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n2:pmcid: PMC0
bibo:doi: 10.1038%2Fncb2086
n5:contains: _:vb8183475 _:vb8183518 _:vb8183511

Subject Item: _:vb8183475
rdf:type: n5:Section
dc:title: results
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Subject Item: _:vb8183476
rdf:type: n2:Context
rdf:value: For instance, depletion of stat from all testis cells causes loss of both stem cell populations, the GSCs and CySCs, while misexpression of unpaired leads to self-renewal of both lineages away from the niche>>1<<,2. However, we recently found that germ cell-intrinsic STAT activation could not renew germ cells away from the niche3. Instead, intrinsic STAT activation in somatic cells renewed not only CySCs, but also permitted GSC renewal away from
n2:mentions: n3:11752575

Subject Item: _:vb8183477
rdf:type: n2:Context
rdf:value: For instance, depletion of stat from all testis cells causes loss of both stem cell populations, the GSCs and CySCs, while misexpression of unpaired leads to self-renewal of both lineages away from the niche1,>>2<<. However, we recently found that germ cell-intrinsic STAT activation could not renew germ cells away from the niche3. Instead, intrinsic STAT activation in somatic cells renewed not only CySCs, but also permitted GSC renewal away from the
n2:mentions: n3:11752574

Subject Item: _:vb8183478
rdf:type: n2:Context
rdf:value: However, we recently found that germ cell-intrinsic STAT activation could not renew germ cells away from the niche>>3<<. Instead, intrinsic STAT activation in somatic cells renewed not only CySCs, but also permitted GSC renewal away from the niche3. This suggested that STAT-activated CySCs produce a germline self-renewal signal, and that the previously
n2:mentions: n3:18593558

Subject Item: _:vb8183479
rdf:type: n2:Context
rdf:value: Instead, intrinsic STAT activation in somatic cells renewed not only CySCs, but also permitted GSC renewal away from the niche>>3<<. This suggested that STAT-activated CySCs produce a germline self-renewal signal, and that the previously observed loss of GSCs upon global stat depletion was secondary to CySC loss. To test this, we examined stat-depleted testes by
n2:mentions: n3:18593558

Subject Item: _:vb8183480
rdf:type: n2:Context
rdf:value: (Figure 1A, Zfh-1, green, marks CySCs), but now stat-depleted germline cells were maintained indefinitely in most testes (57/64 or 89% still had spermatogonia eight days after stat depletion; Figure 1C; see Table 1 for quantitation)>>4<<. In particular, amplifying spermatogonia and spermatocytes were observed at all stages of differentiation in their normal organization emanating from the testis apex near the hub, shown by normal Bam accumulation, branching fusomes,
n2:mentions: n3:20143337

Subject Item: _:vb8183481
rdf:type: n2:Context
rdf:value: Normally, GSCs divide perpendicular to the hub surface>>5<<, providing for a mechanism of asymmetric stem cell division, with one daughter cell invariantly pushed away from renewing influences, allowing its differentiation program to initiate.
n2:mentions: n3:114676

Subject Item: _:vb8183482
rdf:type: n2:Context
rdf:value: Such oriented divisions are directly dependent on tethering of one centrosome in the GSC to its interface with the hub>>6<<,7. While centrosome staining of wild-type mitotic GSCs shows invariant orientation6, the division plane of stat-depleted, displaced GSCs became randomized (Figure 2F shows one oriented mitotic GSC (arrow) and one non-oriented mitotic GSC
n2:mentions: n3:12970569

Subject Item: _:vb8183483
rdf:type: n2:Context
rdf:value: Such oriented divisions are directly dependent on tethering of one centrosome in the GSC to its interface with the hub6,>>7<<. While centrosome staining of wild-type mitotic GSCs shows invariant orientation6, the division plane of stat-depleted, displaced GSCs became randomized (Figure 2F shows one oriented mitotic GSC (arrow) and one non-oriented mitotic GSC
n2:mentions: n3:17255513

Subject Item: _:vb8183484
rdf:type: n2:Context
rdf:value: While centrosome staining of wild-type mitotic GSCs shows invariant orientation>>6<<, the division plane of stat-depleted, displaced GSCs became randomized (Figure 2F shows one oriented mitotic GSC (arrow) and one non-oriented mitotic GSC (arrowhead); ratio of oriented:
n2:mentions: n3:12970569

Subject Item: _:vb8183485
rdf:type: n2:Context
rdf:value: cell daughters into one renewing and one differentiating cell, demonstrating the niche’s capacity to function in a less regimented, population mode for self-renewal, where the production of differentiating daughter cells is stochastic>>8<<. We do not know whether GSCs in this alternate niche configuration completely bypass the other characteristics of asymmetric division, including stem cell inheritance of the mother centrosome and the mitotic checkpoint associated with
n2:mentions: n3:10688781

Subject Item: _:vb8183486
rdf:type: n2:Context
rdf:value: GSCs in this alternate niche configuration completely bypass the other characteristics of asymmetric division, including stem cell inheritance of the mother centrosome and the mitotic checkpoint associated with centrosome attachment>>7<<,9.
n2:mentions: n3:17255513

Subject Item: _:vb8183487
rdf:type: n2:Context
rdf:value: GSCs in this alternate niche configuration completely bypass the other characteristics of asymmetric division, including stem cell inheritance of the mother centrosome and the mitotic checkpoint associated with centrosome attachment7,>>9<<.
n2:mentions: n3:18923395

Subject Item: _:vb8183488
rdf:type: n2:Context
rdf:value: This is consistent with previous findings that Zfh-1 controls a CySC signal that can support GSC renewal>>3<<.
n2:mentions: n3:18593558

Subject Item: _:vb8183489
rdf:type: n2:Context
rdf:value: It is known that an intrinsic increase in STAT activation in CySCs causes these cells to outcompete GSCs for niche access>>10<<. Since our strategy to examine stat-depleted GSCs utilized Gal4-driven rescue of stat in CySCs, we considered whether higher than normal STAT levels in CySCs led to excess STAT activation and hence could account for the observed GSC
n2:mentions: n3:19797664

Subject Item: _:vb8183490
rdf:type: n2:Context
rdf:value: However, a principal role in adhesion is sufficient to explain earlier mosaic experiments>>1<<,2. We suggest that stat null GSCs were lost, not because stat was essential for self-renewal, but rather, because an attachment defect led to their displacement from the niche, and nearby wild-type germ cells de-differentiated11 to take
n2:mentions: n3:11752575

Subject Item: _:vb8183491
rdf:type: n2:Context
rdf:value: However, a principal role in adhesion is sufficient to explain earlier mosaic experiments1,>>2<<. We suggest that stat null GSCs were lost, not because stat was essential for self-renewal, but rather, because an attachment defect led to their displacement from the niche, and nearby wild-type germ cells de-differentiated11 to take
n2:mentions: n3:11752574

Subject Item: _:vb8183492
rdf:type: n2:Context
rdf:value: To ensure reliably oriented divisions, an elaborate cortical complex is postulated to be established in GSCs at the hub interface, linking one centrosome to the adherens junction (Figure 3A and C, respectively)>>6<<,7. Thus, in testes with global stat depletion, we examined these components in GSCs prior to overt signs of differentiation, scoring only GSCs that were still located next to the hub and that did not yet express the differentiation marker
n2:mentions: n3:12970569

Subject Item: _:vb8183493
rdf:type: n2:Context
rdf:value: To ensure reliably oriented divisions, an elaborate cortical complex is postulated to be established in GSCs at the hub interface, linking one centrosome to the adherens junction (Figure 3A and C, respectively)6,>>7<<. Thus, in testes with global stat depletion, we examined these components in GSCs prior to overt signs of differentiation, scoring only GSCs that were still located next to the hub and that did not yet express the differentiation marker
n2:mentions: n3:17255513

Subject Item: _:vb8183494
rdf:type: n2:Context
rdf:value: The BMP pathway has been implicated in GSC renewal>>12<<–14. The ligands, gbb and dpp, are each expressed in CySCs (and in hub cells), and pathway activation has been observed in GSCs12–14.
n2:mentions: n3:15238523 n3:14973292 n3:14653996

Subject Item: _:vb8183495
rdf:type: n2:Context
rdf:value: The ligands, gbb and dpp, are each expressed in CySCs (and in hub cells), and pathway activation has been observed in GSCs>>12<<–14. In testes with germline stat depletion, we observed strong BMP pathway activation in the displaced GSCs, shown by accumulation of phosphoSmad and the BMP target gene dad (Figure 4A, B). Interestingly, we noted that the levels of pSmad
n2:mentions: n3:15238523 n3:14973292 n3:14653996

Subject Item: _:vb8183496
rdf:type: n2:Context
rdf:value: Sustained zfh-1 expression in the cyst lineage, which generates ectopic GSCs far from the hub>>3<<, also led to strong BMP pathway activation in the ectopic GSCs (Figure 4C).
n2:mentions: n3:18593558

Subject Item: _:vb8183497
rdf:type: n2:Context
rdf:value: Thus, Jak/STAT activation in CySCs leads to BMP ligand expression>>15<<, likely using zfh-1 as an intermediary.
n2:mentions: n3:18283115

Subject Item: _:vb8183498
rdf:type: n2:Context
rdf:value: A primary role for BMPs in male GSCs makes the testis much more similar to the Drosophila ovary, where BMP pathway activation regulates GSC renewal via repression of the differentiation gene, bam>>16<<,17, as also occurs in the male germline18.
n2:mentions: n3:9695953

Subject Item: _:vb8183499
rdf:type: n2:Context
rdf:value: A primary role for BMPs in male GSCs makes the testis much more similar to the Drosophila ovary, where BMP pathway activation regulates GSC renewal via repression of the differentiation gene, bam16,>>17<<, as also occurs in the male germline18.
n2:mentions: n3:14561403

Subject Item: _:vb8183500
rdf:type: n2:Context
rdf:value: A primary role for BMPs in male GSCs makes the testis much more similar to the Drosophila ovary, where BMP pathway activation regulates GSC renewal via repression of the differentiation gene, bam16,17, as also occurs in the male germline>>18<<. A remaining difference, however, is that, in the male, BMP pathway activation is insufficient for induction of ectopic GSCs.
n2:mentions: n3:2279698

Subject Item: _:vb8183501
rdf:type: n2:Context
rdf:value: The Jak/STAT pathway is also similar between the two niches, with pathway activation primarily required in the somatic cell populations>>3<<,19,20. However, while the testis GSCs use Jak/STAT-regulated centrosome tethering to attach and orient the division plane, the ovary uses Jak/STAT-independent spectrosome localization to ensure an asymmetric outcome to stem cell
n2:mentions: n3:18593558

Subject Item: _:vb8183502
rdf:type: n2:Context
rdf:value: The Jak/STAT pathway is also similar between the two niches, with pathway activation primarily required in the somatic cell populations3,>>19<<,20. However, while the testis GSCs use Jak/STAT-regulated centrosome tethering to attach and orient the division plane, the ovary uses Jak/STAT-independent spectrosome localization to ensure an asymmetric outcome to stem cell divisions21.
n2:mentions: n3:16198292

Subject Item: _:vb8183503
rdf:type: n2:Context
rdf:value: The Jak/STAT pathway is also similar between the two niches, with pathway activation primarily required in the somatic cell populations3,19,>>20<<. However, while the testis GSCs use Jak/STAT-regulated centrosome tethering to attach and orient the division plane, the ovary uses Jak/STAT-independent spectrosome localization to ensure an asymmetric outcome to stem cell divisions21.
n2:mentions: n3:18171682

Subject Item: _:vb8183504
rdf:type: n2:Context
rdf:value: However, while the testis GSCs use Jak/STAT-regulated centrosome tethering to attach and orient the division plane, the ovary uses Jak/STAT-independent spectrosome localization to ensure an asymmetric outcome to stem cell divisions>>21<<.
n2:mentions: n3:9281339

Subject Item: _:vb8183505
rdf:type: n2:Context
rdf:value: The hub cells are architects of the stem cell environment, establishing close proximity of the three cell populations because STAT influences adhesion in nearby germline and somatic cells, probably through adherens junctions (this work)>>22<< and integrin linkages10, respectively. STAT activation also governs self-renewal in the somatic lineage via zfh-1 and chinmo induction3,23.
n2:mentions: n3:18641633

Subject Item: _:vb8183506
rdf:type: n2:Context
rdf:value: of the stem cell environment, establishing close proximity of the three cell populations because STAT influences adhesion in nearby germline and somatic cells, probably through adherens junctions (this work)22 and integrin linkages>>10<<, respectively. STAT activation also governs self-renewal in the somatic lineage via zfh-1 and chinmo induction3,23.
n2:mentions: n3:19797664

Subject Item: _:vb8183507
rdf:type: n2:Context
rdf:value: STAT activation also governs self-renewal in the somatic lineage via zfh-1 and chinmo induction>>3<<,23. These CySCs then activate BMP, which is used to renew neighboring, hub-adherent germ cells. Thus, self-renewal activity relies not just on the hub, but also on one of the resident stem cell lineages. BMPs are expressed from both hub
n2:mentions: n3:18593558

Subject Item: _:vb8183508
rdf:type: n2:Context
rdf:value: STAT activation also governs self-renewal in the somatic lineage via zfh-1 and chinmo induction3,>>23<<. These CySCs then activate BMP, which is used to renew neighboring, hub-adherent germ cells. Thus, self-renewal activity relies not just on the hub, but also on one of the resident stem cell lineages. BMPs are expressed from both hub and
n2:mentions: n3:20412771

Subject Item: _:vb8183509
rdf:type: n2:Context
rdf:value: BMPs are expressed from both hub and CySCs>>12<<,13, possibly to generate sufficient quantity of BMP for renewal; alternatively, the ligand composition from hub versus CySCs may be qualitatively different.
n2:mentions: n3:14653996

Subject Item: _:vb8183510
rdf:type: n2:Context
rdf:value: BMPs are expressed from both hub and CySCs12,>>13<<, possibly to generate sufficient quantity of BMP for renewal; alternatively, the ligand composition from hub versus CySCs may be qualitatively different.
n2:mentions: n3:14973292

Subject Item: _:vb8183511
rdf:type: n5:Section
dc:title: methods
n5:contains: _:vb8183512 _:vb8183513 _:vb8183514 _:vb8183515 _:vb8183516 _:vb8183517

Subject Item: _:vb8183512
rdf:type: n2:Context
rdf:value: Gal4 drivers were Traffic Jam Gal4 (Kyoto Stock Center, DGRC #104055), c587 Gal4 (Erika Matunis), or EyaA3 Gal>>43<< for CySC expression, Nos Gal4 VP16 (Ruth Lehmann and Erica Selva) for germline expression, and Upd Gal4 (Doug Harrison) for hub expression.
n2:mentions: n3:18593558

Subject Item: _:vb8183513
rdf:type: n2:Context
rdf:value: The BMP pathway was dampened by coexpressing either UAS sogCR1 (activated form of sog, FBtp0018246, Ethan Bier)>>24<< or UAS kekkon5 (FBtp0018362, Joe Duffy)25 in the CySC lineage along with UAS stat in statts flies.
n2:mentions: n3:10769238

Subject Item: _:vb8183514
rdf:type: n2:Context
rdf:value: The BMP pathway was dampened by coexpressing either UAS sogCR1 (activated form of sog, FBtp0018246, Ethan Bier)24 or UAS kekkon5 (FBtp0018362, Joe Duffy)>>25<< in the CySC lineage along with UAS stat in statts flies.
n2:mentions: n3:19013143

Subject Item: _:vb8183515
rdf:type: n2:Context
rdf:value: Primary antibodies were incubated overnight in block, and were goat anti-Vasa (1:400, Santa Cruz sc-26877), mouse anti-Fasciclin III (1:50, Developmental Studies Hybridoma Bank), rabbit anti-Zfh-1 (1:5000, Ruth Lehmann), rabbit anti-Stat>>23<< (1:1000, Erika Bach), mouse anti-HA (1:600, Covance), rabbit anti-GFP (1:1000, Molecular Probes), rabbit anti-alpha-spectrin (1:
n2:mentions: n3:20412771

Subject Item: _:vb8183516
rdf:type: n2:Context
rdf:value: To label cells in S phase, freshly-dissected testes were incubated for 30 minutes in 10μM Edu>>26<< then fixed. Testes were stained as above with other antibodies, followed by visualization of the Edu label with the Click-iT EdU Alexa Fluor 647 Kit (Invitrogen) according to the manufacturer’s instructions.
n2:mentions: n3:18272492

Subject Item: _:vb8183517
rdf:type: n2:Context
rdf:value: Mitotic germ cells were identified by centrosomin enrichment>>27<<, DNA compaction, and nuclear breakdown (revealed by loss of vasa exclusion from the nucleus).
n2:mentions: n3:8653793

Subject Item: _:vb8183518
rdf:type: n5:Section
dc:title: summary
n5:contains: _:vb8183519 _:vb8183520

Subject Item: _:vb8183519
rdf:type: n2:Context
rdf:value: We show here that the principle signaling pathway implicated in this niche, chemokine activation of STAT>>1<<,2, does not primarily regulate self-renewal of germline stem cells (GSCs), but rather governs GSC adhesion to hub cells.
n2:mentions: n3:11752575

Subject Item: _:vb8183520
rdf:type: n2:Context
rdf:value: We show here that the principle signaling pathway implicated in this niche, chemokine activation of STAT1,>>2<<, does not primarily regulate self-renewal of germline stem cells (GSCs), but rather governs GSC adhesion to hub cells.
n2:mentions: n3:11752574

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