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n2:pmcid: PMC0
bibo:doi: 10.1038%2Fnm.4103
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Subject Item: _:vb51921504
rdf:type: n5:Section
dc:title: introduction
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Subject Item: _:vb51921505
rdf:type: n2:Context
rdf:value: developed, and particularly with the development of NSG mice bearing a targeted deletion of the IL2-receptor gamma chain on the NOD-SCID background, engraftment of many human solid tumors and hematopoietic malignancies became feasible>>1<<. However, human hematopoietic cell engraftment often remained at low levels leading to the development of further strains with improved xenograft efficiency through overexpression or targeted insertion of human cytokines such as SCF,
n2:mentions: n3:22305562

Subject Item: _:vb51921506
rdf:type: n2:Context
rdf:value: cell engraftment often remained at low levels leading to the development of further strains with improved xenograft efficiency through overexpression or targeted insertion of human cytokines such as SCF, GM-CSF, IL-3, and TPO >>2<<-5.
n2:mentions: n3:20686503 n3:21262803 n3:14628073 n3:24633240

Subject Item: _:vb51921507
rdf:type: n2:Context
rdf:value: These mouse strains have been used extensively for the engraftment of human hematopoietic malignancies, particularly acute myeloid leukemia (AML) >>6<<. However, a large proportion of AML patient samples, in particular less aggressive subtypes such as core binding factor mutants and acute promyelocytic leukemia (APL), fail to engraft or do so at low levels that do not mimic human disease
n2:mentions: n3:25762176

Subject Item: _:vb51921508
rdf:type: n2:Context
rdf:value: However, a large proportion of AML patient samples, in particular less aggressive subtypes such as core binding factor mutants and acute promyelocytic leukemia (APL), fail to engraft or do so at low levels that do not mimic human disease >>7<<-9. Additionally, many other hematopoietic neoplasms do not engraft in the currently available mouse strains, as transplantation of MDS, MPN, or multiple myeloma has met with very limited success 10-12, although in myelodysplastic syndrome
n2:mentions: n3:19626050 n3:22699451 n3:21157036

Subject Item: _:vb51921509
rdf:type: n2:Context
rdf:value: Additionally, many other hematopoietic neoplasms do not engraft in the currently available mouse strains, as transplantation of MDS, MPN, or multiple myeloma has met with very limited success >>10<<-12, although in myelodysplastic syndrome (MDS), a recent study utilized a modified NSG engraftment assay through the co-transplantation of mesenchymal stromal cells (MSC) with HSC to identify MDS-initiating cells 13.
n2:mentions: n3:22648449 n3:22733078 n3:23388639

Subject Item: _:vb51921510
rdf:type: n2:Context
rdf:value: In these niches, HSC reside in close contact and bidirectional interaction with a complex network of cells including MSC, osteoblasts, adipocytes, vascular endothelial cells, and Schwann cells >>14<<. These niches not only provide sanctuary sites for HSC, but are also co-opted by leukemia cells in hematopoietic malignancies and can support LSC survival 15,16.
n2:mentions: n3:25100529

Subject Item: _:vb51921511
rdf:type: n2:Context
rdf:value: These niches not only provide sanctuary sites for HSC, but are also co-opted by leukemia cells in hematopoietic malignancies and can support LSC survival >>15<<,16.
n2:mentions: n3:17952057

Subject Item: _:vb51921512
rdf:type: n2:Context
rdf:value: These niches not only provide sanctuary sites for HSC, but are also co-opted by leukemia cells in hematopoietic malignancies and can support LSC survival 15,>>16<<.
n2:mentions: n3:25748932

Subject Item: _:vb51921513
rdf:type: n2:Context
rdf:value: that immature mesenchymal stromal cells from human BM (BM-MSC) can recreate a functional hematopoietic microenvironment in NSG mice upon transplantation into ectopic sites through endochondral ossification to form a humanized ossicle >>17<<. We speculated that these ossicles contain a humanized microenvironment with the proper supply of human niche factors to facilitate superior engraftment and growth of normal and malignant human hematopoietic cells.
n2:mentions: n3:25406351

Subject Item: _:vb51921514
rdf:type: n5:Section
dc:title: results
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Subject Item: _:vb51921515
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rdf:value: We sought to establish a xenotransplantation system for human normal and malignant hematopoietic cells through the generation of in vivo humanized BM niches in NSG mice >>17<< (Fig. 1a and see Supplementary Fig. 1a-f for a detailed protocol for humanized ossicle formation).
n2:mentions: n3:25406351

Subject Item: _:vb51921516
rdf:type: n2:Context
rdf:value: 1e left panel). Daily administration of an anabolic dose of human parathyroid hormone (40 μg/kg) for 28 days after BM-MSC transplantation resulted in a significant increase in the size of the humanized ossicles (Supplementary Fig. 1f).>>18<<,19 In order to confirm the human origin of ossicle bone and stromal niche elements, BM-MSC were transduced with lentivirus to stably express green fluorescent protein (GFP).
n2:mentions: n3:20110425

Subject Item: _:vb51921517
rdf:type: n2:Context
rdf:value: left panel). Daily administration of an anabolic dose of human parathyroid hormone (40 μg/kg) for 28 days after BM-MSC transplantation resulted in a significant increase in the size of the humanized ossicles (Supplementary Fig. 1f).18,>>19<< In order to confirm the human origin of ossicle bone and stromal niche elements, BM-MSC were transduced with lentivirus to stably express green fluorescent protein (GFP).
n2:mentions: n3:15878317

Subject Item: _:vb51921518
rdf:type: n2:Context
rdf:value: Note that in the non-transplanted ossicles, human leukemia cells were almost exclusively localized to the endosteal paratrabecular regions previously demonstrated to be preferred niches for both normal HSC and AML LSC >>15<<,20.
n2:mentions: n3:17952057

Subject Item: _:vb51921519
rdf:type: n2:Context
rdf:value: Note that in the non-transplanted ossicles, human leukemia cells were almost exclusively localized to the endosteal paratrabecular regions previously demonstrated to be preferred niches for both normal HSC and AML LSC 15,>>20<<.
n2:mentions: n3:14574412

Subject Item: _:vb51921520
rdf:type: n2:Context
rdf:value: recapitulate the clonal heterogeneity of patient samples, resulting in engrafting subclones that may be distinct from the dominant founding or relapse clone, presumably due to selective pressure imparted by the mouse BM microenvironment >>21<<. We investigated the clonal composition of human AML cells engrafted in the humanized ossicle niches through targeted re-sequencing of known leukemia mutations in four of our engrafted normal karyotype cases.
n2:mentions: n3:24613412

Subject Item: _:vb51921521
rdf:type: n2:Context
rdf:value: Original studies using limiting dilution xenograft assays suggested that leukemia-initiating cells (LIC) in AML were rare, comprising 1 : 10,000 or fewer cells >>22<<,23. In order to determine if transplantation of AML cells into a humanized microenvironment can facilitate increased engraftment and frequency of LIC, we conducted limiting dilution assays with two AML samples using both direct
n2:mentions: n3:7509044

Subject Item: _:vb51921522
rdf:type: n2:Context
rdf:value: Original studies using limiting dilution xenograft assays suggested that leukemia-initiating cells (LIC) in AML were rare, comprising 1 : 10,000 or fewer cells 22,>>23<<. In order to determine if transplantation of AML cells into a humanized microenvironment can facilitate increased engraftment and frequency of LIC, we conducted limiting dilution assays with two AML samples using both direct intraossicle
n2:mentions: n3:9212098

Subject Item: _:vb51921523
rdf:type: n2:Context
rdf:value: Transplantation of human acute promyelocytic leukemia (APL) into immunocompromised mice has so far been of limited success with low to no detectable engraftment >>9<<,23. Since humanized ossicle niches show superior engraftment of non-APL AML samples, we investigated the engraftment of primary human APL cells by direct intraossicle injection.
n2:mentions: n3:22699451

Subject Item: _:vb51921524
rdf:type: n2:Context
rdf:value: Transplantation of human acute promyelocytic leukemia (APL) into immunocompromised mice has so far been of limited success with low to no detectable engraftment 9,>>23<<. Since humanized ossicle niches show superior engraftment of non-APL AML samples, we investigated the engraftment of primary human APL cells by direct intraossicle injection.
n2:mentions: n3:9212098

Subject Item: _:vb51921525
rdf:type: n2:Context
rdf:value: The successful engraftment of primary human APL cells provided the opportunity to investigate leukemia-initiating cells (LIC) in this disease, which has been a point of controversy in the field.>>24<< First, we isolated a CD34+CD38-CD45RA- fraction that normally contains HSC and MPP from primary APL samples and compared engraftment of these cells to bulk APL cells in humanized ossicle niches by direct intraossicle injection.
n2:mentions: n3:14737069

Subject Item: _:vb51921526
rdf:type: n2:Context
rdf:value: Myelofibrosis (MF) is a myeloproliferative neoplasm (MPN) commonly associated with JAK2V617F and calreticulin (CALR) mutations >>25<<-28 that has shown only limited engraftment with transplantation of large numbers of patient-derived CD34+ cells in conventional xenograft models 29-31.
n2:mentions: n3:24325356 n3:24325359 n3:15858187

Subject Item: _:vb51921527
rdf:type: n2:Context
rdf:value: neoplasm (MPN) commonly associated with JAK2V617F and calreticulin (CALR) mutations 25-28 that has shown only limited engraftment with transplantation of large numbers of patient-derived CD34+ cells in conventional xenograft models >>29<<-31. We examined the ability of humanized ossicles to facilitate the engraftment of MF (see Supplementary Table 6 for patient sample information) through the direct intraossicle injection of FACS-purified CD34+ cells.
n2:mentions: n3:18612101 n3:17764815 n3:23023702

Subject Item: _:vb51921528
rdf:type: n2:Context
rdf:value: The successful engraftment of primary human MF cells provided the opportunity to investigate leukemia-initiating cells in this disease, which have long been hypothesized to reside in the HSC compartment >>32<<. We first fractionated CD34+ cells from MF samples into CD34+CD38+ and CD34+CD38lo/- subfractions and transplanted each directly into multiple humanized ossicles (n = 4 – 7 per sample).
n2:mentions: n3:14820991

Subject Item: _:vb51921529
rdf:type: n2:Context
rdf:value: B into HSC and MPP>>33<<, and tested their engraftment potential through direct intraossicle injection.
n2:mentions: n3:18371405

Subject Item: _:vb51921530
rdf:type: n5:Section
dc:title: discussion
n5:contains: _:vb51921532 _:vb51921533 _:vb51921534 _:vb51921535 _:vb51921531 _:vb51921548 _:vb51921549 _:vb51921550 _:vb51921551 _:vb51921544 _:vb51921545 _:vb51921546 _:vb51921547 _:vb51921540 _:vb51921541 _:vb51921542 _:vb51921543 _:vb51921536 _:vb51921537 _:vb51921538 _:vb51921539

Subject Item: _:vb51921531
rdf:type: n2:Context
rdf:value: Original xenotransplantation studies suggested AML LIC/LSC were rare cells, on the order of 1 : 10,000 – 100,000 enriched in the CD34+CD38- fraction of leukemic blasts >>22<<,23. More recent studies suggested that LIC/LSC are more frequent and can be found in additional immunophenotypically defined subpopulations 7,34,35. Here, we provide evidence that with a humanized niche, the frequency of LIC is
n2:mentions: n3:7509044

Subject Item: _:vb51921532
rdf:type: n2:Context
rdf:value: Original xenotransplantation studies suggested AML LIC/LSC were rare cells, on the order of 1 : 10,000 – 100,000 enriched in the CD34+CD38- fraction of leukemic blasts 22,>>23<<. More recent studies suggested that LIC/LSC are more frequent and can be found in additional immunophenotypically defined subpopulations 7,34,35. Here, we provide evidence that with a humanized niche, the frequency of LIC is substantially
n2:mentions: n3:9212098

Subject Item: _:vb51921533
rdf:type: n2:Context
rdf:value: More recent studies suggested that LIC/LSC are more frequent and can be found in additional immunophenotypically defined subpopulations >>7<<,34,35. Here, we provide evidence that with a humanized niche, the frequency of LIC is substantially higher than that determined in the NSG mouse BM with the same sample, suggesting that AML LIC frequency is critically dependent on
n2:mentions: n3:21157036

Subject Item: _:vb51921534
rdf:type: n2:Context
rdf:value: More recent studies suggested that LIC/LSC are more frequent and can be found in additional immunophenotypically defined subpopulations 7,>>34<<,35. Here, we provide evidence that with a humanized niche, the frequency of LIC is substantially higher than that determined in the NSG mouse BM with the same sample, suggesting that AML LIC frequency is critically dependent on leukemia
n2:mentions: n3:21873988

Subject Item: _:vb51921535
rdf:type: n2:Context
rdf:value: More recent studies suggested that LIC/LSC are more frequent and can be found in additional immunophenotypically defined subpopulations 7,34,>>35<<. Here, we provide evidence that with a humanized niche, the frequency of LIC is substantially higher than that determined in the NSG mouse BM with the same sample, suggesting that AML LIC frequency is critically dependent on leukemia
n2:mentions: n3:21251617

Subject Item: _:vb51921536
rdf:type: n2:Context
rdf:value: In APL, the nature of the hematopoietic cell that is the target of the disease-causing chromosomal translocation is a matter of debate>>24<< with in vitro studies implicating immature compartments able to produce myeloid and erythroid output 36, and alternative reports supporting APL arises from committed myeloid progenitors.
n2:mentions: n3:14737069

Subject Item: _:vb51921537
rdf:type: n2:Context
rdf:value: APL, the nature of the hematopoietic cell that is the target of the disease-causing chromosomal translocation is a matter of debate24 with in vitro studies implicating immature compartments able to produce myeloid and erythroid output >>36<<, and alternative reports supporting APL arises from committed myeloid progenitors.
n2:mentions: n3:8136268

Subject Item: _:vb51921538
rdf:type: n2:Context
rdf:value: chromosomal translocation is a matter of debate24 with in vitro studies implicating immature compartments able to produce myeloid and erythroid output 36, and alternative reports supporting APL arises from committed myeloid progenitors.>>37<<-39 Here, we report successful fulminant engraftment of PML-RARA-positive human APL cells in humanized ossicles, and identification of APL-initiating cells in an aberrant CD34-/lo GMP-like population.
n2:mentions: n3:7520272 n3:9792295 n3:7536493

Subject Item: _:vb51921539
rdf:type: n2:Context
rdf:value: These results are consistent with mouse models that drive expression of PML-RARA from a myeloid lineage-restricted promoter resulting in an APL-like disease >>40<<,41. The absence of PML-RARA translocation in human HSC obtained from APL patient samples is surprising given that PML-RARA on its own induced myeloid commitment rather than self-renewal in hematopoietic progenitors such as GMP 42. In
n2:mentions: n3:19797526

Subject Item: _:vb51921540
rdf:type: n2:Context
rdf:value: These results are consistent with mouse models that drive expression of PML-RARA from a myeloid lineage-restricted promoter resulting in an APL-like disease 40,>>41<<. The absence of PML-RARA translocation in human HSC obtained from APL patient samples is surprising given that PML-RARA on its own induced myeloid commitment rather than self-renewal in hematopoietic progenitors such as GMP 42. In
n2:mentions: n3:9122233

Subject Item: _:vb51921541
rdf:type: n2:Context
rdf:value: The absence of PML-RARA translocation in human HSC obtained from APL patient samples is surprising given that PML-RARA on its own induced myeloid commitment rather than self-renewal in hematopoietic progenitors such as GMP >>42<<. In contrast to a recently published report demonstrating establishment of APL from transduced cord blood-derived CMP, we did not identify PML-RARA fusion events in FACS-purified CMP from primary patient samples 43. How PML-RARA initiates
n2:mentions: n3:10942402

Subject Item: _:vb51921542
rdf:type: n2:Context
rdf:value: In contrast to a recently published report demonstrating establishment of APL from transduced cord blood-derived CMP, we did not identify PML-RARA fusion events in FACS-purified CMP from primary patient samples >>43<<. How PML-RARA initiates a self-renewal program in human progenitors to cause APL is an important question for further studies of APL pathogenesis.
n2:mentions: n3:25369030

Subject Item: _:vb51921543
rdf:type: n2:Context
rdf:value: Similar to APL, our humanized ossicle model facilitates robust engraftment of primary human MF cells, which has only been of limited success with peripheral blood derived cells with conventional NSG models.>>31<<,44 Unlike studies of AML, where disease-initiating cells arise from progenitors 35, we show here that myelofibrosis-initiating cells reside in the HSC compartment, as transplantation of these cells, but not MPP or CMP, results in
n2:mentions: n3:23023702

Subject Item: _:vb51921544
rdf:type: n2:Context
rdf:value: Similar to APL, our humanized ossicle model facilitates robust engraftment of primary human MF cells, which has only been of limited success with peripheral blood derived cells with conventional NSG models.31,>>44<< Unlike studies of AML, where disease-initiating cells arise from progenitors 35, we show here that myelofibrosis-initiating cells reside in the HSC compartment, as transplantation of these cells, but not MPP or CMP, results in engraftment
n2:mentions: n3:20858855

Subject Item: _:vb51921545
rdf:type: n2:Context
rdf:value: 31,44 Unlike studies of AML, where disease-initiating cells arise from progenitors >>35<<, we show here that myelofibrosis-initiating cells reside in the HSC compartment, as transplantation of these cells, but not MPP or CMP, results in engraftment of JAK2V617F or CALR-mutant cells.
n2:mentions: n3:21251617

Subject Item: _:vb51921546
rdf:type: n2:Context
rdf:value: Recently, several newly developed xenotransplantation models have been reported for improved engraftment of human hematopoietic cells including modified NSG strains with overexpression or knock-in of human cytokines >>2<<,4,5,45,46, co-transplantation of hematopoietic cells with mesenchymal stromal cells 13, and transplantation of engineered bio-scaffolds designed to resemble human BM niches 47.
n2:mentions: n3:20686503

Subject Item: _:vb51921547
rdf:type: n2:Context
rdf:value: Recently, several newly developed xenotransplantation models have been reported for improved engraftment of human hematopoietic cells including modified NSG strains with overexpression or knock-in of human cytokines 2,>>4<<,5,45,46, co-transplantation of hematopoietic cells with mesenchymal stromal cells 13, and transplantation of engineered bio-scaffolds designed to resemble human BM niches 47.
n2:mentions: n3:24633240

Subject Item: _:vb51921548
rdf:type: n2:Context
rdf:value: Recently, several newly developed xenotransplantation models have been reported for improved engraftment of human hematopoietic cells including modified NSG strains with overexpression or knock-in of human cytokines 2,4,>>5<<,45,46, co-transplantation of hematopoietic cells with mesenchymal stromal cells 13, and transplantation of engineered bio-scaffolds designed to resemble human BM niches 47.
n2:mentions: n3:21262803

Subject Item: _:vb51921549
rdf:type: n2:Context
rdf:value: Recently, several newly developed xenotransplantation models have been reported for improved engraftment of human hematopoietic cells including modified NSG strains with overexpression or knock-in of human cytokines 2,4,5,>>45<<,46, co-transplantation of hematopoietic cells with mesenchymal stromal cells 13, and transplantation of engineered bio-scaffolds designed to resemble human BM niches 47.
n2:mentions: n3:21262827

Subject Item: _:vb51921550
rdf:type: n2:Context
rdf:value: Recently, several newly developed xenotransplantation models have been reported for improved engraftment of human hematopoietic cells including modified NSG strains with overexpression or knock-in of human cytokines 2,4,5,45,>>46<<, co-transplantation of hematopoietic cells with mesenchymal stromal cells 13, and transplantation of engineered bio-scaffolds designed to resemble human BM niches 47.
n2:mentions: n3:21697012

Subject Item: _:vb51921551
rdf:type: n2:Context
rdf:value: strains with overexpression or knock-in of human cytokines 2,4,5,45,46, co-transplantation of hematopoietic cells with mesenchymal stromal cells 13, and transplantation of engineered bio-scaffolds designed to resemble human BM niches >>47<<. While each of these approaches has advantages compared to conventional NSG mouse models, they have not demonstrated the rapid and high leukemic engraftment observed with our humanized ossicle niche model, where direct intraossicle
n2:mentions: n3:22653974

Subject Item: _:vb51921552
rdf:type: n5:Section
dc:title: online methods
n5:contains: _:vb51921556 _:vb51921557 _:vb51921558 _:vb51921559 _:vb51921553 _:vb51921554 _:vb51921555 _:vb51921560 _:vb51921561 _:vb51921562

Subject Item: _:vb51921553
rdf:type: n2:Context
rdf:value: Human BM samples were obtained according to Medical University of Graz Ethikkommission (Institutional Review Board-approved protocol, MUG Graz IRB no. 19-252). BM-MSC were isolated and expanded as previously described.>>17<<,49,50 Age of healthy BM donors ranged from 21 – 45 years. Briefly, BM-MNCs were washed out from BM collection filters by flushing the filters with pre-warmed PBS.
n2:mentions: n3:25406351

Subject Item: _:vb51921554
rdf:type: n2:Context
rdf:value: Human BM samples were obtained according to Medical University of Graz Ethikkommission (Institutional Review Board-approved protocol, MUG Graz IRB no. 19-252). BM-MSC were isolated and expanded as previously described.17,49,>>50<< Age of healthy BM donors ranged from 21 – 45 years. Briefly, BM-MNCs were washed out from BM collection filters by flushing the filters with pre-warmed PBS.
n2:mentions: n3:18620484

Subject Item: _:vb51921555
rdf:type: n2:Context
rdf:value: in four-layered cell factories (CF-4, Thermo Fisher, Nunc, Pittsburg, PA) with half-weekly medium changes until cells reached confluence. At the end of passage 1 (p1), BM-MSC were profiled for expression of consensus MSC surface markers.>>51<< Briefly, trypsinized cells were blocked with sheep serum (10% v/v) and thereafter stained with following monoclonal antibodies for 30 min, 4°C in the dark:
n2:mentions: n3:16923606

Subject Item: _:vb51921556
rdf:type: n2:Context
rdf:value: subcutaneous injections of human parathyroid hormone (PTH [1-34]; R&D Systems, Minneapolis, MN; 40 μg/kg body weight; dorsal neck fold) for 28 consecutive days to further promote bone marrow niche formation, as previously described.>>18<<,19 Eight – 10 weeks post BM-MSC application the site of injection was shaved and transplants were evaluated for bone and marrow formation by palpation and by visual inspection (development of a purple hue is indicative of bone marrow
n2:mentions: n3:20110425

Subject Item: _:vb51921557
rdf:type: n2:Context
rdf:value: subcutaneous injections of human parathyroid hormone (PTH [1-34]; R&D Systems, Minneapolis, MN; 40 μg/kg body weight; dorsal neck fold) for 28 consecutive days to further promote bone marrow niche formation, as previously described.18,>>19<< Eight – 10 weeks post BM-MSC application the site of injection was shaved and transplants were evaluated for bone and marrow formation by palpation and by visual inspection (development of a purple hue is indicative of bone marrow niche
n2:mentions: n3:15878317

Subject Item: _:vb51921558
rdf:type: n2:Context
rdf:value: For single cell transplantation, Lin-CD34+CD38- cells were sorted into single wells of Terasaki plates (Nunc MiniTrays, Thermo Fisher) prefilled with 10 μL IMDM 10% FBS as previously described >>52<< using FACS AriaII.
n2:mentions: n3:21737740

Subject Item: _:vb51921559
rdf:type: n2:Context
rdf:value: Allele-specific JAK2 genotyping assays were performed as described previously.>>53<< Briefly, engrafted human B cells (huCD45+CD19+) and myeloid cells (huCD45+CD33+) were sorted on an Aria II flow cytometer (BD) for analysis.
n2:mentions: n3:25728669

Subject Item: _:vb51921560
rdf:type: n2:Context
rdf:value: The native samples (prior to xenotransplantation) were genotyped by whole exome sequencing (SeqCap EZ Exome SR kit v3.0, Roche/Nimblegen) as previously described >>54<<. All kits were used following manufacturer's instructions (Roche/Nimblegen). Illumina HiSeq 2000, HiSeq 2500, or NextSeq 500 instruments were used for sequencing.
n2:mentions: n3:24550281

Subject Item: _:vb51921561
rdf:type: n2:Context
rdf:value: Hematoxylin-eosin (H/E) staining and IHC staining were performed using standard protocols >>17<<. For IHC staining, temperature-based antigen retrieval was performed (70 °C, 160 W, 40 min) followed by a descending alcohol series. Endogenous peroxidases were blocked with hydrogen peroxide (10 min) and nonspecific antibody binding with
n2:mentions: n3:25406351

Subject Item: _:vb51921562
rdf:type: n2:Context
rdf:value: The absolute number of total mouse BM cells was extrapolated as previously described.>>55<< To calculate total human leukemia cells engrafted within the total humanized ossicle-niche space as compared to the mouse BM, the percentage of human CD45+CD33+ (determined by flow-cytometry) was multiplied with the maximal number of
n2:mentions: n3:6711557

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