WormMine: AnatomyTerm WBbt:0005062

• Definition: male gonad cell, leads gonad during morphogenesis and initiates union with cloaca
• Name: linker cell
• Primary Identifier: WBbt:0005062
• Synonym: gon_male_link

2 Children

Definition	Name	Synonym	Primary Identifier
linker cell of the pedigree Z1.paa.	linker cell Z1		WBbt:0008239
linker cell of the pedigree Z4.aaa.	linker cell Z4		WBbt:0008240

0 Expression Clusters

42 Expression Patterns

• Primary Identifier: Expr4882
• Remark: Picture: Fig. 2C.
• Pattern: Although the signal was very faint, expression of the full-length mig-24::venus translational fusion construct in hermaphrodites, which fully rescues the mig-24 phenotype, was detected only in DTC nuclei. Expression of mig-24::venus was observed in males, where the signal was detected specifically in MLCs from L2 through L4 stages. Thus, MIG-24 is expressed specifically in gonadal leader cells both in hermaphrodites and males.
• Subcellular Localization: Expressed in nuclei.

• Primary Identifier: Expr4645
• Pattern: In the presence of the 6-kb promoter region, the vulval expression is identical to that of the 8-kb nhr-67::[Delta]pes-10 constructs (see Expr4642). Besides the previously reported expression in head neurons, expression was also observed in the anchor cell (AC) (during mid to late L3 stage) in hermaphrodites and the linker cell in males.

• Primary Identifier: Expr1330
• Pattern: In hermaphrodites, GFP is not observed in gonads from first-stage(L1) larvae. Its first gonadal expression occurs in L2 and is limited to the DTCs. GFP continues to be expressed in DTCs through L4, but is faint or not detectable in the adult. GFP is similarly expressed in the male linker cell. In addition to its expression in leader cells, the gon-1 promoter also drives GFP in muscle cells throughout development.

• Primary Identifier: Expr3278
• Pattern: In the embryo, the upstream promoter (ten-1a) is most active in the descendants of the C and EMS blastomers. During postembryonic development, GFP expression was detected in the pharynx, gut, coelomocytes, posterior body wall muscles, vulva muscles in hermaphrodites, and diagonal muscles in males. The ten-1a promoter is also active in some hypodermal cells including the hyp-11 cell, hypodermal seam cells, and rectal hypodermis. In the somatic gonad, it is active throughout its development starting with z1 and z4 cells in the embryo. During gonad development, it is expressed in the distal tip cells and the linker cell in males, in gonad and spermatheca sheath cells, and the utse cells of the uterus. In males, ten-1a is active in the vas deferens and spicule socket cells. Furthermore, GFP expression in DVB neurons and a few ring interneurons could be detected.

• Primary Identifier: Expr12030
• Pattern: Both YFP::LNKN-1 and LNKN-1::YFP are similarly localized to the plasma membrane of many cells. LNKN-1 begins to be expressed in all somatic gonadal cells of the male, including the LC, the vas deferens precursor cells, and seminal vesicle precursor cells, starting in the early L3 stage and continuing through adulthood. It is also expressed in all somatic gonadal cells of the hermaphrodite, including the distal tip cells, anchor cell, uterine precursor cells, and spermatheca precursor cells. Other expression occurs in pharynx, pharyngeal-intestinal valve, intestine, excretory cell and canal, seam cells, a specialized subset of hypodermal cells, the vulval precursor cells of the hermaphrodite, and hook precursor cells in the male. YFP-tagged LNKN-1 is localized to the plasma membrane, exhibiting stronger localization to the sides of cell-cell contact in tissues such as the intestine, seam, and gonad.

• Primary Identifier: Expr8158
• Remark: Lineage expression: B lineage. New Anatomy_term: hyp13. Picture: Figure 2, Figure 4B.
• Pattern: In males, dmd-3::YFP was expressed in a number of sexually dimorphic or sex-specific cells, including the tail tip, hindgut, B lineage, ray RnA neurons and somatic gonad. By contrast, hermaphrodites exhibited strong dmd-3::YFP expression only in the anchor cell. Non-sex-specific expression of these reporters was weak, occurring primarily in the body hypodermis. In addition, expression in phasmid neurons of both sexes was sometimes seen during L3 and L4. In hyp8 to 11, dmd-3::YFP expression was male specific and coincided with morphogenesis. Tail tip expression initiated in early-mid L4 males, first in hyp8, hyp9 and hyp11, and shortly thereafter in hyp10. Authors occasionally observed weak expression in hyp9 in late L3 males. Expression levels peaked during tail tip retraction and decreased rapidly upon its completion. dmd-3 was also expressed in hyp13. Importantly, dmd-3::YFP was not expressed in hermaphrodite hyp8 to 11 at any stage.

• Primary Identifier: Expr962
• Remark: This 4.6 kb fragment contains sufficient regulatory sequence to rescue cell migration and axon guidance phenotypes of unc-5 mutants when fused to an unc-5 cDNA (Hamelin et al., 1993; M.-W. S. and J.G.C., unpublished data).
• Pattern: Expression was observed in the hermaphrodite DTCs and MLCs as well as in all five classes of motorneurons that exhibit axon guidance defects in unc-5 mutants (DA, DB, DD, VD, AS). In addition, expression was observed in several classes of neurons in the head that are not visibly affected by unc-5 mutations. These included 12 sensory neurons of the OLQ, OLL and IL2 classes, and the interneurons RIA, RIH and ASE. Expression of lacZ or gfp transcriptional reporters in the DA, DB and DD embryonic motorneurons was first detected in late stage embryos at about the time when these neurons are known to extend pioneer axons along the epidermis (aka, hypodermis) to the dorsal side. Expression in the DA and DB neurons was transient, and only DD staining persisted into larval and adult stages. The VD motorneurons expressed intensely at this and at later times, while the AS neurons expressed faintly and sporadically.

• Primary Identifier: Expr1300
• Remark: No detailed description on cellular expression pattern at hermaphrodites.
• Pattern: LIN-29 accumulates in B cell progeny. In contrast to the LIN-29 accumulation pattern observed in wild-type hermaphrodites, LIN-29 is not detectable in the F, U, Y, or M blast cell nuclei, or in their progeny, in males. LIN-29 is detected in the progeny of B. LIN-29 is first detectable in B cell progeny during the L3 stage. Although all B.a and B.p progeny accumulate LIN-29, the LIN-29 accumulation signals appear weaker and transient in B.p progeny. The identification of these LIN-29-accumulating cells as B progeny is based on their size, shape, and relative position within the male tail and is further supported by laser microsurgery experiments and the examination of LIN-29-accumulation patterns in a mutant with defects in B cell specification. Elimination of the B cell by laser micro-surgery during the L1 stage reduced the number of LIN-29-accumulating nuclei in the male tail by 1015. LIN-29 appears to persist in B.a progeny in the adult stage. LIN-29 accumulates in the linker cell. The first male-specific LIN-29 accumulation is detected in the nucleus of the linker cell (LC) positioned at the tip of the growing end of the gonad. LIN-29 accumulates in the LC during the L3 stage, after the gonad arm has completed the 180 turn. LIN-29 remains detectable until the late L4 stage when its disappearance is presumably due to LC destruction. LIN-29 accumulates in the male tail seam. LIN-29 accumulates in the lateral body seam cell nuclei and in the SET nuclei in L4 stage wild-type males. LIN-29 accumulates in ventral cord nuclei. During the late L3 stage, five to seven nuclei that belong to the preanal ganglion accumulate LIN-29. Four preanal ganglion nuclei accumulates LIN-29 and were identified by position as the CA9, CP9, AS11, and VA11 neurons which are descended from P11.a. LIN-29 accumulation in the preanal ganglion cluster persists through adulthood. In late L4 stage and adult males, additional ventral cord nuclei anterior to the preanal ganglion accumulate LIN-29. The accumulation of LIN-29 in ventral cord neurons is not observed in hermaphrodites. Also in contrast to hermaphrodites, the ventral hypodermal nuclei positioned within the ventral cord do not contain detectable levels of LIN-29 in adult males.
• Subcellular Localization: nuclei

• Primary Identifier: Expr1034
• Remark: 240 minutes of development (author) = late cleavage stage embryo (curator) Lineage expression: Rn descandents.
• Pattern: Except for an earlier onset of GFP expression from both evpPRII.67 and evpPRII.75, all reporters displayed very similar expression patterns. Embryonic GFP expression reported by evpPRII.14 is first visible at approximately 240 minutes of development and is ubiquitous, a pattern that persists until hatching. At the beginning of each larval stage, dividing seam cells express mab-20::GFP, as do both daughter cells in males and hermaphrodites. After the anterior daughter fuses with surrounding hyp7, mab-20::GFP expression is downregulated or shut off in the both daughters. Other hypodermal expression is not evident in males or hermaphrodites. Other cells that express mab-20::GFP in hermaphrodites include the hermaphrodite-specific neurons at the L4 stage, vulva cells A to F throughout development, the migrating distal tip cells during the L4 stage, and several unidentified neurons within the nerve ring and ventral nerve cord. Expression in males include cells in the posterior ganglia, the migrating male linker cell beginning at the L3 to L4 transition, and the same nerve ring and ventral nerve cord expression observed in hermaphrodites. About 35 hours after hatching, mid-L3 males express mab-20::GFP in the 9 R(n) cells that give rise to the ray precursor clusters. Interior neuronal ganglia also express lower levels of mab-20::GFP at this stage. At 38-40 hours, when ray papillae are just visible, the 9 R(n)s and their descendents express mab-20::GFP. However, the underlying neural ganglia express the same level of GFP as the ray precursors at this stage. This pattern of expression continues until adulthood.

• Primary Identifier: Expr11152
• Pattern: yif-1 is expressed in the linker cell.

• Primary Identifier: Expr8941
• Remark: Picture: Fig 5G.
• Subcellular Localization: In the linker cells of L3 stage wild-type males, MIG-2::GFP was evenly distributed throughout the membrane, but during the L4 stage MIG-2 became polarized to the adherent, ventral side.

• Primary Identifier: Expr8940
• Remark: Picture: Fig. 3D. Reporter gene fusion type not specified.
• Pattern: Expressed by the male linker cell from the early L4 stage until LC death in the L4-to-adult molt (n=40/41)

• Primary Identifier: Expr8939
• Remark: Picture: Fig. 3A,B. Reporter gene fusion type not specified.
• Pattern: The unc-5::GFP reporter only becomes visible in linker cell during the turn in the L2-to-L3 molt. Levels of LC UNC-5 expression remained constant during the L3 and L4 stages.

• Primary Identifier: Expr13013
• Pattern: nob-1 is strongly expressed in the linker cell nucleus from the L3 larval stage onward. NOB-1 is thought to function as a posterior Hox gene, and is expressed in many tail cell nuclei.

• Primary Identifier: Expr879
• Pattern: Excepting body wall muscles and gonadal leaders, described below, no other cells were observed to express SP-GFP-hemicentin. Body wall muscles: Beginning in late embryogenesis (approx. 500 minutes), and continuing through adulthood, hemicentin is synthesized by lateral body wall muscle cells along the entire length of the animal, excepting cells six and eight near the nerve ring. Medial body wall muscle cells express little if any hemicentin. Gonadal leaders (hermaphrodite): Beginning soon after their genesis in late L1 stage, hermaphrodite distal-tip cells express hemicentin continuously throughout development. The anchor cell, or proximal leader, expresses hemicentin beginning in early L3 stage soon after cell commitment. SP-GFP-hemicentin diffuses throughout the utse syncytium at this time, but it is not known whether him-4 is transcribed in nuclei other than the anchor. In the adult, all expression within the utse syncytium ceases. Gonadal leaders (male): In the male, distal-tip cells never express hemicentin. The linker cell, or proximal leader, expresses hemicentin beginning at early L2 stage.

• Primary Identifier: Expr14766
• Pattern: EFA-6 is expressed in the linker cell and U.l/rp cells.

• Primary Identifier: Expr3843
• Remark: Reporter gene fusion type not specified.
• Pattern: Both transgenes qIs56 and qIs19 have similar GFP expression in the DTC, but qIs56 has brighter GFP expression in the ventral nerve cord than qIs19. The wild-type male linker cell expresses lag-2::GFP intensely.

• Primary Identifier: Expr12877
• Pattern: An 11-kb regulatory region upstream of the bar-1/b-catenin gene fused to GFP is not expressed in cloacal cells or in the trailing gonad but is strongly expressed in the linker cell.

• Primary Identifier: Expr12878
• Pattern: mig-5/Dishevelled::GFP reporter is expressed in the linker cell.

• Primary Identifier: Expr12879
• Pattern: lin-17/Frizzled::GFP reporter is expressed in the linker cell.

• Primary Identifier: Expr12880
• Pattern: wrm-1/b-catenin is expressed in the linker cell, as well as other cells.

• Primary Identifier: Expr12881
• Pattern: cul-3 is expressed in the linker cell.

• Primary Identifier: Expr12882
• Pattern: rbx-1 is expressed in the linker cell.

• Primary Identifier: Expr12883
• Pattern: let-70 promoter::let-70::GFP or let-70 promoter::GFP transgenes are expressed in the linker cell and that the translational fusion reporter is evenly distributed between the nucleus and cytoplasm. Importantly, the expression of neither reporter is constitutive. Rather, while GFP fluorescence is not detected during migration of the linker cell, it is induced 1-2 hr before obvious morphological features of cell death appear.

• Primary Identifier: Expr11142
• Pattern: glr-2 is expressed in the linker cell.

• Primary Identifier: Expr11143
• Pattern: acr-16 is expressed in the linker cell.

• Primary Identifier: Expr13014
• Pattern: An eor-1::GFP fusion transgene expressed under the control of endogenous eor-1 regulatory sequences shows constitutive linker cell expression, and EOR-1::GFP protein localizes to the linker cell nucleus, among other nuclei.

• Primary Identifier: Expr11144
• Pattern: arx-7 is expressed in the linker cell.

• Primary Identifier: Expr13015
• Pattern: The nhr-67::mcherry transgene is continuously expressed in the linker cell.

• Primary Identifier: Expr11145
• Remark: Strain CG308.
• Pattern: gar-3 is expressed in the linker cell.

0 Life Stages

2 Parents

Definition	Name	Synonym	Primary Identifier
gonad of a male animal, produces sperm.	male gonad		WBbt:0006794
cell of male gonad.	male gonadal cell	male somatic gonadal cell	WBbt:0008209