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pertaining to the formation of blood or blood cells; "hemopoietic stem cells in bone marrow" (同)haematopoietic, hemopoietic, haemopoietic, hematogenic, haematogenic

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出典(authority):フリー百科事典『ウィキペディア（Wikipedia）』「2013/02/02 20:19:56」(JST)

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Diagram showing the development of different blood cells from haematopoietic stem cell to mature cells

Haematopoiesis (from Ancient Greek: αἷμα, "blood"; ποιεῖν "to make") (or hematopoiesis in American English; sometimes also haemopoiesis or hemopoiesis) is the formation of blood cellular components. All cellular blood components are derived from haematopoietic stem cells. In a healthy adult person, approximately 10¹¹–10¹² new blood cells are produced daily in order to maintain steady state levels in the peripheral circulation.^[1]^[2]

Haematopoietic stem cells (HSCs)

Haematopoietic stem cells (HSCs) reside in the medulla of the bone (bone marrow) and have the unique ability to give rise to all of the different mature blood cell types and tissues. HSCs are self renewing: when they proliferate, at least some of their daughter cells remain as HSCs, so the pool of stem cells does not become depleted. The other daughters of HSCs (myeloid and lymphoid progenitor cells), however can each commit to any of the alternative differentiation pathways that lead to the production of one or more specific types of blood cells, but cannot self-renew. This is one of the vital processes in the body.

Lineages

Comprehensive diagram that shows the development of different blood cells from haematopoietic stem cell to mature cells

All blood cells are divided into three lineages.

Erythroid cells are the oxygen carrying red blood cells. Both reticulocytes and erythrocytes are functional and are released into the blood. In fact, a reticulocyte count estimates the rate of erythropoiesis.
Lymphocytes are the cornerstone of the adaptive immune system. They are derived from common lymphoid progenitors. The lymphoid lineage is primarily composed of T-cells and B-cells (types of white blood cells). This is lymphopoiesis.
Myelocytes, which include granulocytes, megakaryocytes and macrophages and are derived from common myeloid progenitors, are involved in such diverse roles as innate immunity, adaptive immunity, and blood clotting. This is myelopoiesis.

Granulopoiesis (or granulocytopoiesis) is haematopoiesis of granulocytes.

Megakaryocytopoiesis is haematopoiesis of megakaryocytes.

Locations

Sites of haematopoesis (human) in pre- and postnatal periods

In developing embryos, blood formation occurs in aggregates of blood cells in the yolk sac, called blood islands. As development progresses, blood formation occurs in the spleen, liver and lymph nodes. When bone marrow develops, it eventually assumes the task of forming most of the blood cells for the entire organism. However, maturation, activation, and some proliferation of lymphoid cells occurs in secondary lymphoid organs (spleen, thymus, and lymph nodes). In children, haematopoiesis occurs in the marrow of the long bones such as the femur and tibia. In adults, it occurs mainly in the pelvis, cranium, vertebrae, and sternum.

Extramedullary

In some cases, the liver, thymus, and spleen may resume their haematopoietic function, if necessary. This is called extramedullary haematopoiesis. It may cause these organs to increase in size substantially. During fetal development, since bones and thus the bone marrow develop later, the liver functions as the main haematopoetic organ. Therefore, the liver is enlarged during development.

Other vertebrates

In some vertebrates, haematopoiesis can occur wherever there is a loose stroma of connective tissue and slow blood supply, such as the gut, spleen, kidney or ovaries.

Maturation

As a stem cell matures it undergoes changes in gene expression that limit the cell types that it can become and moves it closer to a specific cell type. These changes can often be tracked by monitoring the presence of proteins on the surface of the cell. Each successive change moves the cell closer to the final cell type and further limits its potential to become a different cell type.

Determination

Cell determination appears to be dictated by the location of differentiation.^{[citation needed]} For instance, the thymus provides an ideal environment for thymocytes to differentiate into a variety of different functional T cells. For the stem cells and other undifferentiated blood cells in the bone marrow, the determination is generally explained by the determinism theory of haematopoiesis, saying that colony stimulating factors and other factors of the haematopoietic microenvironment determine the cells to follow a certain path of cell differentiation. This is the classical way of describing haematopoiesis. In fact, however, it is not really true. The ability of the bone marrow to regulate the quantity of different cell types to be produced is more accurately explained by a stochastic theory: Undifferentiated blood cells are determined to specific cell types by randomness. The haematopoietic microenvironment prevails upon some of the cells to survive and some, on the other hand, to perform apoptosis and die. By regulating this balance between different cell types, the bone marrow can alter the quantity of different cells to ultimately be produced.

Haematopoietic growth factors

Diagram including some of the important cytokines that determine which type of blood cell will be created.^[3] SCF= Stem Cell Factor Tpo= Thrombopoietin IL= Interleukin GM-CSF= Granulocyte Macrophage-colony stimulating factor Epo= Erythropoietin M-CSF= Macrophage-colony stimulating factor G-CSF= Granulocyte-colony stimulating factor SDF-1= Stromal cell-derived factor-1 FLT-3 ligand= FMS-like tyrosine kinase 3 ligand TNF-a = Tumour necrosis factor-alpha TGFβ = Transforming growth factor beta ^[4]

Red and white blood cell production is regulated with great precision in healthy humans, and the production of granulocytes is rapidly increased during infection. The proliferation and self-renewal of these cells depend on stem cell factor (SCF). Glycoprotein growth factors regulate the proliferation and maturation of the cells that enter the blood from the marrow, and cause cells in one or more committed cell lines to proliferate and mature. Three more factors that stimulate the production of committed stem cells are called colony-stimulating factors (CSFs) and include granulocyte-macrophage CSF (GM-CSF), granulocyte CSF (G-CSF) and macrophage CSF (M-CSF). These stimulate much granulocyte formation and are active on either progenitor cells or end product cells.

Erythropoietin is required for a myeloid progenitor cell to become an erythrocyte.^[3] On the other hand, thrombopoietin makes myeloid progenitor cells differentiate to megakaryocytes (thrombocyte-forming cells).^[3] Examples of cytokines and the blood cells they give rise to, is shown in the picture to the right.

Transcription factors

Growth factors initiate signal transduction pathways, altering transcription factors, that, in turn activate genes that determine the differentiation of blood cells.

The early committed progenitors express low levels of transcription factors that may commit them to discrete cell lineages. Which cell lineage is selected for differentiation may depend both on chance and on the external signals received by progenitor cells. Several transcription factors have been isolated that regulate differentiation along the major cell lineages. For instance, PU.1 commits cells to the myeloid lineage whereas GATA-1 has an essential role in erythropoietic and megakaryocytic differentiation. The Ikaros, Aiolos and Helios transcription factors play a major role in lymphoid development.^[5]

The myeloid-based model

For a decade now, the evidence is growing that HSC maturation follows a myeloid-based model instead of the 'classical' schoolbook dichotomy model. In the latter model, the HSC first generates a common myeloid-erythroid progenitor (CMEP) and a common lymphoid progenitor (CLP). The CLP produces only T or B cells. The myeloid-based model postulates that HSCs first diverge into the CMEP and a common myelo-lymphoid progenitor (CMLP), which generates T and B cell progenitors through a bipotential myeloid-T progenitor and a myeloid-B progenitor stage. The main difference is that in this new model, all erythroid, T and B lineage branches retain the potential to generate myeloid cells (even after the segregation of T and B cell lineages). The model proposes the idea of erythroid, T and B cells as specialized types of a prototypic myeloid HSC. Read more in Kawamoto et al. 2010.^[6]

References

^ Semester 4 medical lectures at Uppsala University 2008 by Leif Jansson
^ Parslow,T G.;Stites, DP.; Terr, AI.; and Imboden JB.. Medical Immunology (1 ed.). ISBN 0-8385-6278-7.
^ ^a ^b ^c Molecular cell biology. Lodish, Harvey F. 5. ed. : - New York : W. H. Freeman and Co., 2003, 973 s. b ill. ISBN 0-7167-4366-3
^
- For the growth factors also mentioned in previous version File:Hematopoiesis (human) cytokines.jpg: Molecular cell biology. Lodish, Harvey F. 5. ed. : - New York : W. H. Freeman and Co., 2003, 973 s. b ill. ISBN 0-7167-4366-3
- The rest: Rod Flower; Humphrey P. Rang; Maureen M. Dale; Ritter, James M. (2007). Rang & Dale's pharmacology. Edinburgh: Churchill Livingstone. ISBN 0-443-06911-5.
^ Rebollo, A.; C. Schmitt (2003). "Ikaros, Aiolos and Helios: Transcription regulators and lymphoid malignancies". Immunology and Cell Biology 81 (3): 171–175. doi:10.1046/j.1440-1711.2003.01159.x. PMID 12752680.
^ Kawamoto, Wada, Katsura. A revised scheme for developmental pathways of haematopoietic cells: the myeloid-based model. International Immunology 2010.

External links

Granulopoiesis from tulane.edu

UpToDate Contents

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1. 造血幹細胞の概要 overview of hematopoietic stem cells
2. 赤血球産生調節 regulation of erythropoiesis
3. 特発性血球減少症（Idiopathic cytopenias of undetermined significance［ICUS］）、clonal hematopoiesis of indeterminate potential (CHIP)、clonal cytopenias of undetermined significance (CCUS) idiopathic cytopenias of undetermined significance icus clonal hematopoiesis of indeterminate potential chip and clonal cytopenias of undetermined significance ccus
4. 原発性骨髄線維症の臨床症状および診断 clinical manifestations and diagnosis of primary myelofibrosis
5. 原因不明の汎血球減少症が生じた成人へのアプローチ approach to the adult with unexplained pancytopenia

English Journal

The specifically enhanced cellular immune responses in Pacific oyster (Crassostrea gigas) against secondary challenge with Vibrio splendidus.

Zhang T1, Qiu L2, Sun Z1, Wang L2, Zhou Z2, Liu R2, Yue F1, Sun R1, Song L3.Author information 1Key Laboratory of Experimental Marine Biology, Institute of Oceanology, Chinese Academy of Sciences, Qingdao 266071, China; University of Chinese Academy of Sciences, Beijing 100049, China.2Key Laboratory of Experimental Marine Biology, Institute of Oceanology, Chinese Academy of Sciences, Qingdao 266071, China.3Key Laboratory of Experimental Marine Biology, Institute of Oceanology, Chinese Academy of Sciences, Qingdao 266071, China. Electronic address: lshsong@qdio.ac.cn.AbstractThe increasing experimental evidences suggest that there are some forms of specific acquired immunity in invertebrates, but the underlying mechanism is not fully understood. In the present study, Pacific oyster (Crassostrea gigas) stimulated primarily by heat-killed Vibrio splendidus displayed stronger immune responses at cellular and molecular levels when they encountered the secondary challenge of live V. splendidus. The total hemocyte counts (THC) increased significantly after the primary stimulation of heat-killed V. splendidus, and it increased even higher (p<0.01) and reached the peak earlier (at 6h) after the secondary challenge with live V. splendidus compared with that of the primary stimulation. The number of new generated circulating hemocytes increased dramatically (p<0.01) at 6h after the pre-stimulated oysters received the secondary stimulation with live V. splendidus, and the phagocytic rate was also enhanced significantly (p<0.01) at 12h after the secondary stimulation. Meanwhile, the enhanced phagocytosis of hemocytes was highly specific for V. splendidus and they could distinguish Vibrio anguillarum, Vibrio coralliilyticus, Yarrowia lipolytica, and Micrococcus luteus efficiently. In addition, the mRNA expression of 12 candidate genes related to phagocytosis and hematopoiesis were also monitored, and the expression levels of CgIntegrin, CgPI3K (phosphatidylinositol 3-kinase), CgRho J, CgMAPKK (mitogen-activated protein kinase kinase), CgRab32, CgNADPH (nicotinamide adenine dinucleotide phosphate) oxidase, CgRunx1 and CgBMP7 (bone morphogenetic protein 7) in the hemocytes of pre-stimulated oysters after the secondary stimulation of V. splendidus were higher (p<0.01) than that after the primary stimulation, but there was no statistically significant changes for the genes of CgPKC (protein kinase C), CgMyosin, CgActin, and CgGATA 3. These results collectively suggested that the primary stimulation of V. splendidus led to immune priming in oyster with specifically enhanced phagocytosis and rapidly promoted regeneration of circulating hemocytes when the primed oysters encountered the secondary challenge with V. splendidus.
Developmental and comparative immunology.Dev Comp Immunol.2014 Jul;45(1):141-50. doi: 10.1016/j.dci.2014.02.015. Epub 2014 Mar 6.
The increasing experimental evidences suggest that there are some forms of specific acquired immunity in invertebrates, but the underlying mechanism is not fully understood. In the present study, Pacific oyster (Crassostrea gigas) stimulated primarily by heat-killed Vibrio splendidus displayed stron
PMID 24607288

Consequences of irradiation on bone and marrow phenotypes, and its relation to disruption of hematopoietic precursors.

Green DE1, Rubin CT2.Author information 1Department of Biomedical Engineering, Stony Brook University, Stony Brook, NY 11794-5281, USA. Electronic address: DanielleElyseGreen@gmail.com.2Department of Biomedical Engineering, Stony Brook University, Stony Brook, NY 11794-5281, USA.AbstractThe rising levels of radiation exposure, specifically for medical treatments and accidental exposures, have added great concern for the long term risks of bone fractures. Both the bone marrow and bone architecture are devastated following radiation exposure. Even sub-lethal doses cause a deficit to the bone marrow microenvironment, including a decline in hematopoietic cells, and this deficit occurs in a dose dependent fashion. Certain cell phenotypes though are more susceptible to radiation damage, with mesenchymal stem cells being more resilient than the hematopoietic stem cells. The decline in total bone marrow hematopoietic cells is accompanied with elevated adipocytes into the marrow cavity, thereby inhibiting hematopoiesis and recovery of the bone marrow microenvironment. Poor bone marrow is also associated with a decline in bone architectural quality. Therefore, the ability to maintain the bone marrow microenvironment would hinder much of the trabecular bone loss caused by radiation exposure, ultimately decreasing some comorbidities in patients exposed to radiation.
Bone.Bone.2014 Jun;63C:87-94. doi: 10.1016/j.bone.2014.02.018. Epub 2014 Mar 5.
The rising levels of radiation exposure, specifically for medical treatments and accidental exposures, have added great concern for the long term risks of bone fractures. Both the bone marrow and bone architecture are devastated following radiation exposure. Even sub-lethal doses cause a deficit to
PMID 24607941

The essential roles of core binding factors CfRunt and CfCBFβ in hemocyte production of scallop Chlamys farreri.

Yue F1, Zhou Z2, Wang L3, Sun R1, Jiang Q1, Yi Q1, Zhang T1, Song L4.Author information 1Key Laboratory of Experimental Marine Biology, Institute of Oceanology, Chinese Academy of Sciences, 7 Nanhai Rd., Qingdao 266071, China; University of Chinese Academy of Sciences, Beijing 100049, China.2Key Laboratory of Experimental Marine Biology, Institute of Oceanology, Chinese Academy of Sciences, 7 Nanhai Rd., Qingdao 266071, China.3Key Laboratory of Experimental Marine Biology, Institute of Oceanology, Chinese Academy of Sciences, 7 Nanhai Rd., Qingdao 266071, China. Electronic address: wanglingling@ms.qdio.ac.cn.4Key Laboratory of Experimental Marine Biology, Institute of Oceanology, Chinese Academy of Sciences, 7 Nanhai Rd., Qingdao 266071, China. Electronic address: lshsong@ms.qdio.ac.cn.AbstractCore binding factor (CBF) is a family of heterodimeric transcription factors composed of a DNA-binding CBFα subunit and a non-DNA-binding CBFβ subunit, which plays critical roles in regulating hematopoiesis, osteogenesis and neurogenesis. In the present study, two genes encoding Runt (designed as CfRunt) and CBFβ (designed as CfCBFβ) were cloned and characterized from scallop Chlamys farreri. The full-length cDNA of CfRunt and CfCBFβ consists of 2128bp and 1729bp encoding a predicted polypeptide of 530 and 183 amino acids with a conserved Runt domain and CBFβ domain, respectively. Electrophoretic mobility shift assay demonstrated that the recombinant CfRunt protein (rCfRunt) exhibited solid ability to bind specific DNA, whereas rCfCBFβ could remarkably increase the DNA-binding affinity of rCfRunt. The mRNA transcripts of CfRunt and CfCBFβ could be detected in all tested tissues, especially in hemocytes, heart, hepatopancreas or muscle. After bacterial challenge, the circulating total hemocyte count (THC) of scallop reduced to the lowest level at 6h (P<0.05), and then it recovered gradually to the control level at 48-96h, while the mRNA expressions of CfRunt and CfCBFβ were significant up-regulated between 6 and 48h (P<0.05). After CfRunt gene was silenced by RNA interference, the hemocyte renewal rate and circulating THC both decreased significantly (P<0.05). However, following the RNA interference of CfRunt, the mRNA expression of CfRunt was significantly induced (P<0.05) and the attenuated hemocyte renewal rate and circulating THC could be repaired partially by LPS stimulation in the CfRunt-silenced scallops. The results collectively indicated that CfRunt and CfCBFβ, as conserved transcription factors, played essential roles in regulating hemocyte production of scallop.
Developmental and comparative immunology.Dev Comp Immunol.2014 Jun;44(2):291-302. doi: 10.1016/j.dci.2014.01.008. Epub 2014 Jan 21.
Core binding factor (CBF) is a family of heterodimeric transcription factors composed of a DNA-binding CBFα subunit and a non-DNA-binding CBFβ subunit, which plays critical roles in regulating hematopoiesis, osteogenesis and neurogenesis. In the present study, two genes encoding Runt (designed as
PMID 24462835

Japanese Journal

Effects of Incubational Oxygen Concentration on Fatty Acid Metabolism and Heme Synthesis in Broiler Breeder Embryos

Tsushima Nobumichi,Ohta Yoshiyuki
Journal of poultry science 52(3), 213-216, 2015-07
NAID 40020533452

Effects of Incubational Oxygen Concentration on Fatty Acid Metabolism and Heme Synthesis in Broiler Breeder Embryos

Tsushima Nobumichi,Ohta Yoshiyuki
The Journal of Poultry Science 52(3), 213-216, 2015
Two experiments were conducted to evaluate the effects of incubational oxygen concentration on fatty acid metabolism and heme synthesis in broiler breeder embryos. We measured the hepatic activities …
NAID 130005089402

Effects of Incubational Oxygen Concentration on Fatty Acid Metabolism and Heme Synthesis in Broiler Breeder Embryos

Tsushima Nobumichi,Ohta Yoshiyuki
The Journal of Poultry Science advpub(0), 2015
Two experiments were conducted to evaluate the effects of incubational oxygen concentration on fatty acid metabolism and heme synthesis in broiler breeder embryos. We measured the hepatic activities o …
NAID 130005066317

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