Jackson Laboratory hhip lacz

(A-A’’) Simplified model of HH feedback inhibition. (A) Inactive HH signaling : PTCH1 suppresses SMO in the absence of HH ligand leading to production of GLI repressor (GLIR) and the suppression of HH-dependent gene expression. (A’) Active HH signaling : HH ligand binds to PTCH1, relieving SMO inhibition, and leading to GLI activator (GLIA) production and the initiation of HH target gene expression (including Gli1 , Ptch1 , and <t>Hhip</t> ). (A’’) HH feedback inhibition : HHIP protein sequesters HH ligand, allowing unbound PTCH1 to re-establish SMO repression, and the subsequent repression of HH pathway activity. (B-E) X-Gal staining of whole tongues from Gli1 <t>lacZ/+</t> , Gli2 lacZ /+ , Ptch1 lacZ /+ and Hhip lacZ /+ mice. (B) Gli1l acZ is expressed in all three taste papillae: fungiform (FP, circles) on the anterior tongue, foliate (FOL, bracket) and circumvallate (CV, box, B’) on the posterior tongue. The remaining tongue is covered by Gli1 lacZ negative non-taste filiform papillae (FILIF) on the anterior tongue and on the taste papilla-free intermolar eminence (B’’). The scale bars apply to all whole tongue images. (C-C’’) Gli2 lacZ is present in both taste (FP, CV and FOL) and non-taste FILIF papillae. (D-D’’) Ptch1 lacZ expression is apparent in FP, CV and FOL and in FILIF in intermolar eminence. (E-E’’) Hhip lacZ is expressed in non-taste FILIF in the anterior tongue and intermolar. Insets illustrate expression in FILIF surrounding FP (E) and in the intermolar eminence (E’’). Dotted lines demarcate intermolar eminence from anterior tongue.

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Journal: Developmental dynamics : an official publication of the American Association of Anatomists

Article Title: Hedgehog (HH) pathway endogenous antagonist HHIP: unique lingual expression in filiform papillae during homeostasis and ectopic in fungiform papillae during HH signaling inhibition

doi: 10.1002/dvdy.456

Figure Lengend Snippet: (A-A’’) Simplified model of HH feedback inhibition. (A) Inactive HH signaling : PTCH1 suppresses SMO in the absence of HH ligand leading to production of GLI repressor (GLIR) and the suppression of HH-dependent gene expression. (A’) Active HH signaling : HH ligand binds to PTCH1, relieving SMO inhibition, and leading to GLI activator (GLIA) production and the initiation of HH target gene expression (including Gli1 , Ptch1 , and Hhip ). (A’’) HH feedback inhibition : HHIP protein sequesters HH ligand, allowing unbound PTCH1 to re-establish SMO repression, and the subsequent repression of HH pathway activity. (B-E) X-Gal staining of whole tongues from Gli1 lacZ/+ , Gli2 lacZ /+ , Ptch1 lacZ /+ and Hhip lacZ /+ mice. (B) Gli1l acZ is expressed in all three taste papillae: fungiform (FP, circles) on the anterior tongue, foliate (FOL, bracket) and circumvallate (CV, box, B’) on the posterior tongue. The remaining tongue is covered by Gli1 lacZ negative non-taste filiform papillae (FILIF) on the anterior tongue and on the taste papilla-free intermolar eminence (B’’). The scale bars apply to all whole tongue images. (C-C’’) Gli2 lacZ is present in both taste (FP, CV and FOL) and non-taste FILIF papillae. (D-D’’) Ptch1 lacZ expression is apparent in FP, CV and FOL and in FILIF in intermolar eminence. (E-E’’) Hhip lacZ is expressed in non-taste FILIF in the anterior tongue and intermolar. Insets illustrate expression in FILIF surrounding FP (E) and in the intermolar eminence (E’’). Dotted lines demarcate intermolar eminence from anterior tongue.

Article Snippet: These strains were obtained from The Jackson Laboratory: Gli1 lacZ/+ (Stock No: 008211); Gli2 lacZ/+ (Stock No: 007922); Ptch1 lacZ/+ (Stock no: 00308); Hhip lacZ/+ (Stock No: 006241); Smo fl/fl or Smo tm2Amc/J (Stock No: 004526); Gli2 fl/fl (Stock No: 007926); tetO-Cre (Stock No: 006224); R26 M2rtTA/+ or B6.Cg-Gt(ROSA)26Sor tm1(rtTA*M2)Jae/J (Stock No: 006965). tetO-Gli2ΔC4 mice were generated using a mouse Gli2ΔC4 cDNA.

Techniques: Expressing, Inhibition, Gene Expression, Targeted Gene Expression, Activity Assay, Staining

(A-A’’) H&E staining of FP bracketed with FILIF (A, sagittal section), CV (A’, horizontal section), and FOL (A’’, horizontal section) in Control tongue to illustrate tissue morphology. The taste bud is indicated by an oval in the FP (A) and arrows in CV (A’) and FOL (A’’). (B-B’’) X-Gal staining (blue) in Hhip lacZ tongue sections indicates expression in FILIF only and not in FP (B), CV (B’) or FOL (B’’). (C-C’’) Immunostaining of HHIP (red) and taste bud cells (K8, green) in Control tongue demonstrates protein expression only in FILIF (C, arrow heads) similar to Hhip . Arrows mark taste buds in CV (C’) and FOL (C’’). Dotted lines outline the epithelium. Scale bars in A-A’’ apply to corresponding images in B-B’’ and C-C’’. Asterisk in C denotes non-specific surface staining.

Journal: Developmental dynamics : an official publication of the American Association of Anatomists

Article Title: Hedgehog (HH) pathway endogenous antagonist HHIP: unique lingual expression in filiform papillae during homeostasis and ectopic in fungiform papillae during HH signaling inhibition

doi: 10.1002/dvdy.456

Figure Lengend Snippet: (A-A’’) H&E staining of FP bracketed with FILIF (A, sagittal section), CV (A’, horizontal section), and FOL (A’’, horizontal section) in Control tongue to illustrate tissue morphology. The taste bud is indicated by an oval in the FP (A) and arrows in CV (A’) and FOL (A’’). (B-B’’) X-Gal staining (blue) in Hhip lacZ tongue sections indicates expression in FILIF only and not in FP (B), CV (B’) or FOL (B’’). (C-C’’) Immunostaining of HHIP (red) and taste bud cells (K8, green) in Control tongue demonstrates protein expression only in FILIF (C, arrow heads) similar to Hhip . Arrows mark taste buds in CV (C’) and FOL (C’’). Dotted lines outline the epithelium. Scale bars in A-A’’ apply to corresponding images in B-B’’ and C-C’’. Asterisk in C denotes non-specific surface staining.

Techniques: Staining, Control, Expressing, Immunostaining

(A) H&E staining of soft palate (coronal section) in Control tissue to illustrate morphology. (B) X-Gal staining from Hhip lacZ/+ reporter mouse indicates no expression in soft palate. Taste buds are labeled by K8 (red). Inset indicates positive lacZ expression in soft palate from Gli1 lacZ reporter mouse. (C) Antibody detection of endogenous HHIP (red) and taste bud cells (K8, green) in Control tongue. There is no HHIP expression in soft palate, similar to absence of Hhip gene expression. Taste buds are outlined with oval lines in A-C. Dotted lines in B and C outline the base of the epithelium. Scale bar in A applies to B and C.

Journal: Developmental dynamics : an official publication of the American Association of Anatomists

Article Title: Hedgehog (HH) pathway endogenous antagonist HHIP: unique lingual expression in filiform papillae during homeostasis and ectopic in fungiform papillae during HH signaling inhibition

doi: 10.1002/dvdy.456

Figure Lengend Snippet: (A) H&E staining of soft palate (coronal section) in Control tissue to illustrate morphology. (B) X-Gal staining from Hhip lacZ/+ reporter mouse indicates no expression in soft palate. Taste buds are labeled by K8 (red). Inset indicates positive lacZ expression in soft palate from Gli1 lacZ reporter mouse. (C) Antibody detection of endogenous HHIP (red) and taste bud cells (K8, green) in Control tongue. There is no HHIP expression in soft palate, similar to absence of Hhip gene expression. Taste buds are outlined with oval lines in A-C. Dotted lines in B and C outline the base of the epithelium. Scale bar in A applies to B and C.

Techniques: Expressing, Staining, Control, Labeling, Gene Expression

X-Gal staining of FP (A-E) and FILIF (A’-E’) papillae in sagittal sections from a lacZ negative Control mouse and Gli1 lacZ/+ , Gli2 lacZ /+ , Ptch1 lacZ /+ and Hhip lacZ /+ reporter mice. (A,A’) There is no X-Gal staining in Control tissue. Scale bars apply to all corresponding images from reporter mice. (B) Gli1 lacZ cells are observed in basal epithelial cells of the FP walls, in perigemmal and in stromal cells. (B’) No Gli1 expression is detected in FILIF. (C,C’) Gli2 lacZ is expressed throughout the entire basal epithelial and stromal cells of FP and FILIF. (D,D’) Ptch1 lacZ is expressed in FP basal epithelial cells, perigemmal and stromal cells, and in a subset of FILIF cells. (E,E’) Expression of Hhip lacZ is observed only in FILIF, in a subset of cells. There is no Hhip lacZ expression in FP cells. Black dotted lines outline the base of the epithelium.

Journal: Developmental dynamics : an official publication of the American Association of Anatomists

Article Title: Hedgehog (HH) pathway endogenous antagonist HHIP: unique lingual expression in filiform papillae during homeostasis and ectopic in fungiform papillae during HH signaling inhibition

doi: 10.1002/dvdy.456

Figure Lengend Snippet: X-Gal staining of FP (A-E) and FILIF (A’-E’) papillae in sagittal sections from a lacZ negative Control mouse and Gli1 lacZ/+ , Gli2 lacZ /+ , Ptch1 lacZ /+ and Hhip lacZ /+ reporter mice. (A,A’) There is no X-Gal staining in Control tissue. Scale bars apply to all corresponding images from reporter mice. (B) Gli1 lacZ cells are observed in basal epithelial cells of the FP walls, in perigemmal and in stromal cells. (B’) No Gli1 expression is detected in FILIF. (C,C’) Gli2 lacZ is expressed throughout the entire basal epithelial and stromal cells of FP and FILIF. (D,D’) Ptch1 lacZ is expressed in FP basal epithelial cells, perigemmal and stromal cells, and in a subset of FILIF cells. (E,E’) Expression of Hhip lacZ is observed only in FILIF, in a subset of cells. There is no Hhip lacZ expression in FP cells. Black dotted lines outline the base of the epithelium.

Techniques: Staining, Negative Control, Control, Expressing

(A-A’) H&E staining of filiform papillae (FILIF) in sagittal sections. Yellow dotted lines indicate location of Hhip in FILIF (A). FILIF-specific compartments are labeled (A’). Dotted lines demarcate anterior and posterior columns. Anterior column includes granular cell layers with keratohyalin granules. Between the FILIF is the inter-papilla buttress column. The FILIF are covered by thick cornified layers. (B) H&E: A restricted and distinctive pattern of Hhip in the anterior epithelial face of FILIF is illustrated (yellow dotted lines) based on the expression of gene and protein. Blue dotted lines demarcate the anterior and posterior FILIF columns. (B’) X-Gal: X-Gal staining (blue) from Hhip lacZ /+ tongue reveals expression in a subset of FILIF cells (outlined in yellow). Nuclear fast red is used to label all cells. (B’’) HHIP/DAPI: Immunostaining of HHIP (red) demonstrates an expression pattern and location that directly correspond to Hhip . DAPI stains cell nuclei. (C-F) X-Gal staining of FILIF from Gli1 lacZ/+ , Gli2 lacZ /+ , Ptch1 lacZ /+ and Hhip lacZ /+ reporter mice. (C) Gli1 lacZ cells are not detected in FILIF. (D) Gli2 lacZ cells are present in the basal layer of FILIF epithelium. Both Ptch1 lacZ (E) and Hhip lacZ (F) are observed in a subset of FILIF cells. (G-G’’) X-Gal staining in Ptch1 lacZ tongue followed by HHIP immunostaining (red, G’) reveals overlapping expression of Ptch1 and HHIP outlined with yellow dots (G’,G’’). Asterisks denote non-specific antibody staining after X-Gal reactions in G’. Black/white dotted lines outline the epithelium in all the images in B-G’’.

Journal: Developmental dynamics : an official publication of the American Association of Anatomists

Article Title: Hedgehog (HH) pathway endogenous antagonist HHIP: unique lingual expression in filiform papillae during homeostasis and ectopic in fungiform papillae during HH signaling inhibition

doi: 10.1002/dvdy.456

Figure Lengend Snippet: (A-A’) H&E staining of filiform papillae (FILIF) in sagittal sections. Yellow dotted lines indicate location of Hhip in FILIF (A). FILIF-specific compartments are labeled (A’). Dotted lines demarcate anterior and posterior columns. Anterior column includes granular cell layers with keratohyalin granules. Between the FILIF is the inter-papilla buttress column. The FILIF are covered by thick cornified layers. (B) H&E: A restricted and distinctive pattern of Hhip in the anterior epithelial face of FILIF is illustrated (yellow dotted lines) based on the expression of gene and protein. Blue dotted lines demarcate the anterior and posterior FILIF columns. (B’) X-Gal: X-Gal staining (blue) from Hhip lacZ /+ tongue reveals expression in a subset of FILIF cells (outlined in yellow). Nuclear fast red is used to label all cells. (B’’) HHIP/DAPI: Immunostaining of HHIP (red) demonstrates an expression pattern and location that directly correspond to Hhip . DAPI stains cell nuclei. (C-F) X-Gal staining of FILIF from Gli1 lacZ/+ , Gli2 lacZ /+ , Ptch1 lacZ /+ and Hhip lacZ /+ reporter mice. (C) Gli1 lacZ cells are not detected in FILIF. (D) Gli2 lacZ cells are present in the basal layer of FILIF epithelium. Both Ptch1 lacZ (E) and Hhip lacZ (F) are observed in a subset of FILIF cells. (G-G’’) X-Gal staining in Ptch1 lacZ tongue followed by HHIP immunostaining (red, G’) reveals overlapping expression of Ptch1 and HHIP outlined with yellow dots (G’,G’’). Asterisks denote non-specific antibody staining after X-Gal reactions in G’. Black/white dotted lines outline the epithelium in all the images in B-G’’.

Techniques: Staining, Labeling, Expressing, Immunostaining

(A-A’’) H&E staining of fungiform papilla (FP)/taste bud (TB, oval outline) to illustrate three categories observed during HH pathway inhibition (HPI): (A) I. Typical FP/TB, (A’) II. Atypical FP/TB and (A’’) III. Atypical FP/No TB. (B-E) HHIP immunostaining (red) and TB cells (K8, green) after pharmacologic or genetic HPI. (B-B’’) HPI targeting Smo: SMO inhibition by Sonidegib drug, and (C-C’’) epi- Smo deletion with epithelial conditional deletion of Smo (cSmoKO). In Vehicle and Control, HHIP is consistently observed only in filiform papillae (FILIF) (B,C). After HPI with Sonidegib, in addition to FILIF expression, ectopic HHIP is observed next to the TB remnant (B’, arrows, Atypical FP/TB) and at FP apex in the former TB-bearing location (B’’, arrow, Atypical FP/No TB). Parallel ectopic HHIP expression in Atypical FPs (Type II and III) was seen in cSmoKO (C-C’’). (D-E) Gli2 suppression models: (D-D’’) epi- Gli2 deletion with epithelial conditional deletion of Gli2 ( cGli2KO ), and (E-E’’) epi- Gli2 repression by epithelial conditional activation of repressor form of Gli2 ( cGli2 ΔC4). Similar expression patterns are observed in these HPI models compared to Smo inhibition or deletion. In Controls, HHIP expression is only in FILIF and not in Typical FP/TB while after Gli2 suppression, in Atypical FP/TB and Atypical FP/No TB, there is ectopic expression of HHIP in FP apical regions. In B–E’’, white dotted lines outline the base of the epithelium. Yellow dotted lines label the surface of epithelium. DAPI stains cell nuclei.

Journal: Developmental dynamics : an official publication of the American Association of Anatomists

Article Title: Hedgehog (HH) pathway endogenous antagonist HHIP: unique lingual expression in filiform papillae during homeostasis and ectopic in fungiform papillae during HH signaling inhibition

doi: 10.1002/dvdy.456

Figure Lengend Snippet: (A-A’’) H&E staining of fungiform papilla (FP)/taste bud (TB, oval outline) to illustrate three categories observed during HH pathway inhibition (HPI): (A) I. Typical FP/TB, (A’) II. Atypical FP/TB and (A’’) III. Atypical FP/No TB. (B-E) HHIP immunostaining (red) and TB cells (K8, green) after pharmacologic or genetic HPI. (B-B’’) HPI targeting Smo: SMO inhibition by Sonidegib drug, and (C-C’’) epi- Smo deletion with epithelial conditional deletion of Smo (cSmoKO). In Vehicle and Control, HHIP is consistently observed only in filiform papillae (FILIF) (B,C). After HPI with Sonidegib, in addition to FILIF expression, ectopic HHIP is observed next to the TB remnant (B’, arrows, Atypical FP/TB) and at FP apex in the former TB-bearing location (B’’, arrow, Atypical FP/No TB). Parallel ectopic HHIP expression in Atypical FPs (Type II and III) was seen in cSmoKO (C-C’’). (D-E) Gli2 suppression models: (D-D’’) epi- Gli2 deletion with epithelial conditional deletion of Gli2 ( cGli2KO ), and (E-E’’) epi- Gli2 repression by epithelial conditional activation of repressor form of Gli2 ( cGli2 ΔC4). Similar expression patterns are observed in these HPI models compared to Smo inhibition or deletion. In Controls, HHIP expression is only in FILIF and not in Typical FP/TB while after Gli2 suppression, in Atypical FP/TB and Atypical FP/No TB, there is ectopic expression of HHIP in FP apical regions. In B–E’’, white dotted lines outline the base of the epithelium. Yellow dotted lines label the surface of epithelium. DAPI stains cell nuclei.

Techniques: Expressing, Inhibition, Staining, Immunostaining, Control, Activation Assay

(A-C) Antibody detection of endogenous HHIP (red) and taste bud (TB) cells (K8, green) after 5d whole body Smo deletion. Three categories of FP/TB: Typical FP/TB, Atypical FP/TB and Atypical FP/No TB are assessed. Percentage numbers are quantification of the three FP/TB types in a half tongue (total FP = 28). (A’-C’) X-Gal staining in the same sections demonstrates elimination of Gli1 lacZ HH-responding cells from FP basal and perigemmal cell regions in all the three FP/TB types. HHIP expression in Atypical FP with or without TB (arrows) and not in Typical FP/TB suggests a requirement of both SHH and Gli1 reduction for ectopic HHIP expression. Asterisks in B denote non-specific staining post X-Gal reactions. Dotted lines outline the base of the epithelium. Scale bar in A applies to all images. (D-F) Antibody detection of endogenous HHIP (red) and TB cells (K8, green) after Vehicle and 16d sonidegib treatment followed by 14 days Recovery (discontinuation of drug). Percentage numbers are quantification of Typical FP/TB and Atypical FP/No TB during Recovery from previous findings 4 . There is no HHIP expression in Typical FP/TB in Vehicle (D) or after Recovery (E). Ectopic HHIP expression is apparent in Atypical FP/No TB even after stopping the drug treatment for 14 days (F). (G-I) X-Gal staining in Gli1 lacZ reporter mouse in Vehicle and Sonidegib treated Recovery group showed usual Gli1 lacZ expression, in FP basal, perigemmal and stromal cells, in Vehicle (G) and in the recovered Typical FP/TB (H). In Atypical FP/No TB, while Gli1 lacZ expression is not restored in FP epithelium (I) HHIP expression is maintained at the FP apex (F). Thus, if Gli1l acZ + cells are resumed during recovery the ectopic HHIP expression is simultaneously lost, and when there is no Gli1 lacZ activity HHIP is retained. Dotted lines outline the base of the epithelium. Yellow dotted lines label the surface of epithelium. Scale bar applies to all images.

Journal: Developmental dynamics : an official publication of the American Association of Anatomists

Article Title: Hedgehog (HH) pathway endogenous antagonist HHIP: unique lingual expression in filiform papillae during homeostasis and ectopic in fungiform papillae during HH signaling inhibition

doi: 10.1002/dvdy.456

Figure Lengend Snippet: (A-C) Antibody detection of endogenous HHIP (red) and taste bud (TB) cells (K8, green) after 5d whole body Smo deletion. Three categories of FP/TB: Typical FP/TB, Atypical FP/TB and Atypical FP/No TB are assessed. Percentage numbers are quantification of the three FP/TB types in a half tongue (total FP = 28). (A’-C’) X-Gal staining in the same sections demonstrates elimination of Gli1 lacZ HH-responding cells from FP basal and perigemmal cell regions in all the three FP/TB types. HHIP expression in Atypical FP with or without TB (arrows) and not in Typical FP/TB suggests a requirement of both SHH and Gli1 reduction for ectopic HHIP expression. Asterisks in B denote non-specific staining post X-Gal reactions. Dotted lines outline the base of the epithelium. Scale bar in A applies to all images. (D-F) Antibody detection of endogenous HHIP (red) and TB cells (K8, green) after Vehicle and 16d sonidegib treatment followed by 14 days Recovery (discontinuation of drug). Percentage numbers are quantification of Typical FP/TB and Atypical FP/No TB during Recovery from previous findings 4 . There is no HHIP expression in Typical FP/TB in Vehicle (D) or after Recovery (E). Ectopic HHIP expression is apparent in Atypical FP/No TB even after stopping the drug treatment for 14 days (F). (G-I) X-Gal staining in Gli1 lacZ reporter mouse in Vehicle and Sonidegib treated Recovery group showed usual Gli1 lacZ expression, in FP basal, perigemmal and stromal cells, in Vehicle (G) and in the recovered Typical FP/TB (H). In Atypical FP/No TB, while Gli1 lacZ expression is not restored in FP epithelium (I) HHIP expression is maintained at the FP apex (F). Thus, if Gli1l acZ + cells are resumed during recovery the ectopic HHIP expression is simultaneously lost, and when there is no Gli1 lacZ activity HHIP is retained. Dotted lines outline the base of the epithelium. Yellow dotted lines label the surface of epithelium. Scale bar applies to all images.

Techniques: Inhibition, Expressing, Staining, Activity Assay

(A-F) Immunostaining of the chorda tympani, CT (P2X3, green) and lingual, LN nerves (NF, green), and taste bud cells (K8, red) in Control (no nerve cut) side of tongue and in contralateral tongue after 21 days of CT/LN nerve cut. Three types of fungiform papilla (FP)/taste bud (TB) are assessed: I. Typical FP/TB, II. Atypical FP/TB and III. Atypical FP/No TB. In Control, in Typical FP/TB, the CT nerve innervates TB (A) while the LN nerve is widely distributed to FP epithelial cells (D). After nerve cut, in Atypical FP with or without the TB, the CT nerve is eliminated (B,C) and LN nerve fibers are much reduced (E,F). (D’-F’) X-Gal staining in Hhip lacZ/+ reporter demonstrates expression in FILIF (Control) in Typical FP/TB and an ectopic expression at/near the FP apical region after the CT/LN nerve cut in Atypical FP/TB and Atypical FP/No TB (arrows). (G-I) HHIP immunostaining (red) resembles Hhip expression in Control tongue and in contralateral tongue after CT/LN nerve cut (arrows). K8 labels TB cells (green). Yellow dotted lines label the surface of epithelium in A-I. White dotted lines outline the base of the epithelium in A-I. Scale bar in A applies to the images in B and C. (J) Percentage of Typical FP/TB, Atypical FP/TB and Atypical FP/No TB in Control and CT/LN nerve cut. (K) Percentage of total FP with ectopic Hhip expression in Hhip lacZ reporter, in Control and after CT/LN nerve cut. (L) Percentage of total FP with ectopic HHIP expression in Control and after CT/LN nerve cut. Statistical analysis is independent sample t-test (*p<0.001; **p<0.01). Numbers in parentheses are number of animals analyzed.

Journal: Developmental dynamics : an official publication of the American Association of Anatomists

Article Title: Hedgehog (HH) pathway endogenous antagonist HHIP: unique lingual expression in filiform papillae during homeostasis and ectopic in fungiform papillae during HH signaling inhibition

doi: 10.1002/dvdy.456

Figure Lengend Snippet: (A-F) Immunostaining of the chorda tympani, CT (P2X3, green) and lingual, LN nerves (NF, green), and taste bud cells (K8, red) in Control (no nerve cut) side of tongue and in contralateral tongue after 21 days of CT/LN nerve cut. Three types of fungiform papilla (FP)/taste bud (TB) are assessed: I. Typical FP/TB, II. Atypical FP/TB and III. Atypical FP/No TB. In Control, in Typical FP/TB, the CT nerve innervates TB (A) while the LN nerve is widely distributed to FP epithelial cells (D). After nerve cut, in Atypical FP with or without the TB, the CT nerve is eliminated (B,C) and LN nerve fibers are much reduced (E,F). (D’-F’) X-Gal staining in Hhip lacZ/+ reporter demonstrates expression in FILIF (Control) in Typical FP/TB and an ectopic expression at/near the FP apical region after the CT/LN nerve cut in Atypical FP/TB and Atypical FP/No TB (arrows). (G-I) HHIP immunostaining (red) resembles Hhip expression in Control tongue and in contralateral tongue after CT/LN nerve cut (arrows). K8 labels TB cells (green). Yellow dotted lines label the surface of epithelium in A-I. White dotted lines outline the base of the epithelium in A-I. Scale bar in A applies to the images in B and C. (J) Percentage of Typical FP/TB, Atypical FP/TB and Atypical FP/No TB in Control and CT/LN nerve cut. (K) Percentage of total FP with ectopic Hhip expression in Hhip lacZ reporter, in Control and after CT/LN nerve cut. (L) Percentage of total FP with ectopic HHIP expression in Control and after CT/LN nerve cut. Statistical analysis is independent sample t-test (*p<0.001; **p<0.01). Numbers in parentheses are number of animals analyzed.

Techniques: Expressing, Immunostaining, Control, Staining

(A) X-Gal staining from Gli1 lacZ/+ reporter in Control Typical FP/TB illustrates HH signaling in FP basal, perigemmal and stromal cells. (A’) HHIP immunostaining (red) in the same sections demonstrates expression in FILIF only (white arrows). (A’’) Merged images with TB cells (K8, green) indicate absence of HHIP and presence of Gli1 lacZ in Typical FP/TB. (B-E) After 21 days of chorda tympani (CT)/lingual (LN) nerve cut, X-Gal staining from Gli1 lacZ/+ reporter illustrates graded loss of HH signaling in FP basal and perigemmal cells. (B’-E’) HHIP immunostaining (red) in the same sections demonstrates an ectopic expression at FP apex in addition to the expression in FILIF (white arrows). (B’’-E’’) Merged images of Gli1 lacZ , HHIP and TB cells (K8, green) indicate opposing expression patterns of Gli1 lacZ and HHIP. (B-D) In Atypical FP/TB three different expression patterns of Gli1 lacZ and HHIP are observed. (B-B’’) Gli1 lacZ cells are retained in FP walls and there is no HHIP expression. (C-C’’) Gli1 lacZ cells are gradually lost from one side of FP wall and HHIP expression is observed at the FP apex on the same side where Gli1 lacZ is eliminated. (D-D’’) Gli1 lacZ cells are completely eliminated from FP walls and perigemmal cells, and HHIP ectopically expressed at both sides of FP apex bracketing the TB remnant. (E-E’’) In Atypical FP/No TB there are no Gli1 lacZ cells in FP and acquisition of HHIP in the entire FP apical area.

Journal: Developmental dynamics : an official publication of the American Association of Anatomists

Article Title: Hedgehog (HH) pathway endogenous antagonist HHIP: unique lingual expression in filiform papillae during homeostasis and ectopic in fungiform papillae during HH signaling inhibition

doi: 10.1002/dvdy.456

Figure Lengend Snippet: (A) X-Gal staining from Gli1 lacZ/+ reporter in Control Typical FP/TB illustrates HH signaling in FP basal, perigemmal and stromal cells. (A’) HHIP immunostaining (red) in the same sections demonstrates expression in FILIF only (white arrows). (A’’) Merged images with TB cells (K8, green) indicate absence of HHIP and presence of Gli1 lacZ in Typical FP/TB. (B-E) After 21 days of chorda tympani (CT)/lingual (LN) nerve cut, X-Gal staining from Gli1 lacZ/+ reporter illustrates graded loss of HH signaling in FP basal and perigemmal cells. (B’-E’) HHIP immunostaining (red) in the same sections demonstrates an ectopic expression at FP apex in addition to the expression in FILIF (white arrows). (B’’-E’’) Merged images of Gli1 lacZ , HHIP and TB cells (K8, green) indicate opposing expression patterns of Gli1 lacZ and HHIP. (B-D) In Atypical FP/TB three different expression patterns of Gli1 lacZ and HHIP are observed. (B-B’’) Gli1 lacZ cells are retained in FP walls and there is no HHIP expression. (C-C’’) Gli1 lacZ cells are gradually lost from one side of FP wall and HHIP expression is observed at the FP apex on the same side where Gli1 lacZ is eliminated. (D-D’’) Gli1 lacZ cells are completely eliminated from FP walls and perigemmal cells, and HHIP ectopically expressed at both sides of FP apex bracketing the TB remnant. (E-E’’) In Atypical FP/No TB there are no Gli1 lacZ cells in FP and acquisition of HHIP in the entire FP apical area.

Techniques: Staining, Control, Immunostaining, Expressing

(A) X-Gal staining in Ptch1 lacZ/+ mouse gavaged with Vehicle reveals expression in fungiform papilla (FP) basal, perigemmal and stromal cells and in a subset of filiform papillae (FILIF) cells. (A’) HHIP immunostaining (red) and TB cells (K8, green) in the same sections demonstrate HHIP expression in FILIF only (white arrow). (A’’) Merged images with Ptch1 lacZ indicate overlap with HHIP in FILIF. (B-D) After Sonidegib treatment for HH pathway inhibition, X-Gal staining from Ptch1 lacZ reporter demonstrates retained expression in FILIF while the expression in FP basal and perigemmal cells is lost. There is remaining Ptch1 lacZ expression in FP apical basal layer (Atypical FP/TB, black arrows) and an ectopic expression in basal and suprabasal layers at a former TB-bearing location (Atypical FP/No TB, black arrows). (B’-D’) HHIP immunostaining (red) and TB cells (K8, green) in the same sections demonstrate ectopic HHIP expression at FP apex (yellow dotted lines). (B’’-D’’) Merged images of Ptch1 lacZ , HHIP and K8 indicate partial co-expression pattern of Ptch1 lacZ and HHIP (yellow dotted lines). Black/white dotted lines outline the base of the epithelium. Asterisks denote non-specific staining post X-Gal reactions. (E) Percentage of total FP with Typical and Ectopic Ptch1 lacZ expression in Vehicle and after Sonidegib treatment. Statistical analysis is independent sample t-test. (*p<0.001). Numbers in parentheses are number of tongues analyzed.

Journal: Developmental dynamics : an official publication of the American Association of Anatomists

Article Title: Hedgehog (HH) pathway endogenous antagonist HHIP: unique lingual expression in filiform papillae during homeostasis and ectopic in fungiform papillae during HH signaling inhibition

doi: 10.1002/dvdy.456

Figure Lengend Snippet: (A) X-Gal staining in Ptch1 lacZ/+ mouse gavaged with Vehicle reveals expression in fungiform papilla (FP) basal, perigemmal and stromal cells and in a subset of filiform papillae (FILIF) cells. (A’) HHIP immunostaining (red) and TB cells (K8, green) in the same sections demonstrate HHIP expression in FILIF only (white arrow). (A’’) Merged images with Ptch1 lacZ indicate overlap with HHIP in FILIF. (B-D) After Sonidegib treatment for HH pathway inhibition, X-Gal staining from Ptch1 lacZ reporter demonstrates retained expression in FILIF while the expression in FP basal and perigemmal cells is lost. There is remaining Ptch1 lacZ expression in FP apical basal layer (Atypical FP/TB, black arrows) and an ectopic expression in basal and suprabasal layers at a former TB-bearing location (Atypical FP/No TB, black arrows). (B’-D’) HHIP immunostaining (red) and TB cells (K8, green) in the same sections demonstrate ectopic HHIP expression at FP apex (yellow dotted lines). (B’’-D’’) Merged images of Ptch1 lacZ , HHIP and K8 indicate partial co-expression pattern of Ptch1 lacZ and HHIP (yellow dotted lines). Black/white dotted lines outline the base of the epithelium. Asterisks denote non-specific staining post X-Gal reactions. (E) Percentage of total FP with Typical and Ectopic Ptch1 lacZ expression in Vehicle and after Sonidegib treatment. Statistical analysis is independent sample t-test. (*p<0.001). Numbers in parentheses are number of tongues analyzed.

Techniques: Expressing, Inhibition, Staining, Immunostaining

(A-C) HH signaling components in fungiform (FP) and filiform (FILIF) papilla. (A) Control: In Typical FP/TB, SHH ligand (green) is present with the TB basal cells. Gli1+ HH-responding cells (blue) are in FP basal epithelial and perigemmal cells. HHIP (red) is within FILIF only. Hedgehog pathway inhibition: (B) In Type II Atypical FP/TB there is reduction of SHH+ cells from TB, loss of Gli1 + cells in one side of the FP wall and associated ectopic HHIP expression at the FP apex on the same side as loss of Gli1+ cells. (C) When Gli1 + cells are completely eliminated from the FP walls along with loss of SHH in Type III Atypical FP/No TB, ectopic HHIP expression occurs in the FP apex in a location previously occupied by the TB. The FP has acquired a conical FILIF-like morphology. (D) In a proposed mechanism of action: In FP, SHH directly and/or indirectly via activating Gli1+ cells inhibits HHIP expression in the papilla. In FILIF, HHIP blocks expression of SHH and downstream Gli1 activity. (E) We propose that the SHH ligand is secreted along FP basal epithelial cells signaling to Gli1+ cells of the FP wall. Concurrently, HHIP from a FILIF cell subset is secreted along basal epithelial cells and inhibits HH signaling. At the rete peg region of the FP/FILIF epithelia, signaling via Gli1+ cells is inhibited.

Journal: Developmental dynamics : an official publication of the American Association of Anatomists

Article Title: Hedgehog (HH) pathway endogenous antagonist HHIP: unique lingual expression in filiform papillae during homeostasis and ectopic in fungiform papillae during HH signaling inhibition

doi: 10.1002/dvdy.456

Figure Lengend Snippet: (A-C) HH signaling components in fungiform (FP) and filiform (FILIF) papilla. (A) Control: In Typical FP/TB, SHH ligand (green) is present with the TB basal cells. Gli1+ HH-responding cells (blue) are in FP basal epithelial and perigemmal cells. HHIP (red) is within FILIF only. Hedgehog pathway inhibition: (B) In Type II Atypical FP/TB there is reduction of SHH+ cells from TB, loss of Gli1 + cells in one side of the FP wall and associated ectopic HHIP expression at the FP apex on the same side as loss of Gli1+ cells. (C) When Gli1 + cells are completely eliminated from the FP walls along with loss of SHH in Type III Atypical FP/No TB, ectopic HHIP expression occurs in the FP apex in a location previously occupied by the TB. The FP has acquired a conical FILIF-like morphology. (D) In a proposed mechanism of action: In FP, SHH directly and/or indirectly via activating Gli1+ cells inhibits HHIP expression in the papilla. In FILIF, HHIP blocks expression of SHH and downstream Gli1 activity. (E) We propose that the SHH ligand is secreted along FP basal epithelial cells signaling to Gli1+ cells of the FP wall. Concurrently, HHIP from a FILIF cell subset is secreted along basal epithelial cells and inhibits HH signaling. At the rete peg region of the FP/FILIF epithelia, signaling via Gli1+ cells is inhibited.

Techniques: Activity Assay, Expressing, Control, Inhibition

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