Sonic hedgehog protein

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exposed to a low concentration of SHH for a relatively short time and express the transcription factor Olig2. The expression of Olig2 rapidly expands in a dorsal direction concomitantly with the continuous dorsal extension of the SHH gradient over time. However, as the morphogenetic front of SHH ligand moves and begins to grow more concentrated, cells that are exposed to higher levels of the ligand respond by switching off Olig2 and turning on Nkx2.2, creating a sharp boundary between the cells expressing the transcription factor Nkx2.2 ventral to the cells expressing Olig2. It is in this way that each of the domains of the six progenitor cell populations are thought to be successively patterned throughout the neural tube by the SHH concentration gradient. Mutual inhibition between pairs of transcription factors expressed in neighboring domains contributes to the development of sharp boundaries; however, in some cases, inhibitory relationship has been found even between pairs of transcription factors from more distant domains. Particularly,

2862:(innermost of three germ layers) in the distal epithelium, where the embryonic lungs are developing. This suggests that SHH is partially responsible for the branching of the lungs. Further evidence of SHH's role in lung branching has been seen with qPCR. SHH expression occurs in the developing lungs around embryonic day 11 and is strongly expressed in the buds of the fetal lungs but low in the developing bronchi. Mice who are deficient in SHH can develop tracheoesophageal fistula (abnormal connection of the esophagus and trachea). Additionally, a double (SHH-/- ) knockout mouse model exhibited poor lung development. The lungs of the SHH double knockout failed to undergo lobation and branching (i.e., the abnormal lungs only developed one branch, compared to an extensively branched phenotype of the wildtype). 2841:—a signaling center—to provide positional information in both a lateral and planar signaling pattern in tooth development and regulation of tooth cusp growth. SHH in particular is needed for growth of epithelial cervical loops, where the outer and inner epitheliums join and form a reservoir for dental stem cells. After the primary enamel knots are apoptosed, the secondary enamel knots are formed. The secondary enamel knots secrete SHH in combination with other signaling molecules to thicken the oral ectoderm and begin patterning the complex shapes of the crown of a tooth during differentiation and mineralization. In a knockout gene model, absence of SHH is indicative of 2808:. Each of the ventral progenitor domains expresses a highly individualized combination of transcription factors—Nkx2.2, Olig2, Nkx6.1, Nkx6.2, Dbx1, Dbx2, Irx3, Pax6, and Pax7—that is regulated by the SHH gradient. These transcription factors are induced sequentially along the SHH concentration gradient with respect to the amount and time of exposure to SHH ligand. As each population of progenitor cells responds to the different levels of SHH protein, they begin to express a unique combination of transcription factors that leads to neuronal cell fate differentiation. This SHH-induced differential gene expression creates sharp boundaries between the discrete 3015: 2485: 323: 2473: 2461: 300: 222: 197: 7477: 574: 329: 228: 3025: 3035: 42: 2753:), and complex interactions between SHH and other molecules are possible. BMPs are suggested to play a critical role in determining the sensitivity of neural cell to SHH signaling. Evidence supporting this comes from studies using BMP inhibitors that ventralize the fate of the neural plate cell for a given SHH concentration. On the other hand, mutation in BMP antagonists (e.g., 2435:—produces SHH, which travels extracellularly to the ventral region of the neural tube and instructs those cells to form the floor plate. Another view of floor plate induction hypothesizes that some precursor cells located in the notochord are inserted into the neural plate before its formation, later giving rise to the floor plate. 2642:, which are expressed in the neural tube. Mice mutants for Gli1 show normal spinal cord development, suggesting that it is dispensable for mediating SHH activity. However, Gli2 mutant mice show abnormalities in the ventral spinal cord, with severe defects in the floor plate and ventral-most interneurons (V3). Gli3 2388:. Microphthalmia is a condition that affects the eyes, which results in small, underdeveloped tissues in one or both eyes. This can lead to issues ranging from a coloboma to a single small eye to the absence of eyes altogether. Holoprosencephaly is a condition most commonly caused by a mutation of the 2993:

gene might not be well received in a discussion of a serious disorder with a patient or their family. This controversy has largely died down, and the name is now generally seen as a humorous relic of the time before the rise of fast, cheap complete genome sequencing and standardized nomenclature. The

2815:

The spatial and temporal aspect of the progressive induction of genes and cell fates in the ventral neural tube is illustrated by the expression domains of two of the most well-characterized transcription factors, Olig2 and Nkx2.2. Early in development, the cells at the ventral midline have only been

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gene is mutated or absent, the protein Sonic Hedgehog cannot do its job properly. Sonic hedgehog contributes to cell growth, cell specification and formation, structuring and organization of the body plan. This protein functions as a vital morphogenic signaling molecule and plays an important role in

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is a specialized structure, located at the ventral midpoint of the neural tube. Evidence supporting the notochord as the signaling center comes from studies in which a second notochord is implanted near a neural tube in vivo, leading to the formation of an ectopic floor plate within the neural tube.

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gradient, which specifies the dorsal spinal cord. Higher concentrations of the SHH ligand are found in the most ventral aspects of the neural tube and notochord, while lower concentrations are found in the more dorsal regions of the neural tube. The SHH concentration gradient has been visualized in

2853:

Although SHH is most commonly associated with brain and limb digit development, it is also important in lung development. Studies using qPCR and knockouts have demonstrated that SHH contributes to embryonic lung development. The mammalian lung branching occurs in the epithelium of the developing

2836:

SHH plays an important role in organogenesis and, most importantly, craniofacial development. Being that SHH is a signaling molecule, it primarily works by diffusion along a concentration gradient, affecting cells in different manners. In early tooth development, SHH is released from the primary

2827:

SHH expression in the frontonasal ectodermal zone (FEZ), which is a signaling center that is responsible for the patterned development of the upper jaw, regulates craniofacial development mediating through the miR-199 family in the FEZ. Specifically, SHH-dependent signals from the brain regulate

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SHH is suggested to promote the activation function of Gli2 and inhibit repressive activity of Gli3. SHH also seems to promote the activation function of Gli3, but this activity is not strong enough. The graded concentration of SHH gives rise to graded activity of Gli 2 and Gli3, which promote

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SHH is important for regulating dermal adipogenesis by hair follicle transit-amplifying cells (HF-TACs). Specifically, SHH induces dermal angiogenesis by acting directly on adipocyte precursors and promoting their proliferation through their expression of the peroxisome proliferator-activated

68: 3162:

Marigo V, Roberts DJ, Lee SM, Tsukurov O, Levi T, Gastier JM, Epstein DJ, Gilbert DJ, Copeland NG, Seidman CE (July 1995). "Cloning, expression, and chromosomal location of SHH and IHH: two human homologues of the Drosophila segment polarity gene hedgehog".

5116:

Stone DM, Hynes M, Armanini M, Swanson TA, Gu Q, Johnson RL, Scott MP, Pennica D, Goddard A, Phillips H, Noll M, Hooper JE, de Sauvage F, Rosenthal A (November 1996). "The tumour-suppressor gene patched encodes a candidate receptor for Sonic hedgehog".

2893:, thereby preventing proliferation. Inhibiting the activity of Rb seems to allow cells to divide. Therefore, sonic hedgehog—identified as an important regulator of Rb—may also prove to be an important feature in regrowing hair cells after damage. 2379:

gene can cause malformation of components of these systems, which can result in major problems in the developing embryo. The brain and eyes, for example, can be significantly impacted by mutations in this gene and cause disorders such as

2034:, a failure of splitting in the cerebral hemispheres, as demonstrated in an experiment using SHH knock-out mice in which the forebrain midline failed to develop and instead only a single fused telencephalic vesicle resulted. 2989:, which can result in severe brain, skull and facial defects, causing a few clinicians and scientists to criticize the name on the grounds that it sounds too frivolous. It has been noted that mention of a mutation in a 5531:

Sasaki H, Nishizaki Y, Hui C, Nakafuku M, Kondoh H (September 1999). "Regulation of Gli2 and Gli3 activities by an amino-terminal repression domain: implication of Gli2 and Gli3 as primary mediators of Shh signaling".

2828:

genes of the miR-199 family with downregulations of the miR-199 genes increasing SHH expression and resulting in wider faces, while upregulations of the miR-199 genes decrease SHH expression resulting in narrow faces.

3487:

Echelard Y, Epstein DJ, St-Jacques B, Shen L, Mohler J, McMahon JA, McMahon AP (December 1993). "Sonic hedgehog, a member of a family of putative signaling molecules, is implicated in the regulation of CNS polarity".

3586: 336: 235: 3249:

Nanni L, Ming JE, Bocian M, Steinhaus K, Bianchi DW, Die-Smulders C, Giannotti A, Imaizumi K, Jones KL, Campo MD, Martin RA, Meinecke P, Pierpont ME, Robin NH, Young ID, Roessler E, Muenke M (December 1999).

7283:

Nanni L, Ming JE, Du Y, Hall RK, Aldred M, Bankier A, Muenke M (July 2001). "SHH mutation is associated with solitary median maxillary central incisor: a study of 13 patients and review of the literature".

2929:

in the ER. There, SHH undergoes autoprocessing to generate a 20 kDa N-terminal signaling domain (SHH-N) and a 25 kDa C-terminal domain with no known signaling role. The cleavage is catalysed by a

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studies show that incremental two- and threefold changes in SHH concentration give rise to motor neuron and different interneuronal subtypes as found in the ventral spinal cord. These incremental changes

5448:

Matise MP, Epstein DJ, Park HL, Platt KA, Joyner AL (August 1998). "Gli2 is required for induction of floor plate and adjacent cells, but not most ventral neurons in the mouse central nervous system".

2845:. However, SHH activates downstream molecules of Gli2 and Gli3. Mutant Gli2 and Gli3 embryos have abnormal development of incisors that are arrested in early tooth development as well as small molars. 3444:

Krauss S, Concordet JP, Ingham PW (December 1993). "A functionally conserved homolog of the Drosophila segment polarity gene hh is expressed in tissues with polarizing activity in zebrafish embryos".

4145:

Taylor MD, Northcott PA, Korshunov A, Remke M, Cho YJ, Clifford SC, Eberhart CG, Parsons DW, Rutkowski S, Gajjar A, Ellison DW, Lichter P, Gilbertson RJ, Pomeroy SL, Kool M, Pfister SM (April 2012).

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Sonic hedgehog still plays a role in differentiation, proliferation, and maintenance of adult tissues. Abnormal activation of SHH signaling in adult tissues has been implicated in various types of

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Park HL, Bai C, Platt KA, Matise MP, Beeghly A, Hui CC, Nakashima M, Joyner AL (April 2000). "Mouse Gli1 mutants are viable but have defects in SHH signaling in combination with a Gli2 mutation".

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A potential inhibitor of the Hedgehog signaling pathway has been found and dubbed "Robotnikinin"—after Sonic the Hedgehog's nemesis and the main antagonist of the Sonic the Hedgehog game series,

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It is thought that the SHH gradient works to elicit multiple different cell fates by a concentration- and time-dependent mechanism that induces a variety of transcription factors in the ventral

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respond to SHH gradient concentration. Depending upon the nature of their interaction with SHH, they are classified into two groups—class I and class II—and are composed of members from the

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is located on chromosome seven and initiates the production of Sonic Hedgehog protein. This protein sends short- and long-range signals to embryonic tissues to regulate development. If the

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Chiang C, Litingtung Y, Lee E, Young KE, Corden JL, Westphal H, Beachy PA (October 1996). "Cyclopia and defective axial patterning in mice lacking Sonic hedgehog gene function".

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of SHH-N. This modification is required for efficient signaling, resulting in a 30-fold increase in potency over the non-palmitylated form and is carried out by a member of the

5751:

Liem KF, Jessell TM, Briscoe J (November 2000). "Regulation of the neural patterning activity of sonic hedgehog by secreted BMP inhibitors expressed by notochord and somites".

6006:"Notochord-derived Shh concentrates in close association with the apically positioned basal body in neural target cells and forms a dynamic gradient during neural patterning" 2343:(RGC) axons at low concentrations and repels them at higher concentrations. The absence (non-expression) of SHH has been shown to control the growth of nascent hind limbs in 6295:

Bellusci S, Furuta Y, Rush MG, Henderson R, Winnier G, Hogan BL (January 1997). "Involvement of Sonic hedgehog (Shh) in mouse embryonic lung growth and morphogenesis".

5230:

Ericson J, Briscoe J, Rashbass P, van Heyningen V, Jessell TM (1997). "Graded sonic hedgehog signaling and the specification of cell fate in the ventral neural tube".

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ventral and dorsal neuronal subtypes in the ventral spinal cord. Evidence from Gli3 and SHH/Gli3 mutants show that SHH primarily regulates the spatial restriction of

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the neural tube of mice engineered to express a SHH::GFP fusion protein to show this graded distribution of SHH during the time of ventral neural tube patterning.

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SHH also induces other proteins with which it interacts, and these interactions can influence the sensitivity of a cell towards SHH. Hedgehog-interacting protein (

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Placzek M, Tessier-Lavigne M, Yamada T, Jessell T, Dodd J (November 1990). "Mesodermal control of neural cell identity: floor plate induction by the notochord".

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Martí E, Bumcrot DA, Takada R, McMahon AP (May 1995). "Requirement of 19K form of Sonic hedgehog for induction of distinct ventral cell types in CNS explants".

6883:

Stanton BZ, Peng LF, Maloof N, Nakai K, Wang X, Duffner JL, Taveras KM, Hyman JM, Lee SW, Koehler AN, Chen JK, Fox JL, Mandinova A, Schreiber SL (March 2009).

3252:"The mutational spectrum of the sonic hedgehog gene in holoprosencephaly: SHH mutations cause a significant proportion of autosomal dominant holoprosencephaly" 5667:

Pons S, Martí E (January 2000). "Sonic hedgehog synergizes with the extracellular matrix protein vitronectin to induce spinal motor neuron differentiation".

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gene affects several major organ systems, such as the nervous system, cardiovascular system, respiratory system and musculoskeletal system. Mutations in the

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against it shows similar phenotypes. SHH exerts its effects in a concentration-dependent manner, so that a high concentration of SHH results in a local

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Pepinsky RB, Zeng C, Wen D, Rayhorn P, Baker DP, Williams KP, Bixler SA, Ambrose CM, Garber EA, Miatkowski K, Taylor FR, Wang EA, Galdes A (May 1998).

2691:, and V0 interneurons (in ventral to dorsal order). These different progenitor domains are established by "communication" between different classes of 2305:

midline. The sonic hedgehog transcription pathway has also been linked to the formation of specific kinds of cancerous tumors, including the embryonic

1703: 4499: 4787:

van Straaten HW, Hekking JW, Thors F, Wiertz-Hoessels EL, Drukker J (October 1985). "Induction of an additional floor plate in the neural tube".

5488:

Litingtung Y, Chiang C (October 2000). "Specification of ventral neuron types is mediated by an antagonistic interaction between Shh and Gli3".

3716: 2574: 6055:"Boolean modelling reveals new regulatory connections between transcription factors orchestrating the development of the ventral spinal cord" 5837:"Floor plate and motor neuron induction by different concentrations of the amino-terminal cleavage product of sonic hedgehog autoproteolysis" 3144: 7090: 3126: 4098:"Shh and Gli3 regulate formation of the telencephalic-diencephalic junction and suppress an isthmus-like signaling source in the forebrain" 6235:

Hardcastle Z, Mo R, Hui CC, Sharpe PT (August 1998). "The Shh signalling pathway in tooth development: defects in Gli2 and Gli3 mutants".

2460: 3627: 5905:"Establishing and interpreting graded Sonic Hedgehog signaling during vertebrate neural tube patterning: the role of negative feedback" 1930: 2590:

from the signaling tissue (notochord and floor plate) which subsequently differentiates into different neuronal subtypes as it occurs

4605:

de Lussanet MH, Osse JW (2012). "An ancestral axial twist explains the contralateral forebain and the optic chiasm in vertebrates".

1937: 7510: 4012:

Dassule HR, Lewis P, Bei M, Maas R, McMahon AP (November 2000). "Sonic hedgehog regulates growth and morphogenesis of the tooth".

4247:"Paracrine sonic hedgehog signaling contributes significantly to acquired steroidogenesis in the prostate tumor microenvironment" 7297: 6783:

Porter JA, Young KE, Beachy PA (October 1996). "Cholesterol modification of hedgehog signaling proteins in animal development".

2726:, while class II proteins are activated at different thresholds of SHH delineating the dorsal limit of domains. Selective cross- 4663: 4245:

Lubik AA, Nouri M, Truong S, Ghaffari M, Adomat HH, Corey E, Cox ME, Li N, Guns ES, Yenki P, Pham S, Buttyan R (January 2017).

3002:," and "Sonic hedgehog" is largely avoided by using standardized abbreviations when speaking with patients and their families. 2484: 5616:

Chuang PT, McMahon AP (February 1999). "Vertebrate Hedgehog signalling modulated by induction of a Hedgehog-binding protein".

7456: 7357: 7164: 6279: 6170: 3621: 2096: 3109: 2546:. Lower concentration of SHH results in cellular proliferation and induction of various ventral neural cell types. Once the 7649: 2780:. In vertebrates, SHH signaling in the ventral portion of the neural tube is most notably responsible for the induction of 2168: 65: 7503: 7254:

Mullor JL, Sánchez P, Ruiz i Altaba A (December 2002). "Pathways and consequences: Hedgehog signaling in human disease".

5378:

Ruiz i Altaba A (June 1998). "Combinatorial Gli gene function in floor plate and neuronal inductions by Sonic hedgehog".

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Currie PD, Ingham PW (August 1996). "Induction of a specific muscle cell type by a hedgehog-like protein in zebrafish".

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Riddle RD, Johnson RL, Laufer E, Tabin C (December 1993). "Sonic hedgehog mediates the polarizing activity of the ZPA".

2018:

and the organization of the central nervous system, limbs, digits and many other parts of the body. Sonic hedgehog is a

129:, HHG1, HLP3, HPE3, MCOPCB5, SMMCI, TPT, TPTPS, sonic hedgehog, Sonic hedgehog, ShhNC, sonic hedgehog signaling molecule 7370:

Rediscovering biology: Unit 7 Genetics of development .. Expert interview transcripts interview with John Incardona PhD

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of BMP in the ventral neural tube. Interactions of SHH with Fgf and RA have not yet been studied in molecular detail.

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Lewis KE, Eisen JS (September 2001). "Hedgehog signaling is required for primary motoneuron induction in zebrafish".

2905:

SHH undergoes a series of processing steps before it is secreted from the cell. Newly synthesised SHH weighs 45

4544: 4339:"The morphogen sonic hedgehog is an axonal chemoattractant that collaborates with netrin-1 in midline axon guidance" 322: 5265:

Ericson J, Rashbass P, Schedl A, Brenner-Morton S, Kawakami A, van Heyningen V, Jessell TM, Briscoe J (July 1997).

6398:"FGF9 and SHH signaling coordinate lung growth and development through regulation of distinct mesenchymal domains" 2673:

is another protein that is induced by SHH; it acts as an obligate co-factor for SHH signaling in the neural tube.

2400:

gene and subsequent sonic hedgehog protein, earning it the distinction of being an essential gene to development.

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gene is a member of the hedgehog gene family with five variations of DNA sequence alterations or splice variants.

7674: 5788:"Noggin-mediated antagonism of BMP signaling is required for growth and patterning of the neural tube and somite" 4049:"Hedgehog signalling from the zona limitans intrathalamica orchestrates patterning of the zebrafish diencephalon" 3393:

Nüsslein-Volhard C, Wieschaus E (October 1980). "Mutations affecting segment number and polarity in Drosophila".

3113: 3092: 2995: 2650:-dependent manner, promoting dorsal neuronal subtypes. SHH mutant phenotypes can be rescued in a SHH/Gli3 double 299: 7033: 6740:

Ingham PW, Nakano Y, Seger C (June 2011). "Mechanisms and functions of Hedgehog signalling across the metazoa".

3742: 6644:"Hair follicles' transit-amplifying cells govern concurrent dermal adipocyte production through Sonic Hedgehog" 2166:, after his wife Betsy Wilder came home with a magazine containing an advert for the first game in the series, 2159: 2004: 1767: 17: 2084: 2193: 5704:"A homeodomain protein code specifies progenitor cell identity and neuronal fate in the ventral neural tube" 5567:

Persson M, Stamataki D, te Welscher P, Andersson E, Böse J, Rüther U, Ericson J, Briscoe J (November 2002).

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Dorus S, Anderson JR, Vallender EJ, Gilbert SL, Zhang L, Chemnick LG, Ryder OA, Li W, Lahn BT (July 2006).

6595:"Sonic hedgehog initiates cochlear hair cell regeneration through downregulation of retinoblastoma protein" 2269: 1238:

smoothened signaling pathway involved in regulation of cerebellar granule cell precursor cell proliferation

221: 196: 138: 6642:

Zhang B, Tsai PC, Gonzalez-Celeiro M, Chung O, Boumard B, Perdigoto CN, Ezhkova E, Hsu YC (October 2016).

4929:"Two critical periods of Sonic Hedgehog signaling required for the specification of motor neuron identity" 2722:

families. Class I proteins are repressed at different thresholds of SHH delineating ventral boundaries of

2119:

causes the embryos to be covered with denticles, i.e. small pointy projections resembling the spikes of a

7679: 4388:"Sonic hedgehog has a dual effect on the growth of retinal ganglion axons depending on its concentration" 2914: 2824:

expressed in V2 and more dorsal domains, although V3 and V2 are separated by a further domain termed MN.

2750: 2730:

interactions between class I and class II proteins give rise to five cardinal ventral neuronal subtypes.

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regulation of nodal signaling pathway involved in determination of lateral mesoderm left/right asymmetry

7530: 6114:"miR-199 family contributes to regulation of sonic hedgehog expression during craniofacial development" 2515: 335: 234: 7556: 7551: 7449: 7174:

Kim J, Kim P, Hui CC (May 2001). "The VACTERL association: lessons from the Sonic hedgehog pathway".

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is established, cells residing in this region will subsequently express SHH themselves, generating a

2150: 2144: 2073: 125: 5336: 328: 227: 7546: 4650:

Litingtung Y, Chiang C (October 2000). "Control of Shh activity and signaling in the neural tube".

2746: 2719: 2318: 1892: 1841: 146: 7375: 7669: 7566: 7118:"Sonic Hedgehog, a key development gene, experienced intensified molecular evolution in primates" 7064: 2322: 2079: 7495: 4693:

Placzek M (August 1995). "The role of the notochord and floor plate in inductive interactions".

1913: 1888: 1862: 1837: 5569:"Dorsal-ventral patterning of the spinal cord requires Gli3 transcriptional repressor activity" 5331: 5314:

Lum L, Beachy PA (June 2004). "The Hedgehog response network: sensors, switches, and routers".

2886: 2878: 2874: 2793: 2711: 2558: 2551: 2542:. This inhibition causes the floor plate to become thin compared to the lateral regions of the 2443: 2409: 2252: 2104: 5267:"Pax6 controls progenitor cell identity and neuronal fate in response to graded Shh signaling" 2757:) produces severe defects in the ventral-most characteristics of the spinal cord, followed by 3611: 2918: 2797: 2781: 2703: 2695: 2547: 2447: 2420: 2416: 2396:

and facial dysmorphia. Many systems and structures rely heavily on proper expression of the

2340: 7664: 7589: 7442: 6945: 6792: 6352: 6066: 5954:"Wnt signaling plays an essential role in neuronal specification of the dorsal spinal cord" 5625: 5323: 5181: 5126: 5036: 4985: 4880: 4741: 4448: 4303: 4196:"Pediatric medulloblastoma - update on molecular classification driving targeted therapies" 3830: 3661: 3402: 2606: 2217: 888:

positive regulation of epithelial cell proliferation involved in prostate gland development

210: 7485:: A POTENTIAL CATALYTIC SITE WITHIN THE AMINO-TERMINAL SIGNALLING DOMAIN OF SONIC HEDGEHOG 8: 7659: 7526: 4730:"The relationships between notochord and floor plate in vertebrate development revisited" 3872:

Herzog W, Zeng X, Lele Z, Sonntag C, Ting JW, Chang CY, Hammerschmidt M (February 2003).

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domains rather than being inductive, as SHH/Gli3 mutants show intermixing of cell types.

2647: 2643: 2610: 166: 6949: 6796: 6356: 6070: 5629: 5327: 5185: 5130: 5040: 4989: 4884: 4840: 4815: 4745: 4452: 4437:"Developmental basis for hind-limb loss in dolphins and origin of the cetacean bodyplan" 4307: 3834: 3665: 3406: 2654:. Gli proteins have a C-terminal activation domain and an N-terminal repressive domain. 2022:

that patterns the developing embryo using a concentration gradient characterized by the

7338: 7242: 7199: 7082: 7014: 6971: 6909: 6884: 6865: 6816: 6765: 6668: 6643: 6619: 6594: 6575: 6521: 6496: 6427: 6378: 6217: 6138: 6113: 6089: 6054: 6035: 5929: 5904: 5866: 5733: 5649: 5513: 5357: 5296: 5150: 5009: 4958: 4904: 4853: 4675: 4632: 4614: 4471: 4436: 4412: 4387: 4368: 4276: 4222: 4195: 4171: 4146: 4122: 4097: 4078: 3939: 3914: 3854: 3687: 3559: 3513: 3469: 3426: 3370: 3343: 3319: 3292: 3226: 3199: 2946: 2812:

of transcription factor expression, which ultimately patterns the ventral neural tube.

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that mediates signaling activities of the notochord and floor plate. Studies involving

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Roelink H, Porter JA, Chiang C, Tanabe Y, Chang DT, Beachy PA, Jessell TM (May 1995).

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molecule is added to the C-terminus of SHH-N. Thus, the C-terminal domain acts as an

2842: 2535: 2419:. One of the most characterized functions of SHH is its role in the induction of the 2385: 2298: 2031: 2026:. This model has a non-uniform distribution of SHH molecules which governs different 2023: 118: 58: 7203: 7086: 6975: 6869: 6820: 6769: 6579: 6431: 6382: 6221: 6053:

Lovrics A, Gao Y, Juhász B, Bock I, Byrne HM, Dinnyés A, Kovács KA (November 2014).

6039: 5870: 5737: 5517: 5361: 5300: 4962: 4857: 4372: 4082: 3691: 3473: 3014: 2909:

and is referred to as the preproprotein. As a secreted protein, it contains a short

2769:

The concentration- and time-dependent, cell-fate-determining activity of SHH in the

2526:. On the other hand, mice mutants for SHH lack ventral spinal cord characteristics. 2408:

The sonic hedgehog (SHH) signaling molecule assumes various roles in patterning the

2317:

tumours. For SHH to be expressed in the developing embryo limbs, a morphogen called

1118:

positive regulation of mesenchymal cell proliferation involved in ureter development

7364: 7342: 7322: 7293: 7263: 7246: 7224: 7183: 7129: 7072: 6998: 6953: 6904: 6896: 6847: 6800: 6749: 6712: 6704: 6663: 6655: 6614: 6606: 6557: 6516: 6508: 6454: 6409: 6360: 6304: 6244: 6199: 6133: 6125: 6084: 6074: 6017: 5973: 5965: 5924: 5916: 5848: 5807: 5799: 5760: 5715: 5676: 5633: 5588: 5580: 5541: 5497: 5457: 5422: 5387: 5341: 5278: 5239: 5199: 5189: 5154: 5134: 5088: 5044: 5013: 4993: 4940: 4908: 4888: 4835: 4827: 4759: 4749: 4702: 4659: 4636: 4624: 4466: 4456: 4407: 4403: 4399: 4350: 4311: 4280: 4258: 4217: 4207: 4166: 4158: 4117: 4109: 4060: 4021: 3971: 3934: 3930: 3926: 3885: 3858: 3838: 3669: 3563: 3543: 3517: 3497: 3453: 3430: 3410: 3365: 3355: 3314: 3304: 3263: 3221: 3211: 3172: 2926: 2906: 2805: 2754: 2699: 2688: 2677: 2499: 2274: 1398:

regulation of mesenchymal cell proliferation involved in prostate gland development

415: 346: 290: 245: 2885:

that normally cannot return to a proliferative state to divide and differentiate.

1353:

regulation of epithelial cell proliferation involved in prostate gland development

150: 6804: 6079: 3874:"Adenohypophysis formation in the zebrafish and its dependence on sonic hedgehog" 3216: 2738: 2723: 2684: 2659: 2339:

at the ventral midline of the developing spinal cord. Specifically, SHH attracts

2314: 2310: 2282: 2100: 390: 174: 5920: 5243: 5093: 5076: 4113: 3344:"Sonic Hedgehog Signaling in Organogenesis, Tumors, and Tumor Microenvironments" 3149:

National Center for Biotechnology Information, U.S. National Library of Medicine

3131:

National Center for Biotechnology Information, U.S. National Library of Medicine

898:

negative regulation of proteasomal ubiquitin-dependent protein catabolic process

6610: 5786:

McMahon JA, Takada S, Zimmerman LB, Fan CM, Harland RM, McMahon AP (May 1998).

5174:

Proceedings of the National Academy of Sciences of the United States of America

4734:

Proceedings of the National Academy of Sciences of the United States of America

4441:

Proceedings of the National Academy of Sciences of the United States of America

2910: 2809: 2587: 2381: 2278: 2197: 2163: 2132: 2092: 2088: 1323:

mesenchymal smoothened signaling pathway involved in prostate gland development

843:

negative regulation of transcription elongation from RNA polymerase II promoter

491: 7381: 7326: 6885:"A small molecule that binds Hedgehog and blocks its signaling in human cells" 4435:

Thewissen JG, Cohn MJ, Stevens LS, Bajpai S, Heyning J, Horton WE (May 2006).

4162: 1418:

metanephric mesenchymal cell proliferation involved in metanephros development

7643: 6852: 6835: 6512: 4754: 4628: 3765: 3607: 3309: 2974: 2950: 2785: 2742: 2329: 2128: 2042: 2015: 2011: 602: 6546:"Regulation of early lung morphogenesis: questions, facts and controversies" 6248: 5764: 5545: 5461: 5391: 5345: 5194: 5048: 4575: 4461: 4212: 4025: 3975: 3291:

Blaess S, Szabó N, Haddad-Tóvolli R, Zhou X, Álvarez-Bolado G (2015-01-06).

3268: 3251: 551: 429: 7334: 7305: 7275: 7238: 7195: 7143: 7010: 6967: 6918: 6761: 6677: 6659: 6628: 6571: 6530: 6476: 6423: 6213: 6147: 6098: 6031: 5987: 5938: 5803: 5772: 5729: 5688: 5645: 5602: 5553: 5509: 5434: 5426: 5353: 5102: 4849: 4671: 4480: 4421: 4364: 4272: 4231: 4180: 4131: 4074: 4033: 3983: 3948: 3899: 3683: 3379: 3328: 3277: 3235: 3176: 2523: 2519: 2243:

has been found to have the most critical roles in development, acting as a

2054: 2050: 408: 187: 6861: 6812: 6726: 6708: 6374: 6316: 6256: 5862: 5821: 5680: 5469: 5399: 5292: 5251: 5213: 5146: 5056: 5005: 4954: 4900: 4800: 4773: 4714: 4536: 4323: 3850: 3555: 3509: 3465: 3422: 3184: 1977: 1972: 573: 7229: 7212: 7134: 7117: 7077: 7002: 6900: 6836:"Identification of a palmitic acid-modified form of human Sonic hedgehog" 6308: 5077:"The mechanisms of dorsoventral patterning in the vertebrate neural tube" 4337:

Charron F, Stein E, Jeong J, McMahon AP, Tessier-Lavigne M (April 2003).

3360: 2943: 2935: 2838: 2773: 2680: 2670: 2602: 2543: 2439: 2424: 2333: 2256: 2173: 2046: 1812: 1793: 6341:"Sonic hedgehog regulates branching morphogenesis in the mammalian lung" 4927:

Ericson J, Morton S, Kawakami A, Roelink H, Jessell TM (November 1996).

2737:

exerting an effect on the developing neural tube. Many other molecules,

7579: 7382:

GeneReviews/NCBI/NIH/UW entry on Anophthalmia / Microphthalmia Overview

7298:

10.1002/1096-8628(20010722)102:1<1::AID-AJMG1336>3.0.CO;2-U

6022: 6005: 5969: 5584: 5170:"Regulation of patched by sonic hedgehog in the developing neural tube" 4831: 4786: 3293:"Sonic hedgehog signaling in the development of the mouse hypothalamus" 3051: 2890: 2676:

There are five distinct progenitor domains in the ventral neural tube:

2413: 2306: 1358:

epithelial-mesenchymal signaling involved in prostate gland development

307: 204: 154: 6562: 6545: 6414: 6397: 6204: 6187: 4664:

10.1002/1097-0177(2000)9999:9999<::AID-DVDY1050>3.0.CO;2-Q

4263: 4246: 4065: 4048: 3064: 2669:) is induced by SHH, which in turn attenuates its signaling activity. 2371:

the formation of many different structures in developing embryos. The

1443:

epithelial cell proliferation involved in salivary gland morphogenesis

1133:

positive regulation of cerebellar granule cell precursor proliferation

95: 91: 87: 7372:.. explanation of the discovery and naming of the sonic hedgehog gene 6129: 6112:

Richbourg HA, Hu DP, Xu Y, Barczak AJ, Marcucio RS (September 2020).

5138: 4997: 4892: 3842: 3414: 2922: 2882: 2871: 2789: 2777: 2727: 2707: 2428: 2244: 2116: 2027: 2019: 1667: 374: 361: 273: 260: 162: 6958: 6933: 6753: 6693:"Proteolytic processing yields two secreted forms of sonic hedgehog" 5566: 3674: 3649: 2925:. Once translocation is complete, the signal sequence is removed by 2162:. The gene was named by Robert Riddle, a postdoctoral fellow at the 6593:

Lu N, Chen Y, Wang Z, Chen G, Lin Q, Chen ZY, Li H (January 2013).

6497:"Sonic hedgehog signaling in the lung. From development to disease" 6004:

Chamberlain CE, Jeong J, Guo C, Allen BL, McMahon AP (March 2008).

5264: 5229: 5026: 2954: 2931: 2859: 2855: 2796:

that spans the dorso-ventral axis and is antagonized by an inverse

2692: 2531: 2432: 2344: 2290: 2264: 2120: 1961: 7386: 6396:

White AC, Xu J, Yin Y, Smith C, Schmid G, Ornitz DM (April 2006).

6188:"Epithelial-mesenchymal signalling regulating tooth morphogenesis" 5952:

Muroyama Y, Fujihara M, Ikeya M, Kondoh H, Takada S (March 2002).

5637: 5501: 4619: 2994:

problem of the "inappropriateness" of the names of genes such as "

2792:

and ventral floor plate of the developing neural tube to create a

1578:

CD4-positive or CD8-positive, alpha-beta T cell lineage commitment

7525: 7428: 7418: 7407: 7397: 3915:"Patterning the dorsal telencephalon: a role for sonic hedgehog?" 3055: 3024: 2770: 2630:. In vertebrates, three different Gli proteins are present, viz. 2562: 2352: 2302: 2213: 1779: 1734: 1652: 1648: 7313:

Williams JA (December 2005). "Hedgehog and spinal cord injury".

7213:"Shh signaling and pancreatic cancer: implications for therapy?" 3034: 1493:

positive regulation of ureter smooth muscle cell differentiation

1488:

negative regulation of ureter smooth muscle cell differentiation

1468:

negative regulation of kidney smooth muscle cell differentiation

793:

positive regulation of kidney smooth muscle cell differentiation

142: 41: 7538: 6163:

Ten Cate's Oral Histology: Development, Structure, and Function

4147:"Molecular subgroups of medulloblastoma: the current consensus" 2939: 2889:

suppress cell growth by preventing cells from returning to the

2817: 2651: 2594:. Graded SHH signaling is suggested to be mediated through the 2570: 2038: 1945: 1689: 582: 7434: 6641: 1408:

positive regulation of hh target transcription factor activity

788:

positive regulation of immature T cell proliferation in thymus

7574: 7115: 4144: 3486: 3290: 3060: 2958: 2348: 2336: 2030:

according to concentration. Mutations in this gene can cause

7413:

Overview of all the structural information available in the

7392:

Overview of all the structural information available in the

5701: 4926: 3579:"Biologists Find Key Genes That Shape Patterning of Embryos" 3018:

SHH gradient and Gli activity in the vertebrate neural tube.

7628: 7623: 7607: 7602: 7597: 6495:

Kugler MC, Joyner AL, Loomis CA, Munger JS (January 2015).

6294: 5951: 2999: 2962: 2821: 2715: 2666: 2639: 2635: 2631: 2598:

family of proteins, which are vertebrate homologues of the

2595: 2403: 2260: 2000: 1403:

mesenchymal cell proliferation involved in lung development

893:

positive regulation of striated muscle cell differentiation

7253: 7034:"Sonic Hedgehog, DICER, and the Problem With Naming Genes" 6501:

American Journal of Respiratory Cell and Molecular Biology

5530: 4434: 4194:

DeSouza RM, Jones BR, Lowis SP, Kurian KM (22 July 2014).

1028:

negative regulation of dopaminergic neuron differentiation

6989:

Cohen MM (July 2006). "Problems in the naming of genes".

6445:

Miura T (2008). "Modeling lung branching morphogenesis".

5834: 5785: 5115: 4870: 4727: 4294:

Riddle RD, Tabin C (February 1999). "How limbs develop".

3392: 1608:

positive regulation of transcription by RNA polymerase II

1313:

determination of left/right asymmetry in lateral mesoderm

1253:

negative regulation of transcription by RNA polymerase II

953:

negative regulation of mesenchymal cell apoptotic process

783:

positive regulation of skeletal muscle tissue development

738:

positive regulation of skeletal muscle cell proliferation

6494: 6003: 5702:

Briscoe J, Pierani A, Jessell TM, Ericson J (May 2000).

4975: 4728:

Teillet MA, Lapointe F, Le Douarin NM (September 1998).

4336: 4244: 3533: 3248: 3161: 1598:

epithelial tube branching involved in lung morphogenesis

1173:

positive regulation of alpha-beta T cell differentiation

1038:

negative regulation of alpha-beta T cell differentiation

803:

anatomical structure formation involved in morphogenesis

398: 4193: 4046: 2010:

This signaling molecule is key in regulating embryonic

1208:

positive regulation of T cell differentiation in thymus

7159:(6th ed.). Sunderland, Mass: Sinauer Associates. 6934:"Humour of gene names lost in translation to patients" 6882: 6833: 6274:(5th ed.). Oxford University Press. p. 500. 5447: 4816:"The role of Sonic hedgehog in neural tube patterning" 2980: 2881:

activity in rat cochlea, sonic hedgehog allows mature

2561:, there is indirect evidence via the visualization of 2003:. The protein is named after the video game character 1188:

positive regulation of oligodendrocyte differentiation

938:

negative regulation of canonical Wnt signaling pathway

6339:

Pepicelli CV, Lewis PM, McMahon AP (September 1998).

6338: 6234: 4001:(5th ed.). Oxford University Press. p. 500. 3871: 3443: 2934:

within the C-terminal domain. During the reaction, a

878:

positive regulation of mesenchymal cell proliferation

563: 6690: 6111: 6052: 4047:

Scholpp S, Wolf O, Brand M, Lumsden A (March 2006).

3197: 1548:

positive regulation of cell population proliferation

6599:

Biochemical and Biophysical Research Communications

5232:

Cold Spring Harbor Symposia on Quantitative Biology

4011: 2466:

SHH and BMP gradients in the vertebrate neural tube

1068:

positive regulation of transcription, DNA-templated

988:

positive regulation of smoothened signaling pathway

7152: 5750: 5412: 3796: 3778: 3105: 3103: 3101: 3084: 3082: 3080: 2865: 2820:expressed in the V3 domain is reported to inhibit 2289:, located on the posterior side of the embryonic 2285:depends on the secretion of sonic hedgehog by the 2247:involved in patterning many systems—including the 1148:positive regulation of protein import into nucleus 1113:branching involved in prostate gland morphogenesis 948:branching involved in salivary gland morphogenesis 6782: 6739: 6334: 6332: 6330: 6328: 6326: 2985:The gene has been linked to a condition known as 2733:It is important to note that SHH is not the only 1543:polarity specification of anterior/posterior axis 345: 244: 7641: 5487: 4649: 2858:and lungs. SHH expressed throughout the foregut 2328:Sonic hedgehog has also been shown to act as an 1128:branching involved in blood vessel morphogenesis 1093:branching involved in ureteric bud morphogenesis 1008:negative regulation of protein catabolic process 7282: 6691:Bumcrot DA, Takada R, McMahon AP (April 1995). 6395: 5483: 5481: 5479: 5070: 5068: 5066: 4604: 3200:"Getting the measure of positional information" 3098: 3077: 1478:cerebellar granule cell precursor proliferation 958:positive regulation of neuroblast proliferation 6592: 6490: 6488: 6486: 6323: 5999: 5997: 5373: 5371: 5225: 5223: 4922: 4920: 4918: 4385: 3529: 3527: 3110:GRCm38: Ensembl release 89: ENSMUSG00000002633 2913:at its N-terminus, which is recognised by the 2557:Although there is no direct evidence of a SHH 2154:, were named for species of hedgehogs, while 7511: 7450: 6288: 5615: 5377: 4813: 4695:Current Opinion in Genetics & Development 2332:. It has been demonstrated that SHH attracts 2293:. Mutations in the human sonic hedgehog gene 1368:branching morphogenesis of an epithelial tube 1273:regulation of protein localization to nucleus 1043:positive regulation of sclerotome development 6991:American Journal of Medical Genetics. Part A 6982: 6925: 6876: 6537: 6179: 6046: 5902: 5744: 5695: 5660: 5609: 5560: 5524: 5476: 5406: 5307: 5063: 4643: 4598: 4330: 3820: 3341: 2963:Protein-cysteine N-palmitoyltransferase HHAT 2953:, is added to the alpha-amine of N-terminal 2870:Sonic hedgehog may play a role in mammalian 2301:type 3 HPE3, as a result of the loss of the 2099:in 1995 along with developmental geneticist 1628:cell proliferation in external granule layer 1073:positive regulation of Wnt signaling pathway 798:negative regulation of Wnt signaling pathway 7210: 6483: 6228: 5994: 5779: 5441: 5368: 5258: 5220: 5167: 5161: 5109: 5074: 5020: 4969: 4915: 4864: 4780: 4721: 4686: 4293: 3524: 3480: 3437: 3386: 3348:International Journal of Molecular Sciences 3089:GRCh38: Ensembl release 89: ENSG00000164690 1528:regulation of cell population proliferation 1108:positive regulation of cell differentiation 1098:negative regulation of T cell proliferation 1023:negative regulation of cell differentiation 7518: 7504: 7457: 7443: 6389: 6165:(8th ed.). St. Louis, Mo.: Elsevier. 5909:Cold Spring Harbor Perspectives in Biology 4138: 3961: 3703: 3701: 3342:Jeng KS, Chang CF, Lin SS (January 2020). 3063:, an oncogene which was originally named " 2423:and diverse ventral cell types within the 983:regulation of transcription, DNA-templated 502:transitional epithelium of urinary bladder 7228: 7173: 7133: 7076: 6957: 6908: 6851: 6716: 6667: 6618: 6561: 6543: 6520: 6413: 6364: 6203: 6137: 6088: 6078: 6021: 5977: 5928: 5852: 5811: 5719: 5666: 5592: 5335: 5313: 5282: 5203: 5193: 5092: 4944: 4839: 4763: 4753: 4618: 4531: 4529: 4527: 4525: 4494: 4492: 4490: 4470: 4460: 4428: 4411: 4354: 4262: 4221: 4211: 4170: 4121: 4095: 4064: 3990: 3938: 3912: 3889: 3736: 3673: 3369: 3359: 3318: 3308: 3267: 3225: 3215: 3198:Jaeger J, Martinez-Arias A (March 2009). 2764: 2158:was named after the video game character 2095:, which led to the researchers winning a 1638:regulation of signaling receptor activity 1343:negative regulation of cholesterol efflux 7312: 6827: 6185: 5898: 5896: 5894: 5892: 5890: 5888: 5886: 5884: 5882: 5880: 4789:Acta Morphologica Neerlando-Scandinavica 4386:Kolpak A, Zhang J, Bao ZZ (March 2005). 3647: 3033: 3023: 3013: 2442:itself is the initial groundwork of the 2404:Patterning of the central nervous system 2392:gene that causes improper separation or 2077:) was first identified in the fruit fly 1613:anterior/posterior pattern specification 1248:negative regulation of apoptotic process 1168:spinal cord motor neuron differentiation 1058:lymphoid progenitor cell differentiation 873:odontogenesis of dentin-containing tooth 763:bud outgrowth involved in lung branching 7376:‘Sonic Hedgehog’ sounded funny at first 7150: 6931: 6684: 6586: 6447:Current Topics in Developmental Biology 6269: 5828: 4692: 4570: 4568: 4566: 4564: 4562: 4040: 4005: 3996: 3739:"Cliff Tabin: Super Sonic An Interview" 3698: 3570: 2942:and a cholesterol transferase. Another 2569:gene expression, which encodes for the 2066: 903:embryonic digestive tract morphogenesis 14: 7642: 7427:(Mouse Sonic hedgehog protein) at the 7406:(Human Sonic hedgehog protein) at the 7378:.. New York Times November 12, 2006 .. 7093:from the original on December 31, 2020 7062: 7044:from the original on November 12, 2020 6776: 4522: 4487: 4379: 3865: 3814: 3806:. University of Oregon. Archived from 3788:. University of Oregon. Archived from 3774:. Interviewed by Kat Arney. BBC Radio. 3606: 3576: 1593:lung-associated mesenchyme development 1523:positive regulation of gene expression 1053:negative regulation of gene expression 7499: 7438: 7315:Expert Opinion on Therapeutic Targets 7065:"Troublesome gene names get the boot" 7031: 6988: 6444: 6160: 5877: 5168:Marigo V, Tabin CJ (September 1996). 4814:Patten I, Placzek M (November 2000). 4582:. U.S. National Library of Medicine. 3955: 2490:Ventral neural domains in neural tube 2083:in the classic Heidelberg screens of 1348:dorsal/ventral neural tube patterning 1298:negative regulation of cell migration 1288:regulation of prostatic bud formation 1268:embryonic skeletal system development 1048:epithelial-mesenchymal cell signaling 818:spinal cord dorsal/ventral patterning 350: 311: 306: 249: 208: 203: 7286:American Journal of Medical Genetics 4820:Cellular and Molecular Life Sciences 4559: 4510:from the original on 1 November 2020 2831: 2577:throughout the ventral neural tube. 2514:result in floor plate induction and 2431:—a structure derived from the axial 2123:. Investigations aimed at finding a 2103:, identified genes that control the 1603:positive regulation of cell division 1583:striated muscle cell differentiation 1513:determination of left/right symmetry 6840:The Journal of Biological Chemistry 4096:Rash BG, Grove EA (November 2011). 3741:. The Weekly Murmur. Archived from 2981:Former controversy surrounding name 2848: 2622:is a crucial mediator of hedgehog ( 1563:Bergmann glial cell differentiation 1413:prostate epithelial cord elongation 963:dopaminergic neuron differentiation 24: 7108: 5903:Ribes V, Briscoe J (August 2009). 3913:Rash BG, Grove EA (October 2007). 3763: 3757: 3737:Keen A, Tabin C (April 12, 2004). 2917:during the translocation into the 1633:tracheoesophageal septum formation 1498:multicellular organism development 1203:striated muscle tissue development 1083:central nervous system development 968:intermediate filament organization 25: 7691: 7351: 7211:Morton JP, Lewis BC (July 2007). 7188:10.1034/j.1399-0004.2001.590503.x 4586:from the original on 2 April 2020 4316:10.1038/scientificamerican0299-74 3616:. Ebury Press. pp. 113–114. 2741:and mechanisms are active (e.g., 7475: 4547:from the original on 31 May 2020 4537:"UniprotKB - Q15465 (SHH_HUMAN)" 3054:, a retinal protein named after 2959:membrane-bound O-acyltransferase 2921:(ER), the first step in protein 2530:blocking of SHH signaling using 2483: 2471: 2459: 2394:turn of the left and right brain 2313:, as well as the progression of 1618:embryonic hindlimb morphogenesis 1508:intein-mediated protein splicing 1378:embryonic forelimb morphogenesis 1363:dorsal/ventral pattern formation 1318:positive thymic T cell selection 1223:anatomical structure development 1213:cellular response to lithium ion 1063:negative thymic T cell selection 858:primary prostatic bud elongation 823:formation of anatomical boundary 773:T cell differentiation in thymus 572: 334: 327: 321: 298: 233: 226: 220: 195: 40: 7464: 7389:US National Library of Medicine 7056: 7025: 6733: 6635: 6438: 6263: 6154: 6105: 5945: 5075:Wilson L, Maden M (June 2005). 4807: 4287: 4251:International Journal of Cancer 4238: 4187: 4089: 3906: 3730: 3719:from the original on 2018-02-28 3641: 3630:from the original on 2017-05-07 3600: 3589:from the original on 2017-08-17 2996:Mothers against decapentaplegic 2968: 2866:Potential regenerative function 1558:protein localization to nucleus 1328:canonical Wnt signaling pathway 1218:embryonic foregut morphogenesis 828:oligodendrocyte differentiation 778:embryonic pattern specification 7032:White M (September 26, 2014). 6697:Molecular and Cellular Biology 4404:10.1523/JNEUROSCI.4938-04.2005 3931:10.1523/JNEUROSCI.3204-07.2007 3335: 3284: 3242: 3191: 3155: 3137: 3119: 2776:makes it a prime example of a 2586:correspond to the distance of 2176:, two of the three vertebrate 2091:, as published in 1980. These 1123:establishment of cell polarity 753:mesenchymal cell proliferation 583:More reference expression data 552:More reference expression data 13: 1: 7268:10.1016/S0962-8924(02)02405-4 7063:Hopkin M (November 6, 2006). 6544:Cardoso WV, Lü J (May 2006). 6459:10.1016/S0070-2153(07)81010-6 6366:10.1016/S0960-9822(98)70446-4 5721:10.1016/S0092-8674(00)80853-3 5284:10.1016/S0092-8674(00)80323-2 4946:10.1016/S0092-8674(00)81386-0 4356:10.1016/S0092-8674(03)00199-5 3891:10.1016/S0012-1606(02)00124-0 3071: 2900: 2478:Ectopic floor plate formation 2127:equivalent in vertebrates by 2014:in all animals. SHH controls 1333:regulation of gene expression 1178:digestive tract morphogenesis 1033:embryonic digit morphogenesis 923:cell population proliferation 868:telencephalon regionalization 743:pattern specification process 319: 218: 27:Signaling molecule in animals 6805:10.1126/science.274.5285.255 6080:10.1371/journal.pone.0111430 5854:10.1016/0092-8674(95)90397-6 4707:10.1016/0959-437X(95)90055-L 3548:10.1016/0092-8674(93)90626-2 3502:10.1016/0092-8674(93)90627-3 3458:10.1016/0092-8674(93)90628-4 3217:10.1371/journal.pbio.1000081 2270:zona limitans intrathalamica 1283:respiratory tube development 1153:smoothened signaling pathway 1103:embryonic limb morphogenesis 7: 7650:Genes on human chromosome 7 7358:An introductory article on 5921:10.1101/cshperspect.a002014 5244:10.1101/SQB.1997.062.01.053 5094:10.1016/j.ydbio.2005.02.027 4392:The Journal of Neuroscience 4114:10.1016/j.ydbio.2011.08.026 3919:The Journal of Neuroscience 3767:Ingenious: The Cyclops Gene 3045: 2915:signal recognition particle 2287:zone of polarizing activity 2230: 2200:'s books for children) and 2085:Christiane Nüsslein-Volhard 1538:neural crest cell migration 1393:regulation of odontogenesis 1163:camera-type eye development 1158:oligodendrocyte development 1078:ventral midline development 1018:embryonic organ development 933:lung epithelium development 918:hair follicle morphogenesis 908:apoptotic signaling pathway 687:endoplasmic reticulum lumen 10: 7696: 7531:hedgehog signaling pathway 6932:Maclean K (January 2006). 6611:10.1016/j.bbrc.2012.11.088 3650:"Pokémon blocks gene name" 3005: 2321:must be secreted from the 2216:and not for the character 993:androgen metabolic process 838:prostate gland development 758:male genitalia development 464:mucosa of transverse colon 444:pancreatic epithelial cell 7616: 7588: 7565: 7537: 7470: 7327:10.1517/14728222.9.6.1137 6272:Principles of Development 4163:10.1007/s00401-011-0922-z 3999:Principles of Development 3648:Simonite T (2005-12-15). 3613:Emus Can't Walk Backwards 3297:Frontiers in Neuroanatomy 3145:"Mouse PubMed Reference:" 3127:"Human PubMed Reference:" 2975:Dr. Ivo "Eggman" Robotnik 2897:receptor γ (Pparg) gene. 2319:fibroblast growth factors 1976: 1971: 1967: 1960: 1944: 1925: 1910: 1906: 1885: 1881: 1874: 1859: 1855: 1834: 1830: 1823: 1810: 1806: 1791: 1787: 1778: 1765: 1761: 1746: 1742: 1733: 1720: 1716: 1701: 1697: 1688: 1673: 1666: 1662: 1646: 1623:forebrain regionalization 1453:salivary gland cavitation 1293:thyroid gland development 1193:roof of mouth development 1183:hair follicle development 943:regulation of proteolysis 646:glycosaminoglycan binding 601: 597: 580: 571: 562: 549: 498: 489: 436: 427: 397: 389: 385: 368: 355: 318: 297: 288: 284: 267: 254: 217: 194: 185: 181: 136: 133: 123: 116: 111: 84: 79: 62: 57: 52: 48: 39: 34: 7122:Human Molecular Genetics 6853:10.1074/jbc.273.22.14037 6742:Nature Reviews. Genetics 6513:10.1165/rcmb.2014-0132TR 4755:10.1073/pnas.95.20.11733 4629:10.1163/157075611X617102 3310:10.3389/fnana.2014.00156 3256:Human Molecular Genetics 2788:. SHH emanates from the 2554:within the neural tube. 2131:, Andrew P. McMahon and 2115:loss of function mutant 1996:) is encoded for by the 1931:Chr 7: 155.8 – 155.81 Mb 1588:myoblast differentiation 1138:neuroblast proliferation 973:renal system development 6889:Nature Chemical Biology 6648:Genes & Development 6249:10.1242/dev.125.15.2803 6192:Journal of Cell Science 6186:Thesleff I (May 2003). 5958:Genes & Development 5792:Genes & Development 5765:10.1242/dev.127.22.4855 5573:Genes & Development 5546:10.1242/dev.126.17.3915 5462:10.1242/dev.125.15.2759 5392:10.1242/dev.125.12.2203 5346:10.1126/science.1098020 5195:10.1073/pnas.93.18.9346 5049:10.1126/science.2237443 4580:Genetics Home Reference 4462:10.1073/pnas.0602920103 4213:10.3389/fonc.2014.00176 4026:10.1242/dev.127.22.4775 3976:10.1242/dev.128.18.3485 3577:Angier N (1994-01-11). 2887:Retinoblastoma proteins 2683:, motor neurons (MN), 2323:apical ectodermal ridge 2208:(formerly described as 2188:(formerly described as 2080:Drosophila melanogaster 1938:Chr 5: 28.66 – 28.67 Mb 1533:embryonic morphogenesis 1518:vasculature development 1373:lung lobe morphogenesis 1243:cell fate specification 928:metanephros development 748:myotube differentiation 7675:Ligands (biochemistry) 7256:Trends in Cell Biology 6660:10.1101/gad.285429.116 6118:Developmental Dynamics 5804:10.1101/gad.12.10.1438 5427:10.1242/dev.127.8.1593 4652:Developmental Dynamics 4543:. UniProt Consortium. 4506:. Ensembl release 99. 3177:10.1006/geno.1995.1104 3038: 3029: 3019: 2879:retinoblastoma protein 2794:concentration gradient 2765:Morphogenetic activity 2573:binding domain of the 2552:concentration gradient 2540:cellular proliferation 2498:Sonic hedgehog is the 2410:central nervous system 2273:. In vertebrates, the 2180:genes are duplicated: 1990:Sonic hedgehog protein 1573:neuron fate commitment 1503:animal organ formation 1338:osteoblast development 1308:right lung development 666:endopeptidase activity 7155:Developmental biology 6709:10.1128/MCB.15.4.2294 5681:10.1242/dev.127.2.333 5081:Developmental Biology 4200:Frontiers in Oncology 4151:Acta Neuropathologica 4102:Developmental Biology 3878:Developmental Biology 3269:10.1093/hmg/8.13.2479 3037: 3027: 3017: 2919:endoplasmic reticulum 2704:transcription factors 2696:transcription factors 2341:retinal ganglion cell 1568:forebrain development 1458:hindbrain development 1438:trachea morphogenesis 1233:stem cell development 1198:left lung development 913:inner ear development 813:hindgut morphogenesis 538:epithelium of stomach 7590:Transcription factor 7387:SHH – sonic hedgehog 7230:10.4161/cc.6.13.4467 7078:10.1038/news061106-2 7040:. Pacific Standard. 7003:10.1002/ajmg.a.31264 6901:10.1038/nchembio.142 6309:10.1242/dev.124.1.53 3745:on November 10, 2005 3709:"A Gene Named Sonic" 3361:10.3390/ijms21030758 2607:transcription factor 2218:Knuckles the Echidna 2190:tiggywinkle hedgehog 2142:Two of these genes, 2067:Discovery and naming 1383:midbrain development 1303:developmental growth 1228:thalamus development 1143:pancreas development 978:cell fate commitment 808:ectoderm development 722:extracellular matrix 717:extracellular region 452:right adrenal cortex 352:5 B1|5 14.39 cM 313:Chromosome 5 (mouse) 211:Chromosome 7 (human) 80:List of PDB id codes 53:Available structures 7151:Gilbert SF (2000). 6950:2006Natur.439..266M 6846:(22): 14037–14045. 6797:1996Sci...274..255P 6357:1998CBio....8.1083P 6198:(Pt 9): 1647–1648. 6071:2014PLoSO...9k1430L 5630:1999Natur.397..617C 5490:Nature Neuroscience 5328:2004Sci...304.1755L 5322:(5678): 1755–1759. 5186:1996PNAS...93.9346M 5131:1996Natur.384..129S 5041:1990Sci...250..985P 4990:1996Natur.383..407C 4885:1995Natur.375..322M 4746:1998PNAS...9511733T 4740:(20): 11733–11738. 4453:2006PNAS..103.8414T 4308:1999SciAm.280b..74R 4296:Scientific American 3925:(43): 11595–11603. 3835:1996Natur.382..452C 3666:2005Natur.438..897S 3407:1980Natur.287..795N 2611:Cubitus interruptus 2330:axonal guidance cue 2196:, a character from 2071:The hedgehog gene ( 1448:cell-cell signaling 833:trachea development 712:extracellular space 616:calcium ion binding 456:left adrenal cortex 440:right lobe of liver 7680:Sonic the Hedgehog 7135:10.1093/hmg/ddl123 6270:Wolpert L (2015). 6023:10.1242/dev.013086 5970:10.1101/gad.937102 5585:10.1101/gad.243402 4832:10.1007/PL00000652 3997:Wolpert L (2015). 3713:The New York Times 3583:The New York Times 3039: 3030: 3020: 2759:ectopic expression 2735:signaling molecule 2724:progenitor domains 2646:SHH function in a 2504:ectopic expression 2249:anterior pituitary 2169:Sonic the Hedgehog 2160:Sonic the Hedgehog 2005:Sonic the Hedgehog 1768:ENSMUSG00000002633 1553:artery development 1473:somite development 1428:lung morphogenesis 1013:thymus development 998:kidney development 768:limb bud formation 731:Biological process 675:Cellular component 656:hydrolase activity 651:morphogen activity 631:peptidase activity 609:Molecular function 7637: 7636: 7527:Signaling pathway 7493: 7492: 7223:(13): 1553–1557. 7176:Clinical Genetics 7166:978-0-87893-243-6 7128:(13): 2031–2037. 6997:(13): 1483–1484. 6791:(5285): 255–259. 6654:(20): 2325–2338. 6563:10.1242/dev.02310 6415:10.1242/dev.02313 6351:(19): 1083–1086. 6281:978-0-19-967814-3 6243:(15): 2803–2811. 6205:10.1242/jcs.00410 6172:978-0-323-07846-7 5798:(10): 1438–1452. 5759:(22): 4855–4866. 5624:(6720): 617–621. 5579:(22): 2865–2878. 5540:(17): 3915–3924. 5456:(15): 2759–2770. 5386:(12): 2203–2212. 5180:(18): 9346–9351. 5125:(6605): 129–134. 5035:(4983): 985–988. 4984:(6599): 407–413. 4879:(6529): 322–325. 4826:(12): 1695–1708. 4500:"ENSG00000164690" 4447:(22): 8414–8418. 4398:(13): 3432–3441. 4264:10.1002/ijc.30450 4066:10.1242/dev.02248 4020:(22): 4775–4785. 3970:(18): 3485–3495. 3829:(6590): 452–455. 3623:978-0-09-192151-4 3401:(5785): 795–801. 3262:(13): 2479–2488. 3043: 3042: 3028:Processing of SHH 2987:holoprosencephaly 2843:holoprosencephaly 2832:Tooth development 2386:Holoprosencephaly 2299:holoprosencephaly 2194:Mrs. Tiggy-Winkle 2032:holoprosencephaly 2024:French flag model 1987: 1986: 1983: 1982: 1956: 1955: 1921: 1920: 1900: 1899: 1870: 1869: 1849: 1848: 1819: 1818: 1800: 1799: 1774: 1773: 1755: 1754: 1729: 1728: 1710: 1709: 1658: 1657: 1463:blood coagulation 1088:heart development 641:laminin-1 binding 621:metal ion binding 593: 592: 589: 588: 558: 557: 545: 544: 483: 482: 381: 380: 280: 279: 107: 106: 103: 102: 63:Ortholog search: 16:(Redirected from 7687: 7520: 7513: 7506: 7497: 7496: 7479: 7459: 7452: 7445: 7436: 7435: 7365:Davidson College 7346: 7321:(6): 1137–1145. 7309: 7279: 7250: 7232: 7207: 7170: 7158: 7147: 7137: 7103: 7102: 7100: 7098: 7080: 7071:: news061106–2. 7060: 7054: 7053: 7051: 7049: 7029: 7023: 7022: 6986: 6980: 6979: 6961: 6929: 6923: 6922: 6912: 6880: 6874: 6873: 6855: 6831: 6825: 6824: 6780: 6774: 6773: 6737: 6731: 6730: 6720: 6703:(4): 2294–2303. 6688: 6682: 6681: 6671: 6639: 6633: 6632: 6622: 6590: 6584: 6583: 6565: 6556:(9): 1611–1624. 6541: 6535: 6534: 6524: 6492: 6481: 6480: 6442: 6436: 6435: 6417: 6408:(8): 1507–1517. 6393: 6387: 6386: 6368: 6336: 6321: 6320: 6292: 6286: 6285: 6267: 6261: 6260: 6232: 6226: 6225: 6207: 6183: 6177: 6176: 6161:Nanci A (2012). 6158: 6152: 6151: 6141: 6130:10.1002/dvdy.191 6124:(9): 1062–1076. 6109: 6103: 6102: 6092: 6082: 6050: 6044: 6043: 6025: 6016:(6): 1097–1106. 6001: 5992: 5991: 5981: 5949: 5943: 5942: 5932: 5900: 5875: 5874: 5856: 5832: 5826: 5825: 5815: 5783: 5777: 5776: 5748: 5742: 5741: 5723: 5699: 5693: 5692: 5664: 5658: 5657: 5613: 5607: 5606: 5596: 5564: 5558: 5557: 5528: 5522: 5521: 5485: 5474: 5473: 5445: 5439: 5438: 5421:(8): 1593–1605. 5410: 5404: 5403: 5375: 5366: 5365: 5339: 5311: 5305: 5304: 5286: 5262: 5256: 5255: 5227: 5218: 5217: 5207: 5197: 5165: 5159: 5158: 5139:10.1038/384129a0 5113: 5107: 5106: 5096: 5072: 5061: 5060: 5024: 5018: 5017: 4998:10.1038/383407a0 4973: 4967: 4966: 4948: 4924: 4913: 4912: 4893:10.1038/375322a0 4868: 4862: 4861: 4843: 4811: 4805: 4804: 4784: 4778: 4777: 4767: 4757: 4725: 4719: 4718: 4690: 4684: 4683: 4647: 4641: 4640: 4622: 4602: 4596: 4595: 4593: 4591: 4572: 4557: 4556: 4554: 4552: 4533: 4520: 4519: 4517: 4515: 4496: 4485: 4484: 4474: 4464: 4432: 4426: 4425: 4415: 4383: 4377: 4376: 4358: 4334: 4328: 4327: 4291: 4285: 4284: 4266: 4242: 4236: 4235: 4225: 4215: 4191: 4185: 4184: 4174: 4142: 4136: 4135: 4125: 4093: 4087: 4086: 4068: 4044: 4038: 4037: 4009: 4003: 4002: 3994: 3988: 3987: 3959: 3953: 3952: 3942: 3910: 3904: 3903: 3893: 3869: 3863: 3862: 3843:10.1038/382452a0 3818: 3812: 3811: 3804:"Zebrafish SHHb" 3800: 3794: 3793: 3786:"Zebrafish SHHa" 3782: 3776: 3775: 3761: 3755: 3754: 3752: 3750: 3734: 3728: 3727: 3725: 3724: 3705: 3696: 3695: 3677: 3645: 3639: 3638: 3636: 3635: 3604: 3598: 3597: 3595: 3594: 3574: 3568: 3567: 3542:(7): 1401–1416. 3531: 3522: 3521: 3496:(7): 1417–1430. 3484: 3478: 3477: 3452:(7): 1431–1444. 3441: 3435: 3434: 3415:10.1038/287795a0 3390: 3384: 3383: 3373: 3363: 3339: 3333: 3332: 3322: 3312: 3288: 3282: 3281: 3271: 3246: 3240: 3239: 3229: 3219: 3195: 3189: 3188: 3159: 3153: 3152: 3141: 3135: 3134: 3123: 3117: 3107: 3096: 3086: 3010: 3009: 2927:signal peptidase 2877:. By modulating 2849:Lung development 2806:progenitor cells 2700:Trigeminal Nerve 2500:secreted protein 2487: 2475: 2463: 2263:, teeth and the 2212:, named for the 2210:echidna hedgehog 2137:homologous genes 1969: 1968: 1940: 1933: 1916: 1904: 1903: 1895: 1879: 1878: 1875:RefSeq (protein) 1865: 1853: 1852: 1844: 1828: 1827: 1804: 1803: 1785: 1784: 1759: 1758: 1740: 1739: 1714: 1713: 1695: 1694: 1664: 1663: 1433:cell development 1388:skin development 1003:lung development 848:limb development 636:zinc ion binding 599: 598: 585: 576: 569: 568: 554: 494: 492:Top expressed in 487: 486: 432: 430:Top expressed in 425: 424: 404: 403: 387: 386: 377: 364: 353: 338: 331: 325: 314: 302: 286: 285: 276: 263: 252: 237: 230: 224: 213: 199: 183: 182: 177: 128: 121: 98: 77: 76: 71: 50: 49: 44: 32: 31: 21: 7695: 7694: 7690: 7689: 7688: 7686: 7685: 7684: 7640: 7639: 7638: 7633: 7612: 7584: 7561: 7557:Desert hedgehog 7552:Indian hedgehog 7533: 7524: 7494: 7489: 7486: 7480: 7466: 7463: 7354: 7349: 7262:(12): 562–569. 7167: 7111: 7109:Further reading 7106: 7096: 7094: 7061: 7057: 7047: 7045: 7030: 7026: 6987: 6983: 6959:10.1038/439266d 6930: 6926: 6881: 6877: 6832: 6828: 6781: 6777: 6754:10.1038/nrg2984 6738: 6734: 6689: 6685: 6640: 6636: 6591: 6587: 6542: 6538: 6493: 6484: 6469: 6443: 6439: 6394: 6390: 6345:Current Biology 6337: 6324: 6293: 6289: 6282: 6268: 6264: 6233: 6229: 6184: 6180: 6173: 6159: 6155: 6110: 6106: 6065:(11): e111430. 6051: 6047: 6002: 5995: 5950: 5946: 5901: 5878: 5833: 5829: 5784: 5780: 5749: 5745: 5700: 5696: 5665: 5661: 5614: 5610: 5565: 5561: 5529: 5525: 5496:(10): 979–985. 5486: 5477: 5446: 5442: 5411: 5407: 5376: 5369: 5337:10.1.1.476.3902 5312: 5308: 5263: 5259: 5228: 5221: 5166: 5162: 5114: 5110: 5073: 5064: 5025: 5021: 4974: 4970: 4925: 4916: 4869: 4865: 4812: 4808: 4785: 4781: 4726: 4722: 4691: 4687: 4648: 4644: 4603: 4599: 4589: 4587: 4574: 4573: 4560: 4550: 4548: 4535: 4534: 4523: 4513: 4511: 4498: 4497: 4488: 4433: 4429: 4384: 4380: 4335: 4331: 4292: 4288: 4243: 4239: 4192: 4188: 4143: 4139: 4094: 4090: 4045: 4041: 4010: 4006: 3995: 3991: 3960: 3956: 3911: 3907: 3870: 3866: 3819: 3815: 3802: 3801: 3797: 3784: 3783: 3779: 3762: 3758: 3748: 3746: 3735: 3731: 3722: 3720: 3707: 3706: 3699: 3675:10.1038/438897a 3646: 3642: 3633: 3631: 3624: 3605: 3601: 3592: 3590: 3575: 3571: 3532: 3525: 3485: 3481: 3442: 3438: 3391: 3387: 3340: 3336: 3289: 3285: 3247: 3243: 3196: 3192: 3160: 3156: 3143: 3142: 3138: 3125: 3124: 3120: 3108: 3099: 3087: 3078: 3074: 3048: 3008: 2983: 2971: 2911:signal sequence 2903: 2868: 2851: 2834: 2767: 2626:) signaling in 2516:differentiation 2496: 2495: 2494: 2491: 2488: 2479: 2476: 2467: 2464: 2406: 2315:prostate cancer 2311:medulloblastoma 2233: 2172:(1991). In the 2151:Indian hedgehog 2145:desert hedgehog 2135:revealed three 2107:pattern of the 2101:Edward B. Lewis 2069: 2061:and many more. 1978:View/Edit Mouse 1973:View/Edit Human 1936: 1929: 1926:Location (UCSC) 1912: 1891: 1887: 1861: 1840: 1836: 1749:ENSG00000164690 1642: 726: 707:plasma membrane 670: 661:protein binding 626:patched binding 581: 550: 541: 536: 532: 528: 524: 520: 516: 512: 508: 504: 490: 479: 474: 472:urinary bladder 470: 466: 462: 458: 454: 450: 446: 442: 428: 372: 359: 351: 341: 340: 339: 332: 312: 289:Gene location ( 271: 258: 250: 240: 239: 238: 231: 209: 186:Gene location ( 175:SHH - orthologs 137: 124: 117: 86: 64: 28: 23: 22: 15: 12: 11: 5: 7693: 7683: 7682: 7677: 7672: 7670:Cell signaling 7667: 7662: 7657: 7652: 7635: 7634: 7632: 7631: 7626: 7620: 7618: 7614: 7613: 7611: 7610: 7605: 7600: 7594: 7592: 7586: 7585: 7583: 7582: 7577: 7571: 7569: 7563: 7562: 7560: 7559: 7554: 7549: 7547:Sonic hedgehog 7543: 7541: 7535: 7534: 7523: 7522: 7515: 7508: 7500: 7491: 7490: 7488: 7487: 7481: 7474: 7471: 7468: 7467: 7462: 7461: 7454: 7447: 7439: 7433: 7432: 7411: 7390: 7384: 7379: 7373: 7367: 7353: 7352:External links 7350: 7348: 7347: 7310: 7280: 7251: 7208: 7182:(5): 306–315. 7171: 7165: 7148: 7112: 7110: 7107: 7105: 7104: 7055: 7024: 6981: 6924: 6895:(3): 154–156. 6875: 6826: 6775: 6748:(6): 393–406. 6732: 6683: 6634: 6605:(2): 700–705. 6585: 6536: 6482: 6467: 6437: 6388: 6322: 6287: 6280: 6262: 6227: 6178: 6171: 6153: 6104: 6045: 5993: 5964:(5): 548–553. 5944: 5915:(2): a002014. 5876: 5847:(3): 445–455. 5827: 5778: 5743: 5714:(4): 435–445. 5694: 5675:(2): 333–342. 5659: 5608: 5559: 5523: 5475: 5440: 5405: 5367: 5306: 5277:(1): 169–180. 5257: 5219: 5160: 5108: 5062: 5019: 4968: 4939:(4): 661–673. 4914: 4863: 4806: 4779: 4720: 4701:(4): 499–506. 4685: 4658:(2): 143–154. 4642: 4613:(2): 193–216. 4607:Animal Biology 4597: 4558: 4521: 4486: 4427: 4378: 4329: 4286: 4257:(2): 358–369. 4237: 4186: 4157:(4): 465–472. 4137: 4108:(2): 242–250. 4088: 4059:(5): 855–864. 4039: 4004: 3989: 3954: 3905: 3864: 3813: 3810:on 2009-06-26. 3795: 3792:on 2009-06-25. 3777: 3756: 3729: 3715:. 1994-01-11. 3697: 3640: 3622: 3610:(2007-09-06). 3599: 3569: 3523: 3479: 3436: 3385: 3334: 3283: 3241: 3190: 3154: 3136: 3118: 3097: 3075: 3073: 3070: 3069: 3068: 3058: 3047: 3044: 3041: 3040: 3031: 3021: 3007: 3004: 3000:Lunatic fringe 2991:sonic hedgehog 2982: 2979: 2970: 2967: 2902: 2899: 2867: 2864: 2850: 2847: 2833: 2830: 2766: 2763: 2493: 2492: 2489: 2482: 2480: 2477: 2470: 2468: 2465: 2458: 2455: 2454: 2453: 2444:vertebrate CNS 2405: 2402: 2382:Microphthalmia 2255:of the brain, 2232: 2229: 2214:spiny anteater 2198:Beatrix Potter 2156:sonic hedgehog 2133:Clifford Tabin 2089:Eric Wieschaus 2068: 2065: 1985: 1984: 1981: 1980: 1975: 1965: 1964: 1958: 1957: 1954: 1953: 1951: 1949: 1942: 1941: 1934: 1927: 1923: 1922: 1919: 1918: 1908: 1907: 1901: 1898: 1897: 1883: 1882: 1876: 1872: 1871: 1868: 1867: 1857: 1856: 1850: 1847: 1846: 1832: 1831: 1825: 1821: 1820: 1817: 1816: 1808: 1807: 1801: 1798: 1797: 1789: 1788: 1782: 1776: 1775: 1772: 1771: 1763: 1762: 1756: 1753: 1752: 1744: 1743: 1737: 1731: 1730: 1727: 1726: 1718: 1717: 1711: 1708: 1707: 1699: 1698: 1692: 1686: 1685: 1680: 1675: 1671: 1670: 1660: 1659: 1656: 1655: 1644: 1643: 1641: 1640: 1635: 1630: 1625: 1620: 1615: 1610: 1605: 1600: 1595: 1590: 1585: 1580: 1575: 1570: 1565: 1560: 1555: 1550: 1545: 1540: 1535: 1530: 1525: 1520: 1515: 1510: 1505: 1500: 1495: 1490: 1485: 1480: 1475: 1470: 1465: 1460: 1455: 1450: 1445: 1440: 1435: 1430: 1425: 1420: 1415: 1410: 1405: 1400: 1395: 1390: 1385: 1380: 1375: 1370: 1365: 1360: 1355: 1350: 1345: 1340: 1335: 1330: 1325: 1320: 1315: 1310: 1305: 1300: 1295: 1290: 1285: 1280: 1275: 1270: 1265: 1260: 1255: 1250: 1245: 1240: 1235: 1230: 1225: 1220: 1215: 1210: 1205: 1200: 1195: 1190: 1185: 1180: 1175: 1170: 1165: 1160: 1155: 1150: 1145: 1140: 1135: 1130: 1125: 1120: 1115: 1110: 1105: 1100: 1095: 1090: 1085: 1080: 1075: 1070: 1065: 1060: 1055: 1050: 1045: 1040: 1035: 1030: 1025: 1020: 1015: 1010: 1005: 1000: 995: 990: 985: 980: 975: 970: 965: 960: 955: 950: 945: 940: 935: 930: 925: 920: 915: 910: 905: 900: 895: 890: 885: 880: 875: 870: 865: 860: 855: 853:vasculogenesis 850: 845: 840: 835: 830: 825: 820: 815: 810: 805: 800: 795: 790: 785: 780: 775: 770: 765: 760: 755: 750: 745: 740: 734: 732: 728: 727: 725: 724: 719: 714: 709: 704: 699: 694: 689: 684: 678: 676: 672: 671: 669: 668: 663: 658: 653: 648: 643: 638: 633: 628: 623: 618: 612: 610: 606: 605: 595: 594: 591: 590: 587: 586: 578: 577: 566: 560: 559: 556: 555: 547: 546: 543: 542: 540: 539: 535: 531: 530:left lung lobe 527: 523: 519: 515: 511: 507: 503: 499: 496: 495: 484: 481: 480: 478: 477: 476:spinal ganglia 473: 469: 465: 461: 457: 453: 449: 445: 441: 437: 434: 433: 421: 420: 412: 401: 395: 394: 391:RNA expression 383: 382: 379: 378: 370: 366: 365: 357: 354: 349: 343: 342: 333: 326: 320: 316: 315: 310: 304: 303: 295: 294: 282: 281: 278: 277: 269: 265: 264: 256: 253: 248: 242: 241: 232: 225: 219: 215: 214: 207: 201: 200: 192: 191: 179: 178: 135: 131: 130: 122: 114: 113: 109: 108: 105: 104: 101: 100: 82: 81: 73: 72: 61: 55: 54: 46: 45: 37: 36: 26: 18:Sonic hedgehog 9: 6: 4: 3: 2: 7692: 7681: 7678: 7676: 7673: 7671: 7668: 7666: 7663: 7661: 7658: 7656: 7653: 7651: 7648: 7647: 7645: 7630: 7627: 7625: 7622: 7621: 7619: 7615: 7609: 7606: 7604: 7601: 7599: 7596: 7595: 7593: 7591: 7587: 7581: 7578: 7576: 7573: 7572: 7570: 7568: 7564: 7558: 7555: 7553: 7550: 7548: 7545: 7544: 7542: 7540: 7536: 7532: 7528: 7521: 7516: 7514: 7509: 7507: 7502: 7501: 7498: 7484: 7478: 7473: 7472: 7469: 7460: 7455: 7453: 7448: 7446: 7441: 7440: 7437: 7430: 7426: 7425: 7420: 7416: 7412: 7409: 7405: 7404: 7399: 7395: 7391: 7388: 7385: 7383: 7380: 7377: 7374: 7371: 7368: 7366: 7362: 7361: 7356: 7355: 7344: 7340: 7336: 7332: 7328: 7324: 7320: 7316: 7311: 7307: 7303: 7299: 7295: 7291: 7287: 7281: 7277: 7273: 7269: 7265: 7261: 7257: 7252: 7248: 7244: 7240: 7236: 7231: 7226: 7222: 7218: 7214: 7209: 7205: 7201: 7197: 7193: 7189: 7185: 7181: 7177: 7172: 7168: 7162: 7157: 7156: 7149: 7145: 7141: 7136: 7131: 7127: 7123: 7119: 7114: 7113: 7092: 7088: 7084: 7079: 7074: 7070: 7066: 7059: 7043: 7039: 7035: 7028: 7020: 7016: 7012: 7008: 7004: 7000: 6996: 6992: 6985: 6977: 6973: 6969: 6965: 6960: 6955: 6951: 6947: 6944:(7074): 266. 6943: 6939: 6935: 6928: 6920: 6916: 6911: 6906: 6902: 6898: 6894: 6890: 6886: 6879: 6871: 6867: 6863: 6859: 6854: 6849: 6845: 6841: 6837: 6830: 6822: 6818: 6814: 6810: 6806: 6802: 6798: 6794: 6790: 6786: 6779: 6771: 6767: 6763: 6759: 6755: 6751: 6747: 6743: 6736: 6728: 6724: 6719: 6714: 6710: 6706: 6702: 6698: 6694: 6687: 6679: 6675: 6670: 6665: 6661: 6657: 6653: 6649: 6645: 6638: 6630: 6626: 6621: 6616: 6612: 6608: 6604: 6600: 6596: 6589: 6581: 6577: 6573: 6569: 6564: 6559: 6555: 6551: 6547: 6540: 6532: 6528: 6523: 6518: 6514: 6510: 6506: 6502: 6498: 6491: 6489: 6487: 6478: 6474: 6470: 6468:9780123742537 6464: 6460: 6456: 6452: 6448: 6441: 6433: 6429: 6425: 6421: 6416: 6411: 6407: 6403: 6399: 6392: 6384: 6380: 6376: 6372: 6367: 6362: 6358: 6354: 6350: 6346: 6342: 6335: 6333: 6331: 6329: 6327: 6318: 6314: 6310: 6306: 6302: 6298: 6291: 6283: 6277: 6273: 6266: 6258: 6254: 6250: 6246: 6242: 6238: 6231: 6223: 6219: 6215: 6211: 6206: 6201: 6197: 6193: 6189: 6182: 6174: 6168: 6164: 6157: 6149: 6145: 6140: 6135: 6131: 6127: 6123: 6119: 6115: 6108: 6100: 6096: 6091: 6086: 6081: 6076: 6072: 6068: 6064: 6060: 6056: 6049: 6041: 6037: 6033: 6029: 6024: 6019: 6015: 6011: 6007: 6000: 5998: 5989: 5985: 5980: 5975: 5971: 5967: 5963: 5959: 5955: 5948: 5940: 5936: 5931: 5926: 5922: 5918: 5914: 5910: 5906: 5899: 5897: 5895: 5893: 5891: 5889: 5887: 5885: 5883: 5881: 5872: 5868: 5864: 5860: 5855: 5850: 5846: 5842: 5838: 5831: 5823: 5819: 5814: 5809: 5805: 5801: 5797: 5793: 5789: 5782: 5774: 5770: 5766: 5762: 5758: 5754: 5747: 5739: 5735: 5731: 5727: 5722: 5717: 5713: 5709: 5705: 5698: 5690: 5686: 5682: 5678: 5674: 5670: 5663: 5655: 5651: 5647: 5643: 5639: 5638:10.1038/17611 5635: 5631: 5627: 5623: 5619: 5612: 5604: 5600: 5595: 5590: 5586: 5582: 5578: 5574: 5570: 5563: 5555: 5551: 5547: 5543: 5539: 5535: 5527: 5519: 5515: 5511: 5507: 5503: 5502:10.1038/79916 5499: 5495: 5491: 5484: 5482: 5480: 5471: 5467: 5463: 5459: 5455: 5451: 5444: 5436: 5432: 5428: 5424: 5420: 5416: 5409: 5401: 5397: 5393: 5389: 5385: 5381: 5374: 5372: 5363: 5359: 5355: 5351: 5347: 5343: 5338: 5333: 5329: 5325: 5321: 5317: 5310: 5302: 5298: 5294: 5290: 5285: 5280: 5276: 5272: 5268: 5261: 5253: 5249: 5245: 5241: 5237: 5233: 5226: 5224: 5215: 5211: 5206: 5201: 5196: 5191: 5187: 5183: 5179: 5175: 5171: 5164: 5156: 5152: 5148: 5144: 5140: 5136: 5132: 5128: 5124: 5120: 5112: 5104: 5100: 5095: 5090: 5086: 5082: 5078: 5071: 5069: 5067: 5058: 5054: 5050: 5046: 5042: 5038: 5034: 5030: 5023: 5015: 5011: 5007: 5003: 4999: 4995: 4991: 4987: 4983: 4979: 4972: 4964: 4960: 4956: 4952: 4947: 4942: 4938: 4934: 4930: 4923: 4921: 4919: 4910: 4906: 4902: 4898: 4894: 4890: 4886: 4882: 4878: 4874: 4867: 4859: 4855: 4851: 4847: 4842: 4837: 4833: 4829: 4825: 4821: 4817: 4810: 4802: 4798: 4794: 4790: 4783: 4775: 4771: 4766: 4761: 4756: 4751: 4747: 4743: 4739: 4735: 4731: 4724: 4716: 4712: 4708: 4704: 4700: 4696: 4689: 4681: 4677: 4673: 4669: 4665: 4661: 4657: 4653: 4646: 4638: 4634: 4630: 4626: 4621: 4616: 4612: 4608: 4601: 4585: 4581: 4577: 4571: 4569: 4567: 4565: 4563: 4546: 4542: 4538: 4532: 4530: 4528: 4526: 4509: 4505: 4501: 4495: 4493: 4491: 4482: 4478: 4473: 4468: 4463: 4458: 4454: 4450: 4446: 4442: 4438: 4431: 4423: 4419: 4414: 4409: 4405: 4401: 4397: 4393: 4389: 4382: 4374: 4370: 4366: 4362: 4357: 4352: 4348: 4344: 4340: 4333: 4325: 4321: 4317: 4313: 4309: 4305: 4301: 4297: 4290: 4282: 4278: 4274: 4270: 4265: 4260: 4256: 4252: 4248: 4241: 4233: 4229: 4224: 4219: 4214: 4209: 4205: 4201: 4197: 4190: 4182: 4178: 4173: 4168: 4164: 4160: 4156: 4152: 4148: 4141: 4133: 4129: 4124: 4119: 4115: 4111: 4107: 4103: 4099: 4092: 4084: 4080: 4076: 4072: 4067: 4062: 4058: 4054: 4050: 4043: 4035: 4031: 4027: 4023: 4019: 4015: 4008: 4000: 3993: 3985: 3981: 3977: 3973: 3969: 3965: 3958: 3950: 3946: 3941: 3936: 3932: 3928: 3924: 3920: 3916: 3909: 3901: 3897: 3892: 3887: 3883: 3879: 3875: 3868: 3860: 3856: 3852: 3848: 3844: 3840: 3836: 3832: 3828: 3824: 3817: 3809: 3805: 3799: 3791: 3787: 3781: 3773: 3769: 3768: 3760: 3744: 3740: 3733: 3718: 3714: 3710: 3704: 3702: 3693: 3689: 3685: 3681: 3676: 3671: 3667: 3663: 3659: 3655: 3651: 3644: 3629: 3625: 3619: 3615: 3614: 3609: 3603: 3588: 3584: 3580: 3573: 3565: 3561: 3557: 3553: 3549: 3545: 3541: 3537: 3530: 3528: 3519: 3515: 3511: 3507: 3503: 3499: 3495: 3491: 3483: 3475: 3471: 3467: 3463: 3459: 3455: 3451: 3447: 3440: 3432: 3428: 3424: 3420: 3416: 3412: 3408: 3404: 3400: 3396: 3389: 3381: 3377: 3372: 3367: 3362: 3357: 3353: 3349: 3345: 3338: 3330: 3326: 3321: 3316: 3311: 3306: 3302: 3298: 3294: 3287: 3279: 3275: 3270: 3265: 3261: 3257: 3253: 3245: 3237: 3233: 3228: 3223: 3218: 3213: 3209: 3205: 3201: 3194: 3186: 3182: 3178: 3174: 3170: 3166: 3158: 3150: 3146: 3140: 3132: 3128: 3122: 3115: 3111: 3106: 3104: 3102: 3094: 3090: 3085: 3083: 3081: 3076: 3066: 3062: 3059: 3057: 3053: 3050: 3049: 3036: 3032: 3026: 3022: 3016: 3012: 3011: 3003: 3001: 2997: 2992: 2988: 2978: 2976: 2966: 2964: 2960: 2956: 2952: 2948: 2945: 2941: 2937: 2933: 2928: 2924: 2920: 2916: 2912: 2908: 2898: 2894: 2892: 2888: 2884: 2880: 2876: 2873: 2863: 2861: 2857: 2846: 2844: 2840: 2829: 2825: 2823: 2819: 2813: 2811: 2807: 2802: 2799: 2795: 2791: 2787: 2786:motor neurons 2783: 2779: 2775: 2772: 2762: 2760: 2756: 2752: 2748: 2744: 2740: 2736: 2731: 2729: 2725: 2721: 2717: 2713: 2709: 2705: 2701: 2697: 2694: 2690: 2686: 2682: 2679: 2674: 2672: 2668: 2663: 2661: 2655: 2653: 2649: 2645: 2641: 2637: 2633: 2629: 2625: 2621: 2617: 2613: 2612: 2608: 2604: 2601: 2597: 2593: 2589: 2585: 2580: 2576: 2572: 2568: 2566: 2560: 2555: 2553: 2549: 2545: 2541: 2537: 2533: 2529: 2525: 2521: 2517: 2513: 2509: 2505: 2501: 2486: 2481: 2474: 2469: 2462: 2457: 2456: 2452: 2449: 2445: 2441: 2436: 2434: 2430: 2426: 2422: 2418: 2415: 2412:(CNS) during 2411: 2401: 2399: 2395: 2391: 2387: 2383: 2378: 2374: 2369: 2365: 2361: 2356: 2354: 2350: 2346: 2342: 2338: 2335: 2331: 2326: 2324: 2320: 2316: 2312: 2308: 2304: 2300: 2296: 2292: 2288: 2284: 2280: 2276: 2272: 2271: 2266: 2262: 2258: 2254: 2250: 2246: 2242: 2238: 2228: 2226: 2224: 2219: 2215: 2211: 2207: 2203: 2199: 2195: 2191: 2187: 2183: 2179: 2175: 2171: 2170: 2165: 2161: 2157: 2153: 2152: 2147: 2146: 2140: 2138: 2134: 2130: 2129:Philip Ingham 2126: 2122: 2118: 2114: 2111:embryos. The 2110: 2106: 2102: 2098: 2094: 2090: 2086: 2082: 2081: 2076: 2075: 2064: 2062: 2060: 2056: 2052: 2048: 2044: 2040: 2035: 2033: 2029: 2025: 2021: 2017: 2016:organogenesis 2013: 2012:morphogenesis 2008: 2006: 2002: 1999: 1995: 1991: 1979: 1974: 1970: 1966: 1963: 1959: 1952: 1950: 1947: 1943: 1939: 1935: 1932: 1928: 1924: 1917: 1915: 1909: 1905: 1902: 1896: 1894: 1890: 1884: 1880: 1877: 1873: 1866: 1864: 1858: 1854: 1851: 1845: 1843: 1839: 1833: 1829: 1826: 1824:RefSeq (mRNA) 1822: 1815: 1814: 1809: 1805: 1802: 1796: 1795: 1790: 1786: 1783: 1781: 1777: 1770: 1769: 1764: 1760: 1757: 1751: 1750: 1745: 1741: 1738: 1736: 1732: 1725: 1724: 1719: 1715: 1712: 1706: 1705: 1700: 1696: 1693: 1691: 1687: 1684: 1681: 1679: 1676: 1672: 1669: 1665: 1661: 1654: 1650: 1645: 1639: 1636: 1634: 1631: 1629: 1626: 1624: 1621: 1619: 1616: 1614: 1611: 1609: 1606: 1604: 1601: 1599: 1596: 1594: 1591: 1589: 1586: 1584: 1581: 1579: 1576: 1574: 1571: 1569: 1566: 1564: 1561: 1559: 1556: 1554: 1551: 1549: 1546: 1544: 1541: 1539: 1536: 1534: 1531: 1529: 1526: 1524: 1521: 1519: 1516: 1514: 1511: 1509: 1506: 1504: 1501: 1499: 1496: 1494: 1491: 1489: 1486: 1484: 1483:axon guidance 1481: 1479: 1476: 1474: 1471: 1469: 1466: 1464: 1461: 1459: 1456: 1454: 1451: 1449: 1446: 1444: 1441: 1439: 1436: 1434: 1431: 1429: 1426: 1424: 1421: 1419: 1416: 1414: 1411: 1409: 1406: 1404: 1401: 1399: 1396: 1394: 1391: 1389: 1386: 1384: 1381: 1379: 1376: 1374: 1371: 1369: 1366: 1364: 1361: 1359: 1356: 1354: 1351: 1349: 1346: 1344: 1341: 1339: 1336: 1334: 1331: 1329: 1326: 1324: 1321: 1319: 1316: 1314: 1311: 1309: 1306: 1304: 1301: 1299: 1296: 1294: 1291: 1289: 1286: 1284: 1281: 1279: 1278:odontogenesis 1276: 1274: 1271: 1269: 1266: 1264: 1261: 1259: 1256: 1254: 1251: 1249: 1246: 1244: 1241: 1239: 1236: 1234: 1231: 1229: 1226: 1224: 1221: 1219: 1216: 1214: 1211: 1209: 1206: 1204: 1201: 1199: 1196: 1194: 1191: 1189: 1186: 1184: 1181: 1179: 1176: 1174: 1171: 1169: 1166: 1164: 1161: 1159: 1156: 1154: 1151: 1149: 1146: 1144: 1141: 1139: 1136: 1134: 1131: 1129: 1126: 1124: 1121: 1119: 1116: 1114: 1111: 1109: 1106: 1104: 1101: 1099: 1096: 1094: 1091: 1089: 1086: 1084: 1081: 1079: 1076: 1074: 1071: 1069: 1066: 1064: 1061: 1059: 1056: 1054: 1051: 1049: 1046: 1044: 1041: 1039: 1036: 1034: 1031: 1029: 1026: 1024: 1021: 1019: 1016: 1014: 1011: 1009: 1006: 1004: 1001: 999: 996: 994: 991: 989: 986: 984: 981: 979: 976: 974: 971: 969: 966: 964: 961: 959: 956: 954: 951: 949: 946: 944: 941: 939: 936: 934: 931: 929: 926: 924: 921: 919: 916: 914: 911: 909: 906: 904: 901: 899: 896: 894: 891: 889: 886: 884: 881: 879: 876: 874: 871: 869: 866: 864: 863:heart looping 861: 859: 856: 854: 851: 849: 846: 844: 841: 839: 836: 834: 831: 829: 826: 824: 821: 819: 816: 814: 811: 809: 806: 804: 801: 799: 796: 794: 791: 789: 786: 784: 781: 779: 776: 774: 771: 769: 766: 764: 761: 759: 756: 754: 751: 749: 746: 744: 741: 739: 736: 735: 733: 730: 729: 723: 720: 718: 715: 713: 710: 708: 705: 703: 702:membrane raft 700: 698: 695: 693: 690: 688: 685: 683: 680: 679: 677: 674: 673: 667: 664: 662: 659: 657: 654: 652: 649: 647: 644: 642: 639: 637: 634: 632: 629: 627: 624: 622: 619: 617: 614: 613: 611: 608: 607: 604: 603:Gene ontology 600: 596: 584: 579: 575: 570: 567: 565: 561: 553: 548: 537: 533: 529: 525: 521: 517: 513: 509: 505: 501: 500: 497: 493: 488: 485: 475: 471: 467: 463: 459: 455: 451: 447: 443: 439: 438: 435: 431: 426: 423: 422: 419: 417: 413: 411: 410: 406: 405: 402: 400: 396: 392: 388: 384: 376: 371: 367: 363: 358: 348: 344: 337: 330: 324: 317: 309: 305: 301: 296: 292: 287: 283: 275: 270: 266: 262: 257: 247: 243: 236: 229: 223: 216: 212: 206: 202: 198: 193: 189: 184: 180: 176: 172: 168: 164: 160: 156: 152: 148: 144: 140: 132: 127: 120: 115: 110: 99: 97: 93: 89: 83: 78: 75: 74: 70: 67: 60: 56: 51: 47: 43: 38: 33: 30: 19: 7482: 7422: 7401: 7359: 7318: 7314: 7289: 7285: 7259: 7255: 7220: 7216: 7179: 7175: 7154: 7125: 7121: 7097:December 24, 7095:. Retrieved 7068: 7058: 7048:December 24, 7046:. Retrieved 7037: 7027: 6994: 6990: 6984: 6941: 6937: 6927: 6892: 6888: 6878: 6843: 6839: 6829: 6788: 6784: 6778: 6745: 6741: 6735: 6700: 6696: 6686: 6651: 6647: 6637: 6602: 6598: 6588: 6553: 6549: 6539: 6504: 6500: 6450: 6446: 6440: 6405: 6401: 6391: 6348: 6344: 6303:(1): 53–63. 6300: 6296: 6290: 6271: 6265: 6240: 6236: 6230: 6195: 6191: 6181: 6162: 6156: 6121: 6117: 6107: 6062: 6058: 6048: 6013: 6009: 5961: 5957: 5947: 5912: 5908: 5844: 5840: 5830: 5795: 5791: 5781: 5756: 5752: 5746: 5711: 5707: 5697: 5672: 5668: 5662: 5621: 5617: 5611: 5576: 5572: 5562: 5537: 5533: 5526: 5493: 5489: 5453: 5449: 5443: 5418: 5414: 5408: 5383: 5379: 5319: 5315: 5309: 5274: 5270: 5260: 5235: 5231: 5177: 5173: 5163: 5122: 5118: 5111: 5084: 5080: 5032: 5028: 5022: 4981: 4977: 4971: 4936: 4932: 4876: 4872: 4866: 4823: 4819: 4809: 4795:(2): 91–97. 4792: 4788: 4782: 4737: 4733: 4723: 4698: 4694: 4688: 4655: 4651: 4645: 4610: 4606: 4600: 4588:. Retrieved 4579: 4549:. Retrieved 4540: 4512:. Retrieved 4503: 4444: 4440: 4430: 4395: 4391: 4381: 4349:(1): 11–23. 4346: 4342: 4332: 4302:(2): 74–79. 4299: 4295: 4289: 4254: 4250: 4240: 4203: 4199: 4189: 4154: 4150: 4140: 4105: 4101: 4091: 4056: 4052: 4042: 4017: 4013: 4007: 3998: 3992: 3967: 3963: 3957: 3922: 3918: 3908: 3884:(1): 36–49. 3881: 3877: 3867: 3826: 3822: 3816: 3808:the original 3798: 3790:the original 3780: 3771: 3766: 3759: 3747:. Retrieved 3743:the original 3732: 3721:. Retrieved 3712: 3660:(897): 897. 3657: 3653: 3643: 3632:. Retrieved 3612: 3602: 3591:. 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