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n2:pmcid: PMC0
bibo:doi: 10.1084%2Fjem.20110132
n9:contains: _:vb10410031 _:vb10410005 _:vb10409995 _:vb10409956 _:vb10409944

Subject Item: _:vb10409944
rdf:type: n9:Section
dc:title: mesenchymal stem cells (mscs): the main niche players
n9:contains: _:vb10409948 _:vb10409949 _:vb10409950 _:vb10409951 _:vb10409945 _:vb10409946 _:vb10409947 _:vb10409952 _:vb10409953 _:vb10409954 _:vb10409955

Subject Item: _:vb10409945
rdf:type: n2:Context
rdf:value: Support for the hypothesis that one cell type is present at various sites within the marrow niche was obtained by a recent landmark study by Méndez-Ferrer et al. (>>2010<<). This group identified a stromal nestin-expressing MSC population (nestin+ MSC) that is closely associated with putative HSCs.
n2:mentions: n3:20703299

Subject Item: _:vb10409946
rdf:type: n2:Context
rdf:value: nestin+ MSCs are tightly associated with adrenergic nerve fibers of the sympathetic nervous system (SNS) that regulate HSC mobilization and are responsible for the circadian oscillations in circulating HSC numbers (Katayama et al., >>2006<<; Méndez-Ferrer et al., 2008).
n2:mentions: n3:16439213

Subject Item: _:vb10409947
rdf:type: n2:Context
rdf:value: associated with adrenergic nerve fibers of the sympathetic nervous system (SNS) that regulate HSC mobilization and are responsible for the circadian oscillations in circulating HSC numbers (Katayama et al., 2006; Méndez-Ferrer et al., >>2008<<). Strikingly, these MSCs express higher levels of HSC maintenance factor transcripts, including CXCL12, stem cell factor (SCF), angiopoietin-1 (Ang-1), IL-7, vascular cell adhesion molecule 1 (VCAM1), and osteopontin (OPN), compared with
n2:mentions: n3:18256599

Subject Item: _:vb10409948
rdf:type: n2:Context
rdf:value: These cells also seem to mediate the capacity of parathyroid hormone to increase HSC numbers, which had previously been correlated with an increase in putative niche OBs (Calvi et al., >>2003<<; Adams et al., 2007).
n2:mentions: n3:14574413

Subject Item: _:vb10409949
rdf:type: n2:Context
rdf:value: These cells also seem to mediate the capacity of parathyroid hormone to increase HSC numbers, which had previously been correlated with an increase in putative niche OBs (Calvi et al., 2003; Adams et al., >>2007<<). Parathyroid hormone directly stimulated proliferation of nestin+ MSCs while simultaneously promoting their differentiation into OBs (Méndez-Ferrer et al., 2010). Collectively, these data support the hypothesis that nestin+ MSCs are a
n2:mentions: n3:17237769

Subject Item: _:vb10409950
rdf:type: n2:Context
rdf:value: Parathyroid hormone directly stimulated proliferation of nestin+ MSCs while simultaneously promoting their differentiation into OBs (Méndez-Ferrer et al., >>2010<<). Collectively, these data support the hypothesis that nestin+ MSCs are a functional component of the bone marrow HSC niche.
n2:mentions: n3:20703299

Subject Item: _:vb10409951
rdf:type: n2:Context
rdf:value: Most interestingly, nestin+ MSCs show several similarities to recently identified mesenchymal progenitors (Sugiyama et al., >>2006<<; Omatsu et al., 2010). These bipotent adipoosteogenic progenitors were identified in a mouse strain in which GFP was expressed from the endogenous CXCL12 locus (Sugiyama et al., 2006; Omatsu et al., 2010).
n2:mentions: n3:17174120

Subject Item: _:vb10409952
rdf:type: n2:Context
rdf:value: Most interestingly, nestin+ MSCs show several similarities to recently identified mesenchymal progenitors (Sugiyama et al., 2006; Omatsu et al., >>2010<<). These bipotent adipoosteogenic progenitors were identified in a mouse strain in which GFP was expressed from the endogenous CXCL12 locus (Sugiyama et al., 2006; Omatsu et al., 2010).
n2:mentions: n3:20850355

Subject Item: _:vb10409953
rdf:type: n2:Context
rdf:value: These bipotent adipoosteogenic progenitors were identified in a mouse strain in which GFP was expressed from the endogenous CXCL12 locus (Sugiyama et al., >>2006<<; Omatsu et al., 2010).
n2:mentions: n3:17174120

Subject Item: _:vb10409954
rdf:type: n2:Context
rdf:value: These bipotent adipoosteogenic progenitors were identified in a mouse strain in which GFP was expressed from the endogenous CXCL12 locus (Sugiyama et al., 2006; Omatsu et al., >>2010<<). Because of their high CXCL12 expression and their long cellular processes, these cells were named CXCL12-abundant reticular (CAR) cells. The majority of putative HSCs are found in close proximity to CAR cells by immunohistochemistry,
n2:mentions: n3:20850355

Subject Item: _:vb10409955
rdf:type: n2:Context
rdf:value: These data are in agreement with studies in humans, suggesting that virtually all MSC activity is found within the larger pericyte population that associates closely with the vascular system in the entire body (Crisan et al., >>2008<<).
n2:mentions: n3:18786417

Subject Item: _:vb10409956
rdf:type: n9:Section
dc:title: obs and the endosteal niche
n9:contains: _:vb10409988 _:vb10409989 _:vb10409990 _:vb10409991 _:vb10409984 _:vb10409985 _:vb10409986 _:vb10409987 _:vb10409992 _:vb10409993 _:vb10409994 _:vb10409972 _:vb10409973 _:vb10409974 _:vb10409975 _:vb10409968 _:vb10409969 _:vb10409970 _:vb10409971 _:vb10409980 _:vb10409981 _:vb10409982 _:vb10409983 _:vb10409976 _:vb10409977 _:vb10409978 _:vb10409979 _:vb10409957 _:vb10409958 _:vb10409959 _:vb10409964 _:vb10409965 _:vb10409966 _:vb10409967 _:vb10409960 _:vb10409961 _:vb10409962 _:vb10409963

Subject Item: _:vb10409957
rdf:type: n2:Context
rdf:value: This was the first location to be proposed as the putative HSC niche, and HSCs at the endosteum have subsequently been identified by confocal/two-photon intravital imaging (Lord and Hendry, >>1972<<; Nilsson et al., 2001; Calvi et al., 2003; Zhang et al., 2003).
n2:mentions: n3:4550739

Subject Item: _:vb10409958
rdf:type: n2:Context
rdf:value: This was the first location to be proposed as the putative HSC niche, and HSCs at the endosteum have subsequently been identified by confocal/two-photon intravital imaging (Lord and Hendry, 1972; Nilsson et al., >>2001<<; Calvi et al., 2003; Zhang et al., 2003).
n2:mentions: n3:11290590

Subject Item: _:vb10409959
rdf:type: n2:Context
rdf:value: This was the first location to be proposed as the putative HSC niche, and HSCs at the endosteum have subsequently been identified by confocal/two-photon intravital imaging (Lord and Hendry, 1972; Nilsson et al., 2001; Calvi et al., >>2003<<; Zhang et al., 2003). HSCs isolated from endosteal regions by FACS show higher reconstitution activity compared with phenotypically identical cells from the center of the bone marrow (Grassinger et al., 2010).
n2:mentions: n3:14574413

Subject Item: _:vb10409960
rdf:type: n2:Context
rdf:value: location to be proposed as the putative HSC niche, and HSCs at the endosteum have subsequently been identified by confocal/two-photon intravital imaging (Lord and Hendry, 1972; Nilsson et al., 2001; Calvi et al., 2003; Zhang et al., >>2003<<). HSCs isolated from endosteal regions by FACS show higher reconstitution activity compared with phenotypically identical cells from the center of the bone marrow (Grassinger et al., 2010).
n2:mentions: n3:14574412

Subject Item: _:vb10409961
rdf:type: n2:Context
rdf:value: HSCs isolated from endosteal regions by FACS show higher reconstitution activity compared with phenotypically identical cells from the center of the bone marrow (Grassinger et al., >>2010<<).
n2:mentions: n3:20631378

Subject Item: _:vb10409962
rdf:type: n2:Context
rdf:value: Induced depletion of OBs causes mobilization of HSCs/progenitors to the spleen, and some genetically modified mice with augmented OBs show a simultaneous increase in HSC numbers (Calvi et al., >>2003<<; Zhang et al., 2003; Visnjic et al., 2004; Zhu et al., 2007).
n2:mentions: n3:14574413

Subject Item: _:vb10409963
rdf:type: n2:Context
rdf:value: Induced depletion of OBs causes mobilization of HSCs/progenitors to the spleen, and some genetically modified mice with augmented OBs show a simultaneous increase in HSC numbers (Calvi et al., 2003; Zhang et al., >>2003<<; Visnjic et al., 2004; Zhu et al., 2007).
n2:mentions: n3:14574412

Subject Item: _:vb10409964
rdf:type: n2:Context
rdf:value: Induced depletion of OBs causes mobilization of HSCs/progenitors to the spleen, and some genetically modified mice with augmented OBs show a simultaneous increase in HSC numbers (Calvi et al., 2003; Zhang et al., 2003; Visnjic et al., >>2004<<; Zhu et al., 2007). Furthermore, a population of CD45−CD105+ osteoprogenitor cells from fetal long bone or calvaria is sufficient to generate an HSC niche when transplanted under the kidney capsule (Chan et al., 2009).
n2:mentions: n3:14726388

Subject Item: _:vb10409965
rdf:type: n2:Context
rdf:value: of OBs causes mobilization of HSCs/progenitors to the spleen, and some genetically modified mice with augmented OBs show a simultaneous increase in HSC numbers (Calvi et al., 2003; Zhang et al., 2003; Visnjic et al., 2004; Zhu et al., >>2007<<). Furthermore, a population of CD45−CD105+ osteoprogenitor cells from fetal long bone or calvaria is sufficient to generate an HSC niche when transplanted under the kidney capsule (Chan et al., 2009).
n2:mentions: n3:17227831

Subject Item: _:vb10409966
rdf:type: n2:Context
rdf:value: Furthermore, a population of CD45−CD105+ osteoprogenitor cells from fetal long bone or calvaria is sufficient to generate an HSC niche when transplanted under the kidney capsule (Chan et al., >>2009<<). Similarly, adult human bone marrow contains a population of self-renewing CD45−CD146+ perivascular osteoprogenitors that can generate bone and marrow when transplanted under the skin of immunodeficient mice (Sacchetti et al., 2007).
n2:mentions: n3:19078959

Subject Item: _:vb10409967
rdf:type: n2:Context
rdf:value: Similarly, adult human bone marrow contains a population of self-renewing CD45−CD146+ perivascular osteoprogenitors that can generate bone and marrow when transplanted under the skin of immunodeficient mice (Sacchetti et al., >>2007<<). Moreover, HSCs express the calcium ion receptor, which has been demonstrated to be important for HSC engraftment at the endosteum by enabling HSCs to follow the Ca2+ gradient that results from bone remodeling processes occurring at the
n2:mentions: n3:17956733

Subject Item: _:vb10409968
rdf:type: n2:Context
rdf:value: calcium ion receptor, which has been demonstrated to be important for HSC engraftment at the endosteum by enabling HSCs to follow the Ca2+ gradient that results from bone remodeling processes occurring at the endosteum (Adams et al., >>2006<<). Finally, OBs produce factors that are known to be involved in HSC retention and maintenance, including CXCL12, OPN, and N-cadherin, in addition to factors that keep HSCs in a quiescent state, including Ang-1, membrane-bound SCF, and
n2:mentions: n3:16382241

Subject Item: _:vb10409969
rdf:type: n2:Context
rdf:value: are known to be involved in HSC retention and maintenance, including CXCL12, OPN, and N-cadherin, in addition to factors that keep HSCs in a quiescent state, including Ang-1, membrane-bound SCF, and thrombopoietin (TPO; Arai et al., >>2004<<; Yoshihara et al., 2007; Thoren et al., 2008).
n2:mentions: n3:15260986

Subject Item: _:vb10409970
rdf:type: n2:Context
rdf:value: in HSC retention and maintenance, including CXCL12, OPN, and N-cadherin, in addition to factors that keep HSCs in a quiescent state, including Ang-1, membrane-bound SCF, and thrombopoietin (TPO; Arai et al., 2004; Yoshihara et al., >>2007<<; Thoren et al., 2008).
n2:mentions: n3:18371409

Subject Item: _:vb10409971
rdf:type: n2:Context
rdf:value: maintenance, including CXCL12, OPN, and N-cadherin, in addition to factors that keep HSCs in a quiescent state, including Ang-1, membrane-bound SCF, and thrombopoietin (TPO; Arai et al., 2004; Yoshihara et al., 2007; Thoren et al., >>2008<<).
n2:mentions: n3:18250409

Subject Item: _:vb10409972
rdf:type: n2:Context
rdf:value: Accordingly, a recent study suggests that osterix+ osteoprogenitors rather than mature osteocalcin+ OBs are required for the integrity of the niche (Raaijmakers et al., >>2010<<). In addition, MSCs seem to express much higher levels of some HSC maintenance factors compared with OBs (CXCL12, SCF, IL-7, VCAM1, and OPN; Méndez-Ferrer et al., 2010). However, to what degree the expression levels of these factors
n2:mentions: n3:20305640

Subject Item: _:vb10409973
rdf:type: n2:Context
rdf:value: In addition, MSCs seem to express much higher levels of some HSC maintenance factors compared with OBs (CXCL12, SCF, IL-7, VCAM1, and OPN; Méndez-Ferrer et al., >>2010<<). However, to what degree the expression levels of these factors influence niche activity remains unclear. Importantly, sinusoidal endothelium with associated MSC/CAR cells is also found directly at the endosteum. This raises the
n2:mentions: n3:20703299

Subject Item: _:vb10409974
rdf:type: n2:Context
rdf:value: Conversely, some bone marrow HSCs localized far away from bone surfaces might simply have been caught in histological sections during migration from a niche to a vessel (Wright et al., >>2001<<). Moreover, mobilized HSCs present at extramedullary sites like the spleen have no contact with osteoblastic cells, although whether the splenic environment maintains HSC self-renewal as well as the bone marrow remains questionable.
n2:mentions: n3:11729320

Subject Item: _:vb10409975
rdf:type: n2:Context
rdf:value: Although it has been suggested that the endosteum is not important for HSC function, it may be too early to dismiss OBs as critical niche components (Kiel et al., >>2005<<; Méndez-Ferrer et al., 2010). OBs express several factors important for HSC function, including membrane-bound SCF, and they seem to be the exclusive source of TPO (Arai et al., 2009).
n2:mentions: n3:15989959

Subject Item: _:vb10409976
rdf:type: n2:Context
rdf:value: Although it has been suggested that the endosteum is not important for HSC function, it may be too early to dismiss OBs as critical niche components (Kiel et al., 2005; Méndez-Ferrer et al., >>2010<<). OBs express several factors important for HSC function, including membrane-bound SCF, and they seem to be the exclusive source of TPO (Arai et al., 2009).
n2:mentions: n3:20703299

Subject Item: _:vb10409977
rdf:type: n2:Context
rdf:value: OBs express several factors important for HSC function, including membrane-bound SCF, and they seem to be the exclusive source of TPO (Arai et al., >>2009<<). Importantly, TPO signaling via the c-MPL receptor has been shown to mediate HSC quiescence (Qian et al., 2007; Yoshihara et al., 2007), which is a typical feature of the most potent HSCs in steady-state bone marrow (Wilson et al., 2008).
n2:mentions: n3:19796231

Subject Item: _:vb10409978
rdf:type: n2:Context
rdf:value: Importantly, TPO signaling via the c-MPL receptor has been shown to mediate HSC quiescence (Qian et al., >>2007<<; Yoshihara et al., 2007), which is a typical feature of the most potent HSCs in steady-state bone marrow (Wilson et al., 2008).
n2:mentions: n3:18371408

Subject Item: _:vb10409979
rdf:type: n2:Context
rdf:value: Importantly, TPO signaling via the c-MPL receptor has been shown to mediate HSC quiescence (Qian et al., 2007; Yoshihara et al., >>2007<<), which is a typical feature of the most potent HSCs in steady-state bone marrow (Wilson et al., 2008).
n2:mentions: n3:18371409

Subject Item: _:vb10409980
rdf:type: n2:Context
rdf:value: Importantly, TPO signaling via the c-MPL receptor has been shown to mediate HSC quiescence (Qian et al., 2007; Yoshihara et al., 2007), which is a typical feature of the most potent HSCs in steady-state bone marrow (Wilson et al., >>2008<<). Moreover, although c-MPL−/− mice are born with normal numbers of HSCs, their frequency progressively declines with age, demonstrating a critical role for OB-derived TPO in adult HSC maintenance in vivo (Qian et al., 2007; Yoshihara et
n2:mentions: n3:19062086

Subject Item: _:vb10409981
rdf:type: n2:Context
rdf:value: Moreover, although c-MPL−/− mice are born with normal numbers of HSCs, their frequency progressively declines with age, demonstrating a critical role for OB-derived TPO in adult HSC maintenance in vivo (Qian et al., >>2007<<; Yoshihara et al., 2007).
n2:mentions: n3:18371408

Subject Item: _:vb10409982
rdf:type: n2:Context
rdf:value: although c-MPL−/− mice are born with normal numbers of HSCs, their frequency progressively declines with age, demonstrating a critical role for OB-derived TPO in adult HSC maintenance in vivo (Qian et al., 2007; Yoshihara et al., >>2007<<).
n2:mentions: n3:18371409

Subject Item: _:vb10409983
rdf:type: n2:Context
rdf:value: various putative niche cell types have been experimentally ablated using the DTR or thymidine kinase systems, in none of these experiments (MSC, CAR, or OB) did HSCs disappear completely or become mobilized as a whole (Visnjic et al., >>2004<<; Zhu et al., 2007; Méndez-Ferrer et al., 2010; Omatsu et al., 2010). This suggests either that the depletion systems are inefficient or that there is some redundancy in the system (or both).
n2:mentions: n3:14726388

Subject Item: _:vb10409984
rdf:type: n2:Context
rdf:value: cell types have been experimentally ablated using the DTR or thymidine kinase systems, in none of these experiments (MSC, CAR, or OB) did HSCs disappear completely or become mobilized as a whole (Visnjic et al., 2004; Zhu et al., >>2007<<; Méndez-Ferrer et al., 2010; Omatsu et al., 2010). This suggests either that the depletion systems are inefficient or that there is some redundancy in the system (or both).
n2:mentions: n3:17227831

Subject Item: _:vb10409985
rdf:type: n2:Context
rdf:value: ablated using the DTR or thymidine kinase systems, in none of these experiments (MSC, CAR, or OB) did HSCs disappear completely or become mobilized as a whole (Visnjic et al., 2004; Zhu et al., 2007; Méndez-Ferrer et al., >>2010<<; Omatsu et al., 2010). This suggests either that the depletion systems are inefficient or that there is some redundancy in the system (or both).
n2:mentions: n3:20703299

Subject Item: _:vb10409986
rdf:type: n2:Context
rdf:value: using the DTR or thymidine kinase systems, in none of these experiments (MSC, CAR, or OB) did HSCs disappear completely or become mobilized as a whole (Visnjic et al., 2004; Zhu et al., 2007; Méndez-Ferrer et al., 2010; Omatsu et al., >>2010<<). This suggests either that the depletion systems are inefficient or that there is some redundancy in the system (or both).
n2:mentions: n3:20850355

Subject Item: _:vb10409987
rdf:type: n2:Context
rdf:value: dormant during homeostasis, they can be induced to reenter the cell cycle in response to cytokines like type I or II IFN, G-CSF, or injury signals such as those generated by chemotherapy-induced myelosuppression (Wilson et al., >>2008<<; Essers et al., 2009; Foudi et al., 2009; Schaniel and Moore, 2009; Baldridge et al., 2010; Trumpp et al., 2010; Takizawa et al., 2011).
n2:mentions: n3:19062086

Subject Item: _:vb10409988
rdf:type: n2:Context
rdf:value: homeostasis, they can be induced to reenter the cell cycle in response to cytokines like type I or II IFN, G-CSF, or injury signals such as those generated by chemotherapy-induced myelosuppression (Wilson et al., 2008; Essers et al., >>2009<<; Foudi et al., 2009; Schaniel and Moore, 2009; Baldridge et al., 2010; Trumpp et al., 2010; Takizawa et al., 2011).
n2:mentions: n3:19212321

Subject Item: _:vb10409989
rdf:type: n2:Context
rdf:value: be induced to reenter the cell cycle in response to cytokines like type I or II IFN, G-CSF, or injury signals such as those generated by chemotherapy-induced myelosuppression (Wilson et al., 2008; Essers et al., 2009; Foudi et al., >>2009<<; Schaniel and Moore, 2009; Baldridge et al., 2010; Trumpp et al., 2010; Takizawa et al., 2011).
n2:mentions: n3:19060879

Subject Item: _:vb10409990
rdf:type: n2:Context
rdf:value: cell cycle in response to cytokines like type I or II IFN, G-CSF, or injury signals such as those generated by chemotherapy-induced myelosuppression (Wilson et al., 2008; Essers et al., 2009; Foudi et al., 2009; Schaniel and Moore, >>2009<<; Baldridge et al., 2010; Trumpp et al., 2010; Takizawa et al., 2011). Thus, we postulate the existence of two highly similar niches, one harboring dormant HSCs and one housing self-renewing HSCs (Fig.
n2:mentions: n3:19796230

Subject Item: _:vb10409991
rdf:type: n2:Context
rdf:value: to cytokines like type I or II IFN, G-CSF, or injury signals such as those generated by chemotherapy-induced myelosuppression (Wilson et al., 2008; Essers et al., 2009; Foudi et al., 2009; Schaniel and Moore, 2009; Baldridge et al., >>2010<<; Trumpp et al., 2010; Takizawa et al., 2011). Thus, we postulate the existence of two highly similar niches, one harboring dormant HSCs and one housing self-renewing HSCs (Fig.
n2:mentions: n3:20535209

Subject Item: _:vb10409992
rdf:type: n2:Context
rdf:value: I or II IFN, G-CSF, or injury signals such as those generated by chemotherapy-induced myelosuppression (Wilson et al., 2008; Essers et al., 2009; Foudi et al., 2009; Schaniel and Moore, 2009; Baldridge et al., 2010; Trumpp et al., >>2010<<; Takizawa et al., 2011). Thus, we postulate the existence of two highly similar niches, one harboring dormant HSCs and one housing self-renewing HSCs (Fig.
n2:mentions: n3:20182459

Subject Item: _:vb10409993
rdf:type: n2:Context
rdf:value: or injury signals such as those generated by chemotherapy-induced myelosuppression (Wilson et al., 2008; Essers et al., 2009; Foudi et al., 2009; Schaniel and Moore, 2009; Baldridge et al., 2010; Trumpp et al., 2010; Takizawa et al., >>2011<<). Thus, we postulate the existence of two highly similar niches, one harboring dormant HSCs and one housing self-renewing HSCs (Fig.
n2:mentions: n3:21300914

Subject Item: _:vb10409994
rdf:type: n2:Context
rdf:value: 2; Trumpp et al., >>2010<<). Both niches may rely on perivascular MSCs, but the endosteal niche could maintain a small dormant HSC population via the additional presence of TPO-producing OBs embedded in a calcium-rich microenvironment.
n2:mentions: n3:20182459

Subject Item: _:vb10409995
rdf:type: n9:Section
dc:title: macrophages join the niche
n9:contains: _:vb10409996 _:vb10409997 _:vb10409998 _:vb10409999 _:vb10410000 _:vb10410001 _:vb10410002 _:vb10410003 _:vb10410004

Subject Item: _:vb10409996
rdf:type: n2:Context
rdf:value: studies (including Chow et al. and Christopher et al. in the previous issue of the JEM) add to this complexity by reporting a role for bone marrow mononuclear phagocytes in promoting maintenance and retention of HSCs (Winkler et al., >>2010<<). First, Chow et al. (2011) developed a strategy to subdivide neutrophils, Gr-1hi and Gr-1lo monocytes, and Gr1−F4/80+CD169+ macrophages.
n2:mentions: n3:20713966

Subject Item: _:vb10409997
rdf:type: n2:Context
rdf:value: First, Chow et al. (>>2011<<) developed a strategy to subdivide neutrophils, Gr-1hi and Gr-1lo monocytes, and Gr1−F4/80+CD169+ macrophages.
n2:mentions: n3:21282381

Subject Item: _:vb10409998
rdf:type: n2:Context
rdf:value: Because CXCL12-mediated activation of the CXCR4 receptor on HSCs is a critical niche retention signal (Lapidot and Petit, >>2002<<; Broxmeyer, 2008), these data provide a plausible explanation for the phenotype observed in these mice.
n2:mentions: n3:12225788

Subject Item: _:vb10409999
rdf:type: n2:Context
rdf:value: Because CXCL12-mediated activation of the CXCR4 receptor on HSCs is a critical niche retention signal (Lapidot and Petit, 2002; Broxmeyer, >>2008<<), these data provide a plausible explanation for the phenotype observed in these mice.
n2:mentions: n3:18043246

Subject Item: _:vb10410000
rdf:type: n2:Context
rdf:value: Thus, as the CD169-driven DTR model was the most restricted, the loss of CD169+ macrophages rather than monocytes appears to be critical for the mobilization of HSCs (Chow et al., >>2011<<).
n2:mentions: n3:21282381

Subject Item: _:vb10410001
rdf:type: n2:Context
rdf:value: A critical role for monocytes/macrophages was also suggested by Winkler et al. (>>2010<<). After phagocyte depletion using clodronate liposomes or the MAFIA model, they also observed mobilization of functional HSCs with repopulating activity.
n2:mentions: n3:20713966

Subject Item: _:vb10410002
rdf:type: n2:Context
rdf:value: Most strikingly, clodronate depletion was associated with the loss of F4/80+ macrophages specifically associated with the endosteal lining, which have previously been characterized as osteomacs (Chang et al., >>2008<<; Winkler et al., 2010).
n2:mentions: n3:18606677

Subject Item: _:vb10410003
rdf:type: n2:Context
rdf:value: Most strikingly, clodronate depletion was associated with the loss of F4/80+ macrophages specifically associated with the endosteal lining, which have previously been characterized as osteomacs (Chang et al., 2008; Winkler et al., >>2010<<). The loss of phagocytes caused a significant decrease in osteoblastic activity at the bone surface, as the proportion of bone surface lined with OBs and the amount of newly formed bone matrix decreased.
n2:mentions: n3:20713966

Subject Item: _:vb10410004
rdf:type: n2:Context
rdf:value: The latter data conflict with the study by Chow et al. (>>2011<<), who did not observe reduced OB numbers or decreased OB expression of HSC maintenance factors when phagocytes were depleted.
n2:mentions: n3:21282381

Subject Item: _:vb10410005
rdf:type: n9:Section
dc:title: g-csf induces hsc mobilization by targeting monocytes
n9:contains: _:vb10410016 _:vb10410017 _:vb10410018 _:vb10410019 _:vb10410020 _:vb10410021 _:vb10410022 _:vb10410023 _:vb10410024 _:vb10410025 _:vb10410026 _:vb10410027 _:vb10410028 _:vb10410029 _:vb10410030 _:vb10410006 _:vb10410007 _:vb10410008 _:vb10410009 _:vb10410010 _:vb10410011 _:vb10410012 _:vb10410013 _:vb10410014 _:vb10410015

Subject Item: _:vb10410006
rdf:type: n2:Context
rdf:value: The recent studies also addressed the mechanism of G-CSF–induced mobilization of HSCs into the peripheral blood (Winkler et al., >>2010<<; Chow et al., 2011; Christopher et al., 2011).
n2:mentions: n3:20713966

Subject Item: _:vb10410007
rdf:type: n2:Context
rdf:value: The recent studies also addressed the mechanism of G-CSF–induced mobilization of HSCs into the peripheral blood (Winkler et al., 2010; Chow et al., >>2011<<; Christopher et al., 2011).
n2:mentions: n3:21282381

Subject Item: _:vb10410008
rdf:type: n2:Context
rdf:value: The recent studies also addressed the mechanism of G-CSF–induced mobilization of HSCs into the peripheral blood (Winkler et al., 2010; Chow et al., 2011; Christopher et al., >>2011<<). This method has been used for many years to harvest functional HSCs for stem cell transplantation (Gertz, 2010), but the cellular and molecular mechanisms for this striking indirect effect (HSCs do not express G-CSF receptor) remain
n2:mentions: n3:21282380

Subject Item: _:vb10410009
rdf:type: n2:Context
rdf:value: This method has been used for many years to harvest functional HSCs for stem cell transplantation (Gertz, >>2010<<), but the cellular and molecular mechanisms for this striking indirect effect (HSCs do not express G-CSF receptor) remain poorly understood.
n2:mentions: n3:20636438

Subject Item: _:vb10410010
rdf:type: n2:Context
rdf:value: Christopher et al. (>>2011<<) show that G-CSF receptor expression on cells of the monocytic lineage is sufficient for G-CSF–induced mobilization of HSCs (Fig.
n2:mentions: n3:21282380

Subject Item: _:vb10410011
rdf:type: n2:Context
rdf:value: debate about the role of neutrophils and neutrophil-derived proteases (neutrophil elastase, cathepsin G, and MMP9), which were previously thought to be the main drivers of HSC mobilization in response to G-CSF (Lévesque et al., >>2001<<; Heissig et al., 2002; Petit et al., 2002; Christopherson et al., 2003).
n2:mentions: n3:11520773

Subject Item: _:vb10410012
rdf:type: n2:Context
rdf:value: role of neutrophils and neutrophil-derived proteases (neutrophil elastase, cathepsin G, and MMP9), which were previously thought to be the main drivers of HSC mobilization in response to G-CSF (Lévesque et al., 2001; Heissig et al., >>2002<<; Petit et al., 2002; Christopherson et al., 2003).
n2:mentions: n3:12062105

Subject Item: _:vb10410013
rdf:type: n2:Context
rdf:value: and neutrophil-derived proteases (neutrophil elastase, cathepsin G, and MMP9), which were previously thought to be the main drivers of HSC mobilization in response to G-CSF (Lévesque et al., 2001; Heissig et al., 2002; Petit et al., >>2002<<; Christopherson et al., 2003). Although not excluding a contribution of neutrophils to G-CSF–induced mobilization, Christopher et al. (2011) demonstrate that monocytic cells are the key player in this process, at least in mice.
n2:mentions: n3:12068293

Subject Item: _:vb10410014
rdf:type: n2:Context
rdf:value: (neutrophil elastase, cathepsin G, and MMP9), which were previously thought to be the main drivers of HSC mobilization in response to G-CSF (Lévesque et al., 2001; Heissig et al., 2002; Petit et al., 2002; Christopherson et al., >>2003<<). Although not excluding a contribution of neutrophils to G-CSF–induced mobilization, Christopher et al. (2011) demonstrate that monocytic cells are the key player in this process, at least in mice.
n2:mentions: n3:12576320

Subject Item: _:vb10410015
rdf:type: n2:Context
rdf:value: Although not excluding a contribution of neutrophils to G-CSF–induced mobilization, Christopher et al. (>>2011<<) demonstrate that monocytic cells are the key player in this process, at least in mice.
n2:mentions: n3:21282380

Subject Item: _:vb10410016
rdf:type: n2:Context
rdf:value: The data by Christopher et al. (>>2011<<) suggest that G-CSF–stimulated monocytes cause the mobilization of HSCs out of their niche, but how does this work?
n2:mentions: n3:21282380

Subject Item: _:vb10410017
rdf:type: n2:Context
rdf:value: been generally attributed to an overall decrease of bone marrow and serum CXCL12 levels, which activates the CXCR4 receptor to promote HSC retention—an interaction that is blocked by the mobilizing compound AMD3100 (Broxmeyer et al., >>2005<<; De Clercq, 2009). Macrophages have now been identified as niche components that positively regulate MSCs and OBs, which in turn provide HSCs with the necessary extracellular retention factors, including CXCL12.
n2:mentions: n3:15837815

Subject Item: _:vb10410018
rdf:type: n2:Context
rdf:value: to an overall decrease of bone marrow and serum CXCL12 levels, which activates the CXCR4 receptor to promote HSC retention—an interaction that is blocked by the mobilizing compound AMD3100 (Broxmeyer et al., 2005; De Clercq, >>2009<<). Macrophages have now been identified as niche components that positively regulate MSCs and OBs, which in turn provide HSCs with the necessary extracellular retention factors, including CXCL12.
n2:mentions: n3:19161986

Subject Item: _:vb10410019
rdf:type: n2:Context
rdf:value: Christopher et al. (>>2011<<) use flow cytometry to show that inflammatory and resident monocytes were reduced up to fivefold 3 d after G-CSF treatment.
n2:mentions: n3:21282380

Subject Item: _:vb10410020
rdf:type: n2:Context
rdf:value: Although it seems unlikely that these two monocyte populations also contain macrophages, the study by Winkler et al. (>>2010<<) showed that F4/80+ osteomacs, which form a structure similar to a canopy around endosteal OBs, were undetectable as early as 2 d after G-CSF stimulation, returning 4 d after cessation of G-CSF treatment.
n2:mentions: n3:20713966

Subject Item: _:vb10410021
rdf:type: n2:Context
rdf:value: loss of monocytes/macrophages is similar to the situation in the aforementioned phagocyte depletion models, it is not surprising that G-CSF treatment resulted in the rapid depletion of endosteal osteoblastic activity (Winkler et al., >>2010<<). Histomorphometric methods revealed a 90% reduction of the bone formation rate, specifically at the endosteum.
n2:mentions: n3:20713966

Subject Item: _:vb10410022
rdf:type: n2:Context
rdf:value: This effect was apparent within 2 d of G-CSF treatment, preceding the appearance of HSCs in the blood (Winkler et al., >>2010<<). As OBs themselves do not express the G-CSFR, this effect is indirect and likely to be a consequence of G-CSF–mediated depletion of monocytes/macrophages. A positive effect of macrophages on OBs was further supported by data obtained in
n2:mentions: n3:20713966

Subject Item: _:vb10410023
rdf:type: n2:Context
rdf:value: A positive effect of macrophages on OBs was further supported by data obtained in co-culture studies suggesting that macrophages produce factors that support the growth, survival, and mineralization of OBs (Chang et al., >>2008<<; Christopher et al., 2011).
n2:mentions: n3:18606677

Subject Item: _:vb10410024
rdf:type: n2:Context
rdf:value: of macrophages on OBs was further supported by data obtained in co-culture studies suggesting that macrophages produce factors that support the growth, survival, and mineralization of OBs (Chang et al., 2008; Christopher et al., >>2011<<).
n2:mentions: n3:21282380

Subject Item: _:vb10410025
rdf:type: n2:Context
rdf:value: First, phagocyte depletion inhibited the proliferation of MSCs and their expression of osteoblastic differentiation genes such as osteoglycin, bone morphogenetic protein 4, and adiponectin (Méndez-Ferrer et al., >>2010<<). The expression of transcripts encoding HSC maintenance factors including Ang-1, SCF, and VCAM1 by MSCs also decreased. Most importantly, the expression of CXCL12, which is not produced by macrophages themselves, decreased by
n2:mentions: n3:20703299

Subject Item: _:vb10410026
rdf:type: n2:Context
rdf:value: Most importantly, the expression of CXCL12, which is not produced by macrophages themselves, decreased by approximately fivefold in nestin+ MSCs in response to G-CSF stimulation (Méndez-Ferrer et al., >>2010<<).
n2:mentions: n3:20703299

Subject Item: _:vb10410027
rdf:type: n2:Context
rdf:value: SNS fibers are intermingled with nestin+ MSCs and inhibit HSC retention by negatively affecting nestin+ MSC activity (Katayama et al., >>2006<<; Méndez-Ferrer et al., 2008, 2010).
n2:mentions: n3:16439213

Subject Item: _:vb10410028
rdf:type: n2:Context
rdf:value: SNS fibers are intermingled with nestin+ MSCs and inhibit HSC retention by negatively affecting nestin+ MSC activity (Katayama et al., 2006; Méndez-Ferrer et al., >>2008<<, 2010). The G-CSF–mediated increase in sympathetic tone promotes progenitor cell egress, and thus represents a macrophage/monocyte-independent pathway for G-CSF–mediated mobilization. This provides some explanation as to why G-CSF is a
n2:mentions: n3:18256599

Subject Item: _:vb10410029
rdf:type: n2:Context
rdf:value: SNS fibers are intermingled with nestin+ MSCs and inhibit HSC retention by negatively affecting nestin+ MSC activity (Katayama et al., 2006; Méndez-Ferrer et al., 2008, >>2010<<). The G-CSF–mediated increase in sympathetic tone promotes progenitor cell egress, and thus represents a macrophage/monocyte-independent pathway for G-CSF–mediated mobilization. This provides some explanation as to why G-CSF is a much
n2:mentions: n3:20703299

Subject Item: _:vb10410030
rdf:type: n2:Context
rdf:value: This provides some explanation as to why G-CSF is a much more effective HSC mobilizer than is phagocyte depletion (Chow et al., >>2011<<).
n2:mentions: n3:21282381

Subject Item: _:vb10410031
rdf:type: n9:Section
dc:title: the hsc niche unit
n9:contains: _:vb10410032 _:vb10410033 _:vb10410034 _:vb10410035

Subject Item: _:vb10410032
rdf:type: n2:Context
rdf:value: For example, sinusoidal endothelial cells have been suggested to play a role, as they produce angiocrine factors such as Notch ligands, which promote HSC self-renewal and regeneration after bone marrow injury (Butler et al., >>2010<<; Kobayashi et al., 2010).
n2:mentions: n3:20207228

Subject Item: _:vb10410033
rdf:type: n2:Context
rdf:value: endothelial cells have been suggested to play a role, as they produce angiocrine factors such as Notch ligands, which promote HSC self-renewal and regeneration after bone marrow injury (Butler et al., 2010; Kobayashi et al., >>2010<<). Whether sinusoidal endothelial cells also contribute to G-CSF–mediated mobilization remains to be addressed.
n2:mentions: n3:20972423

Subject Item: _:vb10410034
rdf:type: n2:Context
rdf:value: A recent study suggested that adipocytes, another cell type produced by MSCs, have an adverse effect on HSC maintenance (Naveiras et al., >>2009<<). Finally, models suggesting the existence of two distinct HSC niches in the bone marrow remain to be experimentally confirmed (Fig. 2; Trumpp et al., 2010). Although such a model would incorporate the currently available data about the
n2:mentions: n3:19516257

Subject Item: _:vb10410035
rdf:type: n2:Context
rdf:value: Finally, models suggesting the existence of two distinct HSC niches in the bone marrow remain to be experimentally confirmed (Fig. 2; Trumpp et al., >>2010<<). Although such a model would incorporate the currently available data about the niche components and link it to the finding that dormant and actively self-renewing HSCs coexist in the marrow, it remains very difficult to identify and
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