. Thus stoichiometry and rate of reaction are important regulatory influences that allow discrimination between signaling processes with different outcomes (Molecular cell biology. Lodish, Harvey F. 5. ed. )

1.3.2. Cooperative Interaction of Cytokines

For efficient in vitro proliferation and differentiation, pluripotent and multipotent progenitor cells require a combination of cytokines [e.g. SCF, IL-1, IL-3, IL-6, GM-CSF, and colony stimulating factor-1, (CSF-1)]. Immature haematopoietic cells have been shown to co-express a number of different lineage specific receptors at low levels. As these immature cells develop, they lose receptors for [some cytokines e.g. SCF and IL-3, while retaining receptors for later acting cytokines such as CSF-1.]

Cytokines exert their action through high-affinity receptors on the cell surface that are linked to pathways of cellular activation, survival, proliferation and differentiation. Cross-linking of receptor subunits on the outside of the cell wall leads to abutting of kinases associated with the intracellular receptor tails, either as intrinsic activities, or because of pre-association of secondary kinase molecules. This intracellular association of signalling molecules results in phosphorylation of tyrosine residues in the receptor tail and binding of further signalling molecules that have phospho-tyrosine-binding domains. Several aspects of the downstream intracellular pathways of cytokines are similar, because different activated receptor cytoplasmic domains often bind a common signalling molecule or family of signalling molecules. Thus stoichiometry and rate of reaction are important regulatory influences that allow discrimination between signalling processes with different outcomes.

Cooperative Interaction of Cytokines in Proliferation and Differentiation

For efficient in vitro proliferation and differentiation, PPSCs and multi-potent progenitor cells require a combination of cytokines. (e.g., SCF, IL-1, IL-3, IL-6, GM-CSF, and CSF-1).23,24 As might be expected from this observation, immature haematopoietic cells have been shown to co-express a number of different lineage specific receptors at low levels.25 As these immature cells develop, they lose receptors for some cytokines e.g., SCF and IL-3, while retaining receptors for later acting cytokines such as CSF-1.


23. Bradley TR, Hodgson GS. Detection of primitive macrophage progenitor cells in mouse bone marrow. Blood 1979; 54:1446–1450.

24. Stanley ER, Bartocci A, Patinkin D et al. Regulation of very primitive multipotent haematopoietic cells by Hemopoietin–1. Cell 1986; 45:667–674.

25. Cross MA, Enver T. The lineage commitment of haematopoietic progenitor cells. Curr Opin Genet Devel 1997; 7:609-613.

Anmerkungen

Identical, without any part of it marked as a citation.

Pak names a general source, but this passage is not to be found there.

Sichter
(Graf Isolan) Singulus

[38.] Pak/Fragment 033 01 - Diskussion
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[NOD/SCID mice were bred from the breeding] pairs originally obtained from Taconic Bomholt, Denmark and maintained in the animal facility located at the GSF-Haematology, Munich. All animals were handled under sterile conditions and maintained under micro isolators. [xenografts [sic] into immunodeficient NOD-SCID mice are the gold standard readout for human long-term repopulating hematopoietic cells.110-112]
3.1.5. Mice related reagents and equipment

Sterile Syringes: BD Plastipak 1 ml syringe (BD Biosciences, Palo Alto, CA) for injection of cells in mice and Kendall Monoject 3 ml syringes (Tyco Healthcare, UK) for bone marrow flushing and plating of CFC.

Sterile needles: 0.5 x 25 mm for intra venous injection of cells in mice and 0.55 x 25 mm (BD Microlance, Drogheda, Ireland) for bone marrow aspiration from living mice and flushing of bone marrow from extracted bones. 16 X 1.5 inch needles for dispensing and plating Methocult (CFC) media (Stem Cell Technologies, Vancouver, Canada).

Ammonium Chloride solution: For erythrocyte lysis 0.8% NH4Cl with 0.1 mM EDTA (Stem Cell Technologies, Vancouver, Canada).

Heparinized capillaries: (Microvette CB 300) plastic capillaries for collection of blood, containing 15 units (I.E) Lithium heparin per ml of blood (Sarstedt, Numbrecht, Germany).

3.1.6. Bacterial strain

• E. coli DH5á [sic]

3.1.5 Bacterial strain

• E. coli DH5α

[...]

NOD/SCID mice were bred from the breeding pairs originally obtained from Taconic Europe and maintained in the animal facility located at the GSF-Haematology, Munich. All animals were handled under sterile conditions and maintained under micro isolators.

3.1.7 Mice related reagents and equipment:

[...]

Sterile Syringes: BD Plastipak 1 ml syringe (BD Biosciences, Palo Alto, CA) for injection of cells in mice and Kendall Monoject 3 ml syringes (Tyco Healthcare, UK) for bone marrow flushing and plating of CFC.

Sterile needles: 0.5 x 25 mm for intra venous injection of cells in mice and 0.55 x 25 mm (BD Microlance, Drogheda, Ireland) for bone marrow aspiration from living

[page 34]

mice and flushing of bone marrow from extracted bones. 16 X 1.5 inch needles for dispensing and plating Methocult (CFC) media (Stem Cell Technologies, Vancouver, Canada)

Ammonium Chloride solution: For erythrocyte lysis 0.8% NH4Cl with 0.1 mM EDTA (Stem Cell Technologies, Vancouver, Canada)

Heparinized capillaries: (Microvette CB 300) plastic capillaries for collection of blood, containing 15 units (I.E) Lithium heparin per ml of blood (Sarstedt, Numbrecht, Germany).

Anmerkungen

The source is not given.

Sichter
(SleepyHollow02) Schumann

[39.] Pak/Fragment 032 01 - Diskussion
Bearbeitet: 6. April 2014, 06:49 Hindemith
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[Introduction of retroviral vectors into PG13] cells results in the production of retrovirus virions capable of infecting cells from many species excluding mice.

K562: An erythroleukemia cell line derived from a chronic myeloid leukaemia patient in blast crisis. K562 cells were purchased from ATCC.

M2-10B4 j-GCSF-tkneo j-IL-3-hytk: Murine M2-10B4 fibroblasts engineered to produce high levels of both human granulocyte colony-stimulating factor (GCSF) and interleukin-3 (IL-3; 190 and 4 ng/mll, respectively), referred henceforth as M2-10B4 G-CSF / IL-3, were provided courteously provided by Connie Eaves (Terry Fox Laboratory, Vancouver, Canada).

Sl/Sl j-SF-tkneo j-IL-3-hytk: SI/SI fibroblasts engineered to produce high levels of soluble Steel factor (SF), with or without production of the transmembrane form of SF (60 and 4 ng/ ml, respectively), referred henceforth as Sl/Sl SF / IL-3, were provided kindly by Connie Eaves (Terry Fox Laboratory, Vancouver, Canada).

[• MS-5: Mouse stromal cells established by irradiation of the adherent cells in long-term bone marrow cultures derived from C3H/HeNSlc strain mice.]

3.1.4. The NOD-SCID Mice

The NOD/LtSz-scid strain was generated by crossing the SCID mutation from C.B-17- SCID mice onto the NOD background. C.B-17-scid mice lack functional T & B lymphocytes. The NOD strain mouse is an animal model of spontaneous autoimmune T-cell mediated insulin dependent diabetes mellitus (IDDM); however they have multiple defects in innate immunity. They are deficient in NK cell activity; display defects in myeloid development and function, and cannot generate either the classical or alternative pathways of haemolytic complement activation.

The NOD/LtSz-scid lacks an adaptive immune system; due to the absence of T cells, they do not develop autoimmune IDDM and remain insulitis- and diabetes free throughout life. However they carry the innate immune defects present in the parental NOD/Lt stock of mice.109


109.Lowry PA, Shultz LD, Greiner DL, et al. Improved engraftment of human cord blood stem cells in NOD/LtSz-scid/scid mice after irradiation or multiple-day injections into unirradiated recipients. Biol Blood Marrow Transplant. 1996;2:15-23.

[page 32]

Introduction of retroviral vectors into PG13 cells results in the production of retrovirus virions capable of infecting cells from many species excluding mice.

K562: An erythroleukemia cell line derived from a chronic myeloid leukemia patient in blast crisis. K562 cells were purchased from ATCC.

M2-10B4 j-GCSF-tkneo j-IL-3-hytk: Murine M2-10B4 fibroblasts engineered to produce high levels of both human granulocyte colony-stimulating factor (G-CSF) and interleukin-3 (IL-3; 190 and 4 ng/mll, respectively), referred henceforth as M2-10B4 G-CSF / IL-3, were provided courteously provided by Connie Eaves (Terry Fox Laboratory, Vancouver, Canada).

Sl/Sl j-SF-tkneo j-IL-3-hytk: SI/SI fibroblasts engineered to produce high levels of soluble Steel factor (SF), with or without production of the transmembrane form of SF (60 and 4 ng/ ml, respectively), referred henceforth as Sl/Sl SF / IL-3, were provided kindly by Connie Eaves (Terry Fox Laboratory, Vancouver, Canada).

[page 33]

3.1.6 The NOD-SCID Mice:

The NOD/LtSz-scid strain was generated by crossing the SCID mutation from C.B-17-scid mice onto the NOD background. C.B-17-scid mice lack functional T & B lymphocytes. The NOD strain mouse is an animal model of spontaneous autoimmune T-cell mediated insulin dependent diabetes mellitus (IDDM); however they have multiple defects in innate immunity. They are deficient in NK cell activity; display defects in myeloid development and function, and cannot generate either the classical or alternative pathways of haemolytic complement activation. The NOD/LtSz-scid lacks an adaptive immune system; due to the absence of T cells, they do not develop autoimmune IDDM and remain insulitis- and diabetes free throughout life. However they carry the innate immune defects present in the parental NOD/Lt stock of mice (Lowry et al., 1996).


Lowry,P.A., Shultz,L.D., Greiner,D.L., Hesselton,R.M., Kittler,E.L., Tiarks,C.Y., Rao,S.S., Reilly,J., Leif,J.H., Ramshaw,H., Stewart,F.M., and Quesenberry,P.J. (1996). Improved engraftment of human cord blood stem cells in NOD/LtSz-scid/scid mice after irradiation or multiple-day injections into unirradiated recipients. Biol. Blood Marrow Transplant. 2, 15-23.

Anmerkungen

The source is not given.

Sichter
(SleepyHollow02) Schumann

[40.] Pak/Fragment 022 01 - Diskussion
Bearbeitet: 6. April 2014, 06:49 Hindemith
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Some other factors appear to be more lineage-specific in action such as GATA-3, Ikaros, PU.1, GATA-1, CBP, Atf4, c-myb, and E2A, and their absence affects specific haematopoietic lineages. 73-77 The genetic factors involved in regulating fetal liver HSC are: Meis1, which is highly expressed in fetal liver Sca-1 +Lin- cells that are enriched for HSC activity78 and Hoxb4, which causes in vivo and ex vivo expansion of HSC when constitutively expressed. 79,80 Hox proteins interact with another transcription factor Pbx1, which itself interacts with Meis1 and forms a trimeric nuclear complex which is involved in target gene regulation81,82 (Fig.1.3.4a).

HSC self- renewal maintenance in adult BM is regulated by a different set of genes. A number of recent studies point out to nuclear factors such as the Polycomb (PcG) genes Bmi-1 and Rae-28, GATA-2 and TEL for potentially regulating this process. It has been observed that Bmi-1 levels decline during haematopoietic development, and that Bmi-1 deficient mice develop hypocellular BM and die at less than 2 months of age. This led to the speculation that Bmi-1 is involved in maintenance of the HSC pool.83 Rae-28, a known nuclear partner of Bmi-1, also plays a crucial role in maintaining the activity of HSCs during fetal haematopoiesis. 84

The zinc-finger transcription factor GATA-2, a member of GATA family, plays a critical role in maintaining the pool of multipotent progenitors and HSCs, both during embryogenesis and in the adult.85 The zinc-finger transcription factors GATA-1 and its transcriptional cofactor called Friend of GATA-1 (FOG-1), have been found to be essential for erythroid and megakaryocytic differentiation.86,87 PU.1 is a member of the Ets family of transcription factors and is essential in the development of cells of the monocytic, granulocytic and lymphoid lineages.88


73.Bain G, Maandag EC, Izon DJ, et al. E2A proteins are required for proper B cell development and initiation of immunoglobulin gene rearrangements. Cell. 1994;79:885-892.

74.Wang H, Xie Z, Scott RE. JunD phosphorylation, and expression of AP-1 DNA binding activity modulated by serum growth factors in quiescent murine 3T3T cells. Oncogene. 1996;13:2639-2647.

75.Scott G, Ewing J, Ryan D, Abboud C. Stem cell factor regulates human melanocyte-matrix interactions. Pigment Cell Res. 1994;7:44-51.

76.Emambokus N, Vegiopoulos A, Harman B, Jenkinson E, Anderson G, Frampton J. Progression through key stages of haemopoiesis is dependent on distinct threshold levels of c-Myb. EMBO J. 2003;22:4478-4488.

77.Lessard J, Faubert A, Sauvageau G. Genetic programs regulating HSC specification, maintenance and expansion. Oncogene. 2004;23:7199-7209.

78.Pineault N, Helgason CD, Lawrence HJ, Humphries RK. Differential expression of Hox, Meis1, and Pbx1 genes in primitive cells throughout murine hematopoietic ontogeny. Exp Hematol. 2002;30:49-57.

79.Antonchuk J, Sauvageau G, Humphries RK. HOXB4-induced expansion of adult hematopoietic stem cells ex vivo. Cell. 2002;109:39-45.

80.Buske C, Feuring-Buske M, Antonchuk J, et al. Overexpression of HOXA10 perturbs human lymphomyelopoiesis in vitro and in vivo. Blood. 2001;97:2286-2292.

81.Liu JP, Laufer E, Jessell TM. Assigning the positional identity of spinal motor neurons: rostrocaudal patterning of Hox-c expression by FGFs, Gdf11, and retinoids. Neuron. 2001;32:997-1012.

82. Mann RS, Affolter M. Hox proteins meet more partners. Curr Opin Genet Dev. 1998;8:423-429.

83.Lessard J, Schumacher A, Thorsteinsdottir U, van Lohuizen M, Magnuson T, Sauvageau G. Functional antagonism of the Polycomb-Group genes eed and Bmi1 in hemopoietic cell proliferation. Genes Dev. 1999;13:2691-2703.

84.Ohta H, Sawada A, Kim JY, et al. Polycomb group gene rae28 is required for sustaining activity of hematopoietic stem cells. J Exp Med. 2002;195:759-770.

85.Tsai FY, Orkin SH. Transcription factor GATA-2 is required for proliferation/survival of early hematopoietic cells and mast cell formation, but not for erythroid and myeloid terminal differentiation. Blood. 1997;89:3636-3643.

86.Pevny L, Simon MC, Robertson E, et al. Erythroid differentiation in chimaeric mice blocked by a targeted mutation in the gene for transcription factor GATA-1. Nature. 1991;349:257-260.

87.Vyas P, McDevitt MA, Cantor AB, Katz SG, Fujiwara Y, Orkin SH. Different sequence requirements for expression in erythroid and megakaryocytic cells within a regulatory element upstream of the GATA- 1 gene. Development. 1999;126:2799-2811.

88.Scott EW, Simon MC, Anastasi J, Singh H. Requirement of transcription factor PU.1 in the development of multiple hematopoietic lineages. Science. 1994;265:1573-1577.

Some other factors appear to be more lineage-specific in action such as GATA-3, Ikaros, PU.1, GATA-1, CBP, Atf4, c-myb, and E2A, and their absence affects specific haematopoietic lineages (Bain et al 1994, Scott 1994, Wang et al 1996 Mauoka 2002, Wang 1997, Emambokus 2003). [...] The genetic factors involved in regulating fetal liver HSC are: Meis1, which is highly expressed in fetal liver Sca-1 + Lin- cells that are enriched for HSC activity (Pineault et al., 2002) and Hoxb4, the overexpression of which causes in vivo and ex vivo expansion of HSC (Antonchuk et al., 2001; Buske et al., 2002). Hox proteins interact with another transcription factor Pbx which itself interacts with Meis1 and forms a trimeric nuclear complex which is involved in target gene regulation (Liu et al., 2001; Swift et al., 1998). HSC self-renewal maintenance in adult BM is regulated by a different set of genes. A number of recent studies point out to nuclear factors such as the Polycomb-Group (PcG) genes Bmi-1 and Rae-28, GATA-2 and TEL for potentially regulating this process. It has been observed that Bmi-1 levels decline during haematopoietic development, and that Bmi-1 deficient mice develop hypocellular BM and die at less than 2 months of age. This led to the speculation that Bmi-1 is involved in maintenance of the HSC pool (Lessard et al., 2004). Rae-28, a known nuclear partner of Bmi-1 also plays a crucial role in maintaining the activity of HSC during fetal haematopoiesis (Ohta et al., 2002). The zinc-finger transcription factor GATA-2, a member of GATA family, plays a critical role in

[page 8]

maintaining the pool of multipotent progenitors and HSC, both during embryogenesis and in the adult (Tsai et al., 1994).

[...] The zinc-finger transcription factor GATA-1 and its transcriptional cofactor called Friend of GATA-1 (FOG-1), have been found to be essential for erythroid and megakaryocytic differentiation (Pevny et al., 1991; Tsang et al., 1998; Vyas et al., 1999). PU.1 is a member of the Ets family of transcription factors and is essential in the development of cells of the monocytic, granulocytic and lymphoid lineages (Scott et al., 1994).


Antonchuk,J., Sauvageau,G., and Humphries,R.K. (2001). HOXB4 overexpression mediates very rapid stem cell regeneration and competitive haematopoietic repopulation. Exp. Hematol. 29, 1125-1134.

Buske,C., Feuring-Buske,M., Abramovich,C., Spiekermann,K., Eaves,C.J., Coulombel,L., Sauvageau,G., Hogge,D.E., and Humphries,R.K. (2002). Deregulated expression of HOXB4 enhances the primitive growth activity of human hematopoietic cells. Blood 100, 862-868.

Lessard,J., Faubert,A., and Sauvageau,G. (2004). Genetic programs regulating HSC specification, maintenance and expansion. Oncogene 23, 7199-7209.

Liu,Y., MacDonald,R.J., and Swift,G.H. (2001). DNA binding and transcriptional activation by a PDX1.PBX1b.MEIS2b trimer and cooperation with a pancreas-specific basic helix-loop-helix complex. J Biol. Chem. 276, 17985-17993.

Ohta,H., Sawada,A., Kim,J.Y., Tokimasa,S., Nishiguchi,S., Humphries,R.K., Hara,J., and Takihara,Y. (2002). Polycomb group gene rae28 is required for sustaining activity of hematopoietic stem cells. J Exp. Med. 195, 759-770.

Pevny,L., Simon,M.C., Robertson,E., Klein,W.H., Tsai,S.F., D'Agati,V., Orkin,S.H., and Costantini,F. (1991). Erythroid differentiation in chimaeric mice blocked by a targeted mutation in the gene for transcription factor GATA-1. Nature 349, 257-260.

Pineault,N., Helgason,C.D., Lawrence,H.J., and Humphries,R.K. (2002). Differential expression of Hox, Meis1, and Pbx1 genes in primitive cells throughout murine hematopoietic ontogeny. Exp. Hematol. 30, 49-57.

Scott,E.W., Simon,M.C., Anastasi,J., and Singh,H. (1994). Requirement of transcription factor PU.1 in the development of multiple hematopoietic lineages. Science 265, 1573-1577.

Swift,G.H., Liu,Y., Rose,S.D., Bischof,L.J., Steelman,S., Buchberg,A.M., Wright,C.V., and MacDonald,R.J. (1998). An endocrine-exocrine switch in the activity of the pancreatic homeodomain protein PDX1 through formation of a trimeric complex with PBX1b and MRG1 (MEIS2). Mol. Cell Biol. 18, 5109-5120.

Tsai,F.Y., Keller,G., Kuo,F.C., Weiss,M., Chen,J., Rosenblatt,M., Alt,F.W., and Orkin,S.H. (1994). An early haematopoietic defect in mice lacking the transcription factor GATA-2. Nature 371, 221-226.

Tsang,A.P., Fujiwara,Y., Hom,D.B., and Orkin,S.H. (1998). Failure of megakaryopoiesis and arrested erythropoiesis in mice lacking the GATA-1 transcriptional cofactor FOG. Genes Dev. 12, 1176-1188.

Vyas,P., Ault,K., Jackson,C.W., Orkin,S.H., and Shivdasani,R.A. (1999). Consequences of GATA-1 deficiency in megakaryocytes and platelets. Blood 93, 2867-2875.

Wang,J.C., Doedens,M., and Dick,J.E. (1997). Primitive human hematopoietic cells are enriched in cord blood compared with adult bone marrow or mobilized peripheral blood as measured by the quantitative in vivo SCID-repopulating cell assay. Blood 89, 3919-3924.

Anmerkungen

The source is not given.

Note: Bain et al (1994) is not listed in the list of references of the source, neither is Wang et al. (1996), Mauoka (2002) and Emambokus (2003).

Sichter
(Hindemith), SleepyHollow02

[41.] Pak/Fragment 021 01 - Diskussion
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The production of mature blood cells from HSC requires three distinct genetic programs. These include: (a) the specification of HSC, (b) their self-renewal and c) their commitment/ proliferation/ differentiation. Most of the studies leading to the knowledge of genes involved in HSC genetic programs have been carried by assaying haematopoietic cells from animals deficient for the gene of interest. More recently, expression profiling strategies have been used to determine genetic and molecular signatures of HSC.68,69 Genes involved in deciding fate of HSCs during early embryogenesis include: SCL and Rbtn2/Lmo-2, which are necessary for primitive and definitive haematopoiesis.70,71 GATA-2 and AML1 are specifically required for definitive haematopoiesis.72

68.Ivanova M, Rozemuller E, Tyufekchiev N, Michailova A, Tilanus M, Naumova E. HLA polymorphism in Bulgarians defined by high-resolution typing methods in comparison with other populations. Tissue Antigens. 2002;60:496-504.

69.Georgantas RW, 3rd, Tanadve V, Malehorn M, et al. Microarray and serial analysis of gene expression analyses identify known and novel transcripts overexpressed in hematopoietic stem cells. Cancer Res. 2004;64:4434-4441.

70.Shivdasani RA, Mayer EL, Orkin SH. Absence of blood formation in mice lacking the T-cell leukaemia oncoprotein tal-1/SCL. Nature. 1995;373:432-434.

71.Warren HS, Kinnear BF, Skipsey LJ, Pembrey RG. Differential expression of CD45R0 on natural killer (NK) cells in patients with an NK lymphocytosis. Immunol Cell Biol. 1994;72:500-507.

72.Okuda T, van Deursen J, Hiebert SW, Grosveld G, Downing JR. AML1, the target of multiple chromosomal translocations in human leukemia, is essential for normal fetal liver hematopoiesis. Cell. 1996;84:321-330.

The production of mature blood cells from HSC requires three distinct genetic programs. These include: a) the specification of HSC, b) their self-renewal and c) their commitment/proliferation/differentiation (fig 1.2).

[page 7]

Most of the studies leading to the knowledge of genes involved in HSC genetic programs have been carried by assaying haematopoietic cells from animals deficient for the gene of interest. More recently, expression profiling strategies have been used to determine genetic and molecular signatures of HSC (Ivanova et al., 2002).

Genes involved in specifying HSC during early embryogenesis include: SCL and Rbtn2/Lmo-2 which are necessary for primitive and definitive haematopoiesis (Shivdasani et al., 1995; Warren et al., 1994). GATA-2 and AML1 are specifically required for definitive haematopoiesis (Okuda et al., 1996).


Ivanova,N.B., Dimos,J.T., Schaniel,C., Hackney,J.A., Moore,K.A., and Lemischka,I.R. (2002). A stem cell molecular signature. Science 298, 601-604.

Okuda,T., van Deursen,J., Hiebert,S.W., Grosveld,G., and Downing,J.R. (1996). AML1, the target of multiple chromosomal translocations in human leukemia, is essential for normal fetal liver hematopoiesis. Cell 84, 321-330.

Shivdasani,R.A., Mayer,E.L., and Orkin,S.H. (1995). Absence of blood formation in mice lacking the T-cell leukaemia oncoprotein tal-1/SCL. Nature 373, 432-434.

Warren,A.J., Colledge,W.H., Carlton,M.B., Evans,M.J., Smith,A.J., and Rabbitts,T.H. (1994). The oncogenic cysteine-rich LIM domain protein rbtn2 is essential for erythroid development. Cell 78, 45-57.

Anmerkungen

The source is not mentioned. Interestingly, the author gives different publications of Warren et al. (1994) and Ivanova et al. (2002) compared to the source.

Sichter
(Hindemith), SleepyHollow02

[42.] Pak/Fragment 008 25 - Diskussion
Bearbeitet: 5. April 2014, 20:53 Hindemith
Erstellt: 10. March 2014, 02:55 (Hindemith)
Bryder et al 2006, Fragment, Gesichtet, Pak, SMWFragment, Schutzlevel sysop, Verschleierung

Typus
Verschleierung
Bearbeiter
Hindemith
Gesichtet
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Untersuchte Arbeit:
Seite: 8, Zeilen: 25-28
Quelle: Bryder_et_al_2006
Seite(n): 339, Zeilen: r.col: 10-16
Because of the very short life span of most effecter-cells [sic], mature blood cell production is an ongoing process, with the production of 1.5x106 blood cells every second in an adult human. This high turnover rate necessitates some homeostatic controlmechanisms; the primary level of which resides with the HSCs. Because of the very short life span of most effector cells, mature blood cell production is an ongoing process with estimates suggesting the production of 1.5x106 blood cells every second in an adult human. This high turnover rate necessitates profound homeostatic control mechanisms, the primary level of which resides with the HSCs.
Anmerkungen

The source is not mentioned here.

Sichter
(Hindemith) Schumann

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