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Title: Multiple phenotypes resulting from a mutagenesis screen for pharynx muscle mutations in Caenorhabditis elegans .
Authors: Ferrier A ; Charron A ; Sadozai Y ; Switaj L ; Szutenbach A ; Smith PA
Journal: PLoS One
Year: 2011
Doc ID: WBPaper00040399
Bibliographic Information
Abstract
Matching Sentences
SECTION: results. Phenotype * Number Anterior bulb morphology 18 APH ( anterior pharynx absent ) 5 Pharynx basement membrane defect 11 Short pharynx 20 GFP-expressing cells outside of pharynx 14 Isthmus defect 11 Pha ( pharynx absent ) 1 Pharynx asymmetric 21 Pharynx GFP weak / nearly absent 6 Posterior bulb defect 9 Procorpus defect 29 Pun ( Pharynx unattached ) 7 Thin , cylindrical pharynx 25 * Worm strains may have overlapping phenotypes . doi : 10 . 1371 / journal . pone . [Field: results, subscore: 18.00]
SECTION: results. Gene Location Phenotype C26E6 . 6 III : 22 . 34 Body morphology variant C35D10 . 5 III : 22 . 42 Short Pharynx , large head rnp-4 III : 23 . 18 Pharynx asymmetry dcn-1 III : 23 . 19 Body morphology variant M88 . 2 III : 23 . 12 Short Pharynx , Slightly Dpy xbp-1 III : 23 . 70 myo-2 : : GFP expression limited ccdc-55 III : 23 . 72 Pharynx asymmetry wht-3 III : 23 . 79 Short Pharynx , Slightly Dpy doi : 10 . 1371 / journal . pone . [Field: results, subscore: 10.00]
SECTION: results. 10 I : 6 . 41 Pharynx thin and asymmetric hmr-1 I : 6 . 61 Short pharynx rsr-1 I : 7 . 25 Pharynx asymmetry F56G4 . 4 I : 7 . 96 Pharynx asymmetry doi : 10 . 1371 / journal . pone . [Field: results, subscore: 8.00]
SECTION: discussion. The lack of consistent staining AJM-1 antibody in the pharynx of PAS136 embryos suggests that a disruption to the adherence of one cell to another is contributing to the mutant phenotype , specifically in the pharynx since adherens junction staining in other areas of the embryo is nearly normal . This implies that the allele mutated in PAS136 may be specific to the pharynx , although it is possible that AJM-1 or accessory proteins such as DLG-1 , LET-413 , or HMP-1 are specifically missing pharyngeal enhancers necessary for it issue specific expression1 [ 30 ] . [Field: discussion, subscore: 6.00]
SECTION: introduction. This report describes multiple classes of mutant pharynx phenotypes isolated from an Ethyl Methanesulfonate ( EMS ) mutagenesis screen for larvae with phenotypes such as short pharynx , thin-cylindrical pharynx , non-adherent cells , anterior pharynx absent , pharynx cells outside the basement membrane , and pharynx unattached . [Field: introduction, subscore: 6.00]
SECTION: results. Gene Name Location Phenotype mom-5 I : 4 . 12 Pharynx Unattached blmp-1 I : 4 . 99 Amorphous pharynx sec-8 I : 5 . 04 Compressed isthmus phi-56 I : 5 . 06 Anterior bulb / isthmus merged , long , thin procorpus lam-3 I : 5 . 06 Amorphous pharynx Y52B11A . [Field: results, subscore: 6.00]
SECTION: discussion. Because MH27 stains adherens junctions , which are easily detected in the pharynx , the lack of a pharynx was obvious in the chromosomal deficiency ozDf2 , which deletes pha-4 [ 33 ] . [Field: discussion, subscore: 4.00]
SECTION: discussion. Pharynx Phenotype Screen PLoS ONE | www . plosone . org 8 November 2011 | Volume 6 | Issue 11 | e26594 pha-4 alleles have been isolated in numerous screens ; although most were not designed to isolate pharynx phenotypes specifically . [Field: discussion, subscore: 4.00]
SECTION: discussion. The isolation of pha-2 , pha-3 , and various Eat mutants was facilitated by using an unc-31 genetic background to identify slow growing , starved , and abnormally pumping worms under a dissecting microscope [ 19 ] ; while most pharynx abnormalities will result in slow growth or larval arrest , many genes responsible for these phenotypes are not specific to the pharynx . [Field: discussion, subscore: 4.00]
SECTION: discussion. A genetic screen optimized for rapid identification of pharynx mutant phenotypes The genetic screen described in this paper was unique in the approach to observing pharynx morphology phenotypes . [Field: discussion, subscore: 4.00]
SECTION: discussion. As expected , myo- 2 : : GFP : : H2B nuclei appear randomly positioned in the mutant pharynx as compared to the wild-type pharynx , consistent with the hypothesis that abnormal morphology and adhesion are the key defects in PAS136 animals . [Field: discussion, subscore: 4.00]
SECTION: discussion. PAS136 mutants are not lacking pharynx muscle cells The C . elegans pharynx is made up of exactly twenty muscle cells with 37 distinct nuclei as a result of the fusion of adjacent cells in many muscle groups during development [ 4 ] . [Field: discussion, subscore: 4.00]
SECTION: discussion. Since PAS136 appear to only suffer adhesion defects in the pharynx , it is likely the mutation of a gene crucial for pharynx adherens junctions but redundant in those in the hypodermis and other it issues is causing the phenotype . [Field: discussion, subscore: 4.00]
SECTION: discussion. Interestingly , past research has shown that the mor-1 gene also results in the rounded pharynx mouth phenotype seen in many of the short pharynx mutants and is located within the same region on chromosome III as PAS77 [ 21 ] . [Field: discussion, subscore: 4.00]
SECTION: discussion. While the short pharynx phenotype was classified as a pharynx that does not undergo proper pharyngeal elongation ; there was variability in the body length among these strains ranging from those with a normal body length to Dpy worms . [Field: discussion, subscore: 4.00]
SECTION: results. Other pharynx phenotypes Mutant PAS87 , PAS100 , PAS117 , PAS132 , PAS147 , PAS155 , and PAS197 are among the strains that manifest a tube-like pharynx ( Figure 1C , E , J , and not shown ) . [Field: results, subscore: 4.00]
SECTION: results. Compared to the organized and consistent wild-type pharynx , the nuclei appeared in random locations throughout the mutant pharynx ; however , counts of nuclei showed no significant difference between the numbers of muscle cells seen in the two phenotypes ( n = 19 mutant , n = 9 wild type , p = 0 . 82 ) . [Field: results, subscore: 4.00]
SECTION: results. In wild-type worms , the adherens junctions of the pharynx are clearly discernible from those in the Protein Class Ubiquinol cytochrome c reductase assembly protein RNA-binding protein UBA-like ubiquitin ligase Mitochondrial ribosomal protein bHLH transcription factor Uncharacterized conserved protein ATP-binding cassette ( ABC ) transporter Pharynx Phenotype Screen November 2011 | Volume 6 | Issue 11 | e26594 hypodermis and other areas because of increase intensity at the buccal cavity , metacorpus and terminal bulb in the wild-type worms ( Figure 3F ) ( n = 11 ) . [Field: results, subscore: 4.00]
SECTION: results. 0026594 . t003 PLoS ONE | www . plosone . org 5 Protein Class Frizzled family of seven transmembrane receptors Transcription factor Exocyst complex subunit Signal peptidase subunit Laminin alpha-2 High osmolarity signaling pathway Classical cadherin Splicing co-activator Spliceosomal protein Pharynx Phenotype Screen November 2011 | Volume 6 | Issue 11 | e26594 PAS136 mutants exhibit severe disorganization of the pharynx and appear to have misshapen cells . [Field: results, subscore: 4.00]
SECTION: results. III Short pharynx / thin pharynx PAS252 LG . [Field: results, subscore: 4.00]
SECTION: results. A mutant with pharynx muscle cells that do not appear to adhere PAS136 mutant progeny exhibit pharyngeal disorganization and misshapen cells in both anterior and posterior portions of the pharynx ( Figure 1I ) . [Field: results, subscore: 4.00]
SECTION: results. Complementation analysis of both PAS120 and PAS154 heterozygous worms with the homozygous sma-1 ( e30 ) worms produced 50 % mutant phenotypes in the F1 generation ( n = 99 ) , suggesting these mutant strains represent new sma-1 Pharynx Phenotype Screen PLoS ONE | www . plosone . org 2 November 2011 | Volume 6 | Issue 11 | e26594 Pharynx Phenotype Screen PLoS ONE | www . plosone . org 3 November 2011 | Volume 6 | Issue 11 | e26594 alleles . [Field: results, subscore: 4.00]
SECTION: results. A test of the MH27 adherens junction antibody ( AJM-1 ) staining showed the presence of normal junctions between epithelial cell types in both wild-type embryos and the pharynx of PAS77 embryos , although all regions of the pharynx are shorter in the anterior-posterior orientation ( Figure 2F , G ) . [Field: results, subscore: 4.00]
SECTION: results. Some of these short pharynx mutants were viable to the adult stage , while others died during larval development ; four of the short pharynx stains have been chromosomally mapped ( Table 2 ) . [Field: results, subscore: 4.00]
SECTION: results. In total , we identified 83 possible pharynx defective strains suggestive of abnormalities in cell adhesion , cell fate , cell morphology , and migration in both anterior and posterior pharynx regions ( Table 1 ) . [Field: results, subscore: 4.00]
SECTION: abstract. We observed over 83 C . elegans lines with distinctive pharyngeal phenotypes in worms surviving to the L1 larval stage , with phenotypes ranging from short pharynx , unattached pharynx , missing cells , asymmetric morphology , and non-adherent pharynx cells . [Field: abstract, subscore: 3.00]
SECTION: introduction. Thus , the initial goal of our mutagenesis screen was to isolate a gene necessary for either all pharynx muscle , or posterior pharynx muscle specification ; however , we instead found phenotypes that ranged from a short pharynx to pharynxes that are barely distinguishable . [Field: introduction, subscore: 3.00]
SECTION: introduction. We observed over 83 C . elegans lines with distinctive pharyngeal phenotypes in worms surviving to the L1 larval stage , with phenotypes ranging from short pharynx , unattached pharynx , missing cells , asymmetric morphology , and non-adherent pharynx cells . [Field: introduction, subscore: 3.00]
SECTION: abstract. We have also identified new alleles of sma-1 , and our screen suggests many genes directing pharynx assembly and structure may be either pharynx specific or less critical in other it issues . [Field: abstract, subscore: 2.00]
SECTION: discussion. In contrast , the myo-2 : : GFP strain used in our study fluoresced brightly and provided a very distinctive pharynx shape that could be recognized as wild type of mutant immediately . [Field: discussion, subscore: 2.00]
SECTION: discussion. In both cases , the screens were not targeting detecting diverse pharynx phenotypes . [Field: discussion, subscore: 2.00]
SECTION: discussion. Although alleles of genes such as pha-1 , pha-2 and pha-3 , and pha-4 were found in previous mutagenesis screens ; many of these alleles were isolated in screens that did not focus on pharynx structure [ 5 , 9 , 19 ] . [Field: discussion, subscore: 2.00]
SECTION: discussion. Counting the nuclei using myo-2 : : GFP : : H2B allowed us to determine if the PAS136 mutant worms were missing pharynx muscle cells compared to wild-type worms . [Field: discussion, subscore: 2.00]
SECTION: discussion. These basal domains of the apical junctions are known to regulate adhesion in the pharynx [ 31 ] . [Field: discussion, subscore: 2.00]
SECTION: discussion. Although complementation of PAS136 and lam-3 worms and DNA sequence of the PAS136 lam-3 allele does not support lam-3 as the cause of the disorganized pharynx phenotype ; it is possible another nearby gene involved in extracellular matrix organization may be involved . [Field: discussion, subscore: 2.00]
SECTION: discussion. The PAS136 amorphous pharynx phenotype We hypothesize the extreme disorganization of the pharyngeal muscles and marginal cells of PAS136 homozygous L1 progeny are caused by a mutation in a gene responsible for morphology and cell adhesion . [Field: discussion, subscore: 2.00]
SECTION: discussion. Sequence analysis confirmed PAS154 is a novel allele mutant for bH- spectrin , previously documented alleles of sma-1 share the sma- 1 ( lfc1 ) phenotype of a small body phenotype and short , fat pharynx [ 23 ] . [Field: discussion, subscore: 2.00]
SECTION: discussion. Even more , these mutants were chromosomally mapped to chromosome V . Worms with a loss-of-function mutation for sma-1 manifest a short pharynx phenotype similar to that of PAS120 and PAS154 . [Field: discussion, subscore: 2.00]
SECTION: discussion. The PAS154 mutant phenotype is a result of a new allele of sma-1 Through our forward genetic screen we also isolated two other mutants that exhibit similar short pharynx phenotypes : PAS120 and PAS154 . [Field: discussion, subscore: 2.00]
SECTION: discussion. 0026594 . g004 Pharynx Phenotype Screen PLoS ONE | www . plosone . org 7 November 2011 | Volume 6 | Issue 11 | e26594 For instance , pha-2 mutants ( pha-2 encodes a homeodomain transcription factor protein ) manifest a short isthmus [ 26 ] . [Field: discussion, subscore: 2.00]
SECTION: discussion. ( A ) PAS77 mutant phenotype with poorly defined pharynx regions . [Field: discussion, subscore: 2.00]
SECTION: discussion. Moreover , seeing that the pharynx is essential for the grinding and ingestion of food , it is possible that PAS77 dies due to starvation . [Field: discussion, subscore: 2.00]
SECTION: discussion. Also , PAS77 usually developmentally arrests and dies during the L1 stage of development , because while the overall proportions of the pharynx are consistent , the length is greatly diminished . [Field: discussion, subscore: 2.00]
SECTION: discussion. We analyzed the short pharynx mutant strain PAS77 , which exhibits defects in the metacorpus , procorpus , and manifests a short isthmus . [Field: discussion, subscore: 2.00]
SECTION: discussion. Surprisingly , 20 of the 83 pharyngeal mutants were homozygous recessive mutants that expressed the short pharynx and rounded mouth phenotype . [Field: discussion, subscore: 2.00]
SECTION: introduction. This report provides an overview of the results of the screen , and focuses on the mapping and PLoS ONE | www . plosone . org 1 November 2011 | Volume 6 | Issue 11 | e26594 00040399 characterization of short pharynx mutants and non-adherent muscle cell pharynx mutants . [Field: introduction, subscore: 2.00]
SECTION: introduction. The ability to use Green Fluorescent Protein ( GFP ) expressed specifically in pharynx muscle of larva has simplified the screening for mutations that affect pharynx structure in live worms . [Field: introduction, subscore: 2.00]
SECTION: introduction. Many previously described pharynx genes have been found using genetic screens , including alleles of genes pha-1 , pha-2 , pha-3 , and pha-4 ; however , these screens were not optimized for screening of morphological changes in pharynx muscle [ 5 , 9 , 19 ] . [Field: introduction, subscore: 2.00]
SECTION: introduction. Interestingly , the posterior sets of pharynx muscle cells derived from the MS blastomere form normally in the absence of TBX-2 and non-muscle ABa derived pharynx does not appear to require TBX-2 function [ 14 ] . [Field: introduction, subscore: 2.00]
SECTION: introduction. Multiple genes have been identified that are expressed in distinct pharyngeal cell types , such as myo-2 and ceh-22 in pharynx muscle and intermediate filaments in marginal cells ; however only tbx-2 is essential to specify a particular cell fate , in this case , anterior ABa derived pharynx muscle cells [ 6 , 14 , 15 , 16 , 17 , 18 ] . [Field: introduction, subscore: 2.00]
SECTION: introduction. In the cases of pha-4 , glp-1 , tbx-35 , and lag-1 , the loss of pharynx cells is not cell-type specific , rather entire regions of the pharynx are deleted such as the anterior ABa derived-pharynx in glp-1 mutants [ 5 , 8 , 10 , 11 , 12 , 13 ] . [Field: introduction, subscore: 2.00]
SECTION: introduction. While less dramatic , mutations in glp-1 , tbx-35 , or lag-1 result in a loss of all pharynx cells derived from ABa or MS lineage , resulting in formation of a half pharynx . [Field: introduction, subscore: 2.00]
SECTION: introduction. If pha-4 expression is eliminated through mutation or RNA interference , the entire pharynx fails to develop ; ectopic expression of pha-4 in early embryos converts additional cells to become pharynx cells [ 5 , 8 ] . [Field: introduction, subscore: 2.00]
SECTION: introduction. For example , eight different classes of pharynx muscle differ in morphology , producing the distinct bilobed pharynx that enables the worm to pump bacteria from the environment and pulverize this food before it passes into the intestine . [Field: introduction, subscore: 2.00]
SECTION: introduction. We have also identified new alleles of sma-1 , and our screen suggests many genes directing pharynx assembly and structure may be either pharynx specific or less critical in other it issues . [Field: introduction, subscore: 2.00]
SECTION: references. Thatcher JD , Fernandez AP , Beaster-Jones L , Haun C , Okkema PG ( 2001 ) The Caenorhabditis elegans peb-1 gene encodes a novel DNA-binding protein Pharynx Phenotype Screen PLoS ONE | www . plosone . org 10 November 2011 | Volume 6 | Issue 11 | e26594 involved in morphogenesis of the pharynx , vulva , and hindgut . [Field: references, subscore: 2.00]
SECTION: results. The PAS117 , PAS147 , PAS157 , PAS158 , PAS163 , and PAS236 mutant phenotypes show extensive loss of anterior pharynx muscle morphology ( Figure 1E , J , L , M , P , T ) . [Field: results, subscore: 2.00]
SECTION: results. III ; however , some progeny also demonstrated the short pharynx phenotype ( Figure 1V , W ) . [Field: results, subscore: 2.00]
SECTION: results. PAS147 lacks of a distinct anterior bulb , but otherwise has generally symmetric pharynx morphology ( Figure 1J ) . [Field: results, subscore: 2.00]
SECTION: results. However , PAS252 mutant worms with a very similar phenotype did not have beads present in either their pharynx or intestine ( Figure 4C , D ) . [Field: results, subscore: 2.00]
SECTION: results. PAS77 phenotype worms that fed on Fluoresbrite beads for two hours exhibited beads in both their pharynx and intestine , demonstrating their ability to ingest food normally ( Figure 4A , B ) . [Field: results, subscore: 2.00]
SECTION: results. PAS136 mutants have a wild type number of pharynx muscle cells The mutant worms of the PAS136 strains appear to have gaps between muscle cells that are not normally present , which could be the result of some muscle cells not be present at all . [Field: results, subscore: 2.00]
SECTION: results. ( G ) PAS136 embryo with weak and disconnect AJM-1 staining in the pharynx ( ph ) and more normal AJM-1 in the intestine ( it ) . [Field: results, subscore: 2.00]
SECTION: results. ( F ) Wild-type MH27 AJM-1 adherens junction antibody staining showing pharynx ( ph ) and intestine ( it ) localization . [Field: results, subscore: 2.00]
SECTION: results. ( E ) hmr-1 ( W02B9 . 1 ) RNAi results in a Pun phenotype with diminished anterior pharynx cells ( arrow ) . [Field: results, subscore: 2.00]
SECTION: results. ( D ) blmp-1 ( F25D7 . 3 ) RNAi has a less severe PAS136 phenotype ( arrow denotes cell disconnected from the pharynx ) . [Field: results, subscore: 2.00]
SECTION: results. ( B ) pha-4 RNAi used a positive control for pharynx phenotypes , arrow shows lack of myo-2 : : GFP in most of the head . [Field: results, subscore: 2.00]
SECTION: results. ( A ) Probable location of the PAS136 pharynx phenotype allele is between 6 cM and 8 cM on LG . [Field: results, subscore: 2.00]
SECTION: results. PAS136 mapping and pharynx markers . [Field: results, subscore: 2.00]
SECTION: results. ( G ) PAS77 MH27 antibody staining shows four compressed pharynx regions ( arrows ) . [Field: results, subscore: 2.00]
SECTION: results. ( F ) wild-type MH27 AJM-1 adherens junction antibody staining shows four distinct regions in the pharynx ( arrows ) . [Field: results, subscore: 2.00]
SECTION: results. ( E ) xbp-1 ( R74 . 3 ) RNAi eliminates GFP expression in pharynx . [Field: results, subscore: 2.00]
SECTION: results. ( C and D ) C35D10 . 5 RNAi or M88 . 2 RNAi results in a short pharynx phenotype . [Field: results, subscore: 2.00]
SECTION: results. Arrow shows region missing anterior pharynx cells . [Field: results, subscore: 2.00]
SECTION: results. ( A ) Probable location of the PAS77 pharynx phenotype allele is between 24 . 47 cM and 23 . 1 cM relative to the genetic center of LG . [Field: results, subscore: 2.00]
SECTION: results. PAS77 mapping and pharynx markers . [Field: results, subscore: 2.00]
SECTION: results. 0026594 . g001 Pharynx Phenotype Screen PLoS ONE | www . plosone . org 4 November 2011 | Volume 6 | Issue 11 | e26594 Figure 2 . [Field: results, subscore: 2.00]
SECTION: results. ( X ) PAS252 short pharynx . doi : 10 . 1371 / journal . pone . [Field: results, subscore: 2.00]
SECTION: results. ( W ) PAS241 short pharynx . [Field: results, subscore: 2.00]
SECTION: results. ( U ) PAS237 pharynx muscle cells unattached . [Field: results, subscore: 2.00]
SECTION: results. ( R ) PAS196 short pharynx . [Field: results, subscore: 2.00]
SECTION: results. ( P ) PAS163 disorganized pharynx with unattached GFP-expressing cells . [Field: results, subscore: 2.00]
SECTION: results. ( O ) PAS159 short pharynx . [Field: results, subscore: 2.00]
SECTION: results. ( N ) PAS158 Diminished GFP expression and asymmetric anterior pharynx . [Field: results, subscore: 2.00]
SECTION: results. ( L and M ) PAS157 pharynx asymmetry with indistinct isthmus and anterior bulb . [Field: results, subscore: 2.00]
SECTION: results. ( K ) PAS154 short pharynx phenotype . [Field: results, subscore: 2.00]
SECTION: results. ( J ) PAS147 cylindrical pharynx with diminished anterior bulb . [Field: results, subscore: 2.00]
SECTION: results. ( I ) PAS136 pharynx muscle cells do not adhere to each other . [Field: results, subscore: 2.00]
SECTION: results. ( H ) PAS129 short pharynx with head defects . [Field: results, subscore: 2.00]
SECTION: results. ( G ) PAS126 short pharynx . [Field: results, subscore: 2.00]
SECTION: results. ( F ) PAS120 short pharynx and bulbous head . [Field: results, subscore: 2.00]
SECTION: results. ( D ) PAS101 pharynx unattached . [Field: results, subscore: 2.00]
SECTION: results. ( C ) PAS100 thin pharynx with less anterior GFP expression than wild type . [Field: results, subscore: 2.00]
SECTION: results. ( B ) PAS77 short pharynx phenotype . [Field: results, subscore: 2.00]
SECTION: results. X Short pharynx doi : 10 . 1371 / journal . pone . [Field: results, subscore: 2.00]
SECTION: results. X Short pharynx PAS262 LG . [Field: results, subscore: 2.00]
SECTION: results. IV Anterior pharynx defects PAS202 LG . [Field: results, subscore: 2.00]
SECTION: results. IV Short pharynx PAS170 LG . [Field: results, subscore: 2.00]
SECTION: results. I Amorphous pharynx PAS154 LG . [Field: results, subscore: 2.00]
SECTION: results. I Amorphous pharynx PAS138 LG . [Field: results, subscore: 2.00]
SECTION: results. V Thin , cylindrical pharynx PAS136 LG . [Field: results, subscore: 2.00]
SECTION: results. IV Short pharynx PAS132 LG . [Field: results, subscore: 2.00]
SECTION: results. III Short pharynx PAS100 LG . [Field: results, subscore: 2.00]
SECTION: results. Summation of pharynx phenotypes . [Field: results, subscore: 2.00]
SECTION: results. A partially penetrant phenotype similar to that of PAS136 resulted from lam-3 ( T22A3 . 8 ) RNAi and blmp-1 ( F25D7 . 3 ) RNAi ( Figure 3C and D ) ; the laminin gene lam-3 has previously been shown to affect the extracellular matrix and pharynx cohesion , while blmp-1 is described as a homolog of the B lymphocyte-induced maturation protein 1 on WormBase [ 24 , 25 ] . [Field: results, subscore: 2.00]
SECTION: results. Within the region of PAS136 mapping , there are 322 genes ; 39 were chosen for screening using recorded L1 arrest as a necessary parameter ; experiments showing pharynx phenotypes are listed ( Table 4 ) . [Field: results, subscore: 2.00]
SECTION: results. To screen genes in the mapped region as candidates for PAS136 , we used RNAi to screen for phenocopy of the pharynx phenotype with bacterial feeding RNAi using the PD4792 myo- 2 : : GFP reporter strain . [Field: results, subscore: 2.00]
SECTION: results. There are also gaps between pharynx muscle cells , which are not normally present . [Field: results, subscore: 2.00]
SECTION: results. Finally , the PAS241 phenotype results in an extremely wide head and Dpy body with an extremely short pharynx lacking distinct regions ( Figure 1W ) . [Field: results, subscore: 2.00]
SECTION: results. PAS126 , PAS129 , PAS159 , PAS196 , and PAS241 also exhibit the short pharynx phenotype ; although the primary the defect is a failure for the procorpus to elongate ( Figure 1G , H , O , R , W ) . [Field: results, subscore: 2.00]
SECTION: results. V and encodes a spectrin homolog with a similar pharynx phenotype [ 23 ] . [Field: results, subscore: 2.00]
SECTION: results. The strains PAS120 and PAS154 also share the short pharynx phenotype , however , despite their morphological defects , both are viable and fertile as homozygotes ( Figure 1F , K ) . [Field: results, subscore: 2.00]
SECTION: results. The MH4 monoclonal antibody to recognize intermediate filament shows that marginal cells are present , and appear similar to wild type ( Figure 2H ) other than for the compressed shape of the pharynx ( Figure 2I ) . [Field: results, subscore: 2.00]
SECTION: results. While not a PAS77 phenotype , the R74 . 3 ( xpb-1 ) dsRNA bacterial vector resulted in a nearcomplete absence of myo-2 : : GFP without apparent loss of pharynx muscle as seen in DIC imaging ( Figure 2E ) . [Field: results, subscore: 2.00]
SECTION: results. RNAi of M88 . 2 , a mitochondrial ribosomal protein , also produced L1 arrested larvae with a short pharynx phenotype ( Figure 2D ) . [Field: results, subscore: 2.00]
SECTION: results. The C35D10 . 5 RNAi resulted in many embryos with a short pharynx phenotype ; interestingly , C35D10 . 5 encodes a predicted ubiquinol cytochrome c reductase assembly protein ( Figure 2C ) . [Field: results, subscore: 2.00]
SECTION: results. Multiple RNAi bacterial strains resulted in pharynx phenotypes ( Table 3 ) . [Field: results, subscore: 2.00]
SECTION: results. glp-1 was used as a positive control for this experiment because it has a dramatic Anterior Pharynx Defective ( Aph ) phenotype that is easily identified in the PD4792 strain ( Figure 2B ) . [Field: results, subscore: 2.00]
SECTION: results. The short-pharynx phenotypes We identified 20 mutant lines that shared a similar phenotypic trait , a short pharynx and rounded mouth similar to mutant strain PAS77 ; nine are shown in Figure 1 ( Figure 1B , F , G , H , K , O , R , W , X ) . [Field: results, subscore: 2.00]
SECTION: results. Initially , the low-copy number myo-2 : : GFP ( AZ217 ) integrated reporter strain was used in mutagenesis ; however , the strain ' s weak fluorescence made rapid identification of pharynx abnormalities difficult under an epifluorescent stereomicroscope . [Field: results, subscore: 2.00]
SECTION: results. We performed an EMS mutagenesis screen of , 10 , 000 haploid genomes to identify the genes affecting posterior muscle fate in C . elegans , using a myo-2 : : GFP reporter to visualize pharynx morphology in L1s . [Field: results, subscore: 2.00]
SECTION: title. Multiple phenotypes resulting from a mutagenesis screen for pharynx muscle mutations in Caenorhabditis elegans . [Field: title, subscore: 2.00]
SECTION: abstract. Our studies have focused on genetically mapping and functionally testing two phenotypes , the short pharynx and the loss of muscle cohesion phenotypes . [Field: abstract, subscore: 1.00]
SECTION: introduction. No gene has been found that is specifically required for posterior pharynx muscle specification . [Field: introduction, subscore: 1.00]
SECTION: introduction. The pha-1 gene allows for initial development of pharyngeal precursor cells , but then affects differentiation of all pharynx cells types after the 1K-fold stage of embryogenesis when differentiation markers such as pharyngeal myosin and intermediate filaments are normally activated [ 9 ] . [Field: introduction, subscore: 1.00]
SECTION: introduction. In C . elegans , pha-4 is an organ identity gene involved in the specification and differentiation of all cells destined to become the pharynx [ 5 , 6 , 7 ] . [Field: introduction, subscore: 1.00]
SECTION: introduction. Seven different cell types are specified during pharynx organogenesis ; and within these cell types , sub-specialization occurs producing distinct anterior to posterior characteristics [ 4 ] . [Field: introduction, subscore: 1.00]
SECTION: introduction. The C . elegans pharynx exhibits progressive restriction of cell fate during development , ultimately resulting in the expression of differentiation factors and structural proteins essential to its function as a neuromuscular pump [ 1 , 2 , 3 ] . [Field: introduction, subscore: 1.00]
SECTION: introduction. Citation : Ferrier A , Charron A , Sadozai Y , Switaj L , Szutenbach A , et al . ( 2011 ) Multiple Phenotypes Resulting from a Mutagenesis Screen for Pharynx Muscle Mutations in Caenorhabditis elegans . [Field: introduction, subscore: 1.00]
SECTION: introduction. Our studies have focused on genetically mapping and functionally testing two phenotypes , the short pharynx and the loss of muscle cohesion phenotypes . [Field: introduction, subscore: 1.00]
SECTION: introduction. Multiple Phenotypes Resulting from a Mutagenesis Screen for Pharynx Muscle Mutations in Caenorhabditis elegans Andrew Ferrier , Alexandra Charron , Yama Sadozai , Lynn Switaj , Anneliese Szutenbach , Pliny A . Smith * Biology Department , Lake Forest College , Lake Forest , Illinois , United States of America , Abstract [Field: introduction, subscore: 1.00]
SECTION: materials. Pharynx muscle nuclei counts Nuclearly localized myo-2 : : GFP worms were maintained by microinjecting 0 . 05 ng / mL myo-2 : : GFP : : His2B , 40 ng / ml pRF4 , and 60 ng / ml herring sperm DNA into N2 hermaphrodite and screening for rolling progeny . [Field: materials, subscore: 1.00]
SECTION: materials. Sequence data was assem- Pharynx Phenotype Screen PLoS ONE | www . plosone . org 9 November 2011 | Volume 6 | Issue 11 | e26594 bled into a contig and compared to wild type sequences in WormBase ( wormbase . org ) [ 20 ] . [Field: materials, subscore: 1.00]
SECTION: references. Pharynx Phenotype Screen PLoS ONE | www . plosone . org 11 November 2011 | Volume 6 | Issue 11 | e26594 [Field: references, subscore: 1.00]
SECTION: references. Jafari G , Burghoorn J , Kawano T , Mathew M , Morck C , et al . ( 2010 ) Genetics of extracellular matrix remodeling during organ growth using the Caenorhabditis elegans pharynx model . [Field: references, subscore: 1.00]
SECTION: references. Albertson DG , Thomson JN ( 1976 ) The pharynx of Caenorhabditis elegans . [Field: references, subscore: 1.00]
Supplemental links/files: reference in endnote reference in xml Wormbase reference
Score: 195.00
Title: Organ Length Control by an ADAMTS Extracellular Protease in Caenorhabditis elegans .
Authors: Shibata Y ; Kawakado Y ; Hori N ; Tanaka K ; Inoue R ; Takano T ; Kubota Y ; Nishiwaki K
Journal: G3 ( Bethesda )
Year: 2016-03-18
Doc ID: WBPaper00049351
Bibliographic Information
Abstract
Matching Sentences
SECTION: abstract. The number of nuclei in the pharynx and the pumping rate of the pharynx were not affected in the mig-17 mutants , suggesting that the cells constituting the pharynx are elongated although the pharynx functions normally in these mutants . [Field: abstract, subscore: 4.00]
SECTION: discussion. If this is also the case in the larval pharynx , the active elongation of the larval body , or the elongation of the epidermis , should stretch the pharynx during its elongation . [Field: discussion, subscore: 4.00]
SECTION: discussion. Pharynx elongation in mig-17 mutants Our finding indicates that the pharynx of mig-17 mutants is elongated because of the elongation of cells rather than an increase in the cell number . [Field: discussion, subscore: 4.00]
SECTION: discussion. Interestingly , let-2 ( k196 ) and emb-9 ( b117 ) showed an elongated pharynx , whereas emb-9 ( b189 ) and emb-9 ( g34 ) had a shortened pharynx . [Field: discussion, subscore: 4.00]
SECTION: discussion. MIG-17 acts through different molecular mechanisms in pharynx and gonad development An analysis of molecules involved in the MIG-17 pathway that regulates DTC migration revealed distinct aspects of these molecules in the regulation of pharynx length . [Field: discussion, subscore: 4.00]
SECTION: discussion. These results suggest that MIG-17 negatively regulates pharynx elongation from the basal side of the pharynx . [Field: discussion, subscore: 4.00]
SECTION: discussion. DISCUSSION MIG-17 activity is involved in pharynx length regulation In this study , we showed that an ADAMTS family metalloprotease , MIG-17 , which is required for directional migration of gonadal DTCs , also acts in the regulation of pharynx length . [Field: discussion, subscore: 4.00]
SECTION: introduction. The number of nuclei in the pharynx , and the pumping rate of the pharynx , were not affected in cells constituting the pharynx are elongated , although the pharynx functions normally in these mutants . [Field: introduction, subscore: 4.00]
SECTION: results. The mig-6 ( k177 ) class-s mutants , however , showed a rather shortened pharynx ( Figure 9 , A and B ) , suggesting that mig-6 may not function with mig-17 to control the length of the pharynx . [Field: results, subscore: 4.00]
SECTION: results. Catalytic activity , disintegrin ( DI ) domain , and N-glycosylation are essential for pharynx length regulation To explore the function of individual domains of MIG-17 , as well as its glycosylation , in the regulation of pharynx length and DTC migration , we conducted transgenic rescue experiments of mig-17 ( k174 ) using extrachromosomal transgenic arrays ( Ihara and Nishiwaki 2007 ) . [Field: results, subscore: 4.00]
SECTION: results. We measured body length in addition to pharynx length to determine whether mutations affect pharyngeal length specifically or they affect overall size of the animal . Figure 1 Pharynx elongation phenotype and mutation sites of mig-17 mutants . [Field: results, subscore: 4.00]
SECTION: results. We found that the gene is also required to form a pharynx of the proper length : mig-17 mutants exhibited an elongated pharynx phenotype ( Figure 1A ) . [Field: results, subscore: 4.00]
SECTION: discussion. It is worth noting that MIG-17 , which acts in pharynx length regulation in C . elegans , has a structural similarity to ADAMTS10 and 17 in that they all share C-terminal protease and lacunin ( PLAC ) domains ( Ihara and Nishiwaki 2007 ) . [Field: discussion, subscore: 2.00]
SECTION: discussion. However , it is also possible that the larval pharynx expands actively rather than passively by stretching . [Field: discussion, subscore: 2.00]
SECTION: discussion. Loss of MIG-17 activity may enhance the adhesion and therefore confer a greater stretching force , resulting in pharynx elongation . [Field: discussion, subscore: 2.00]
SECTION: discussion. Because the pharyngeal end interacts with the surrounding epidermis through their basement membranes , MIG-17 may normally act to weaken the friction or adhesiveness between these basement membranes to produce a pharynx of proper length . [Field: discussion, subscore: 2.00]
SECTION: discussion. Because epidermal elongation is much greater than pharyngeal elongation , we should postulate some slippage between the pharynx end and the epidermis . [Field: discussion, subscore: 2.00]
SECTION: discussion. Recently , it was shown that the embryonic pharynx behaves like a spring in which the anterior end is attached to the mouth , and the posterior end attached to the surrounding epidermis ( Kelley et al . 2015 ) . [Field: discussion, subscore: 2.00]
SECTION: discussion. The pharynx is $ 60 mm long when worms hatch , and it elongates to $ 130 mm long by the adult stage ( about a twofold increase ) . [Field: discussion, subscore: 2.00]
SECTION: discussion. Thus , it might be possible that endostatin also acts in pharynx length control . [Field: discussion, subscore: 2.00]
SECTION: discussion. We found that cle-1 ( cg120 ) results in pharynx elongation similar to that in the emb-9 ( b117 ) and let-2 ( k196 ) mutants . [Field: discussion, subscore: 2.00]
SECTION: discussion. We speculate that MIG-17 may modulate the structure or conformation of the C-terminal half of the triple-helical domain to achieve a pharynx of the proper length . [Field: discussion, subscore: 2.00]
SECTION: discussion. Therefore it might be possible that glycine substitutions in the C-terminal high-stability region result in pharynx elongation , whereas those in the N-terminal low-stability region result in pharyngeal shortening . [Field: discussion, subscore: 2.00]
SECTION: discussion. We also showed in this study that mutations in type IV collagen affect pharynx length . [Field: discussion, subscore: 2.00]
SECTION: discussion. Volume 6 May 2016 | ADAMTS Control of Organ Length | 1455 Basement membrane collagens are involved in pharynx length regulation Basement membrane type IV collagen plays important roles in it issue morphogenesis ( Morrissey and Sherwood 2015 ) . [Field: discussion, subscore: 2.00]
SECTION: discussion. ( A ) , ( B ) Pharynx length ( A ) and body length ( B ) of young adult hermaphrodites were analyzed . [Field: discussion, subscore: 2.00]
SECTION: discussion. Figure 9 Effects of mutations affecting glycosylation and basement membrane components on pharynx length . [Field: discussion, subscore: 2.00]
SECTION: discussion. ( A ) , ( B ) Pharynx length ( A ) and body length ( B ) of young adult hermaphrodites were analyzed . [Field: discussion, subscore: 2.00]
SECTION: discussion. Figure 8 Effects of fbl-1 and nid-1 mutations on pharynx length . [Field: discussion, subscore: 2.00]
SECTION: discussion. Thus , it might be possible that qualitative and / or quantitative differences in NID-1expression may be the cause for distinct activities of NID-1 in the MIG-17dependent mechanisms in the pharynx and the gonad . [Field: discussion, subscore: 2.00]
SECTION: discussion. nid-1 appears to act antagonistically to mig-17 in the regulation of pharynx length . [Field: discussion, subscore: 2.00]
SECTION: discussion. Therefore , the different suppression outcomes by k196 in the formation of the pharynx and the gonad could reflect the different actions of MIG-17 in these basement membrane regions . [Field: discussion, subscore: 2.00]
SECTION: discussion. Although both let-2 alleles , k193 and k196 , strongly suppress DTC migration defects ( Kubota et al . 2008 ) , only k193 suppressed the pharynx phenotype of mig-17 . k193 and k196 are amino acid substitutions in the C-terminal noncollagenous ( NC1 ) domain and the triple helical domain of type IV collagen , respectively . [Field: discussion, subscore: 2.00]
SECTION: discussion. MIG-18 , a cofactor of MIG-17 in gonadal DTC migration , was not required for pharynx length regulation . [Field: discussion, subscore: 2.00]
SECTION: discussion. By expressing various mutant constructs in the mig-17 ( k174 ) null mutant , we found that the catalytic activity , the C-terminal DI domain and N-glycosylation of MIG-17 are essential for the regulation of pharynx length as in the case of the control of DTC migration . [Field: discussion, subscore: 2.00]
SECTION: discussion. ( A ) , ( B ) Pharynx length ( A ) and body length ( B ) of young adult hermaphrodites were analyzed . [Field: discussion, subscore: 2.00]
SECTION: discussion. Consistent with this observation , MIG-17-Venus strongly accumulated in the pharyngeal basement membrane during Figure 7 Effects of let-2 mutations on pharynx length . [Field: discussion, subscore: 2.00]
SECTION: discussion. We found that the pharynx of mig-17 mutants becomes longer compared with the wild type after the L2 stage . [Field: discussion, subscore: 2.00]
SECTION: introduction. Although pharynx length did not differ between wild-type and mig-17 mutant animals during the first larval stage , the mig-17 pharynx became longer than that of wild type until the adult stage . [Field: introduction, subscore: 2.00]
SECTION: materials. Pharynx volume was assessed by measuring the area of the sagittal optical section of the pharynx . [Field: materials, subscore: 2.00]
SECTION: results. These results indicate that mutations in basement membrane collagens affect the control of the pharynx and body length . [Field: results, subscore: 2.00]
SECTION: results. We also examined an allele of cle-1 , which encodes basement membrane type XVIII collagen ( Ackley et al . 2001 ) and observed a significant increase in the length of the pharynx , as was noted in the emb-9 ( b177 ) mutants ( Figure 9 , A and B ) . [Field: results, subscore: 2.00]
SECTION: results. The b117 allele produced a slightly elongated pharynx , whereas the b189 and g34 alleles had shorter pharynges and bodies compared with the wild type . [Field: results, subscore: 2.00]
SECTION: results. Mutations in basement membrane proteins affect the length of the pharynx and body mig-6 encodes the basement membrane protein papilin . [Field: results, subscore: 2.00]
SECTION: results. Although mig-23 , cogc-1 , and cogc-3 mutants partially compromise the function of MIG-17 in DTC migration through their misglycosylation ( Nishiwaki et al . 2004 ; Kubota et al . 2006 ) , it may still function adequately to control pharynx length . [Field: results, subscore: 2.00]
SECTION: results. Because of the shorter body length , it was not clear whether the other mutants affected pharynx length . [Field: results, subscore: 2.00]
SECTION: results. We examined mutant alleles of these genes , and found that each of these mutations resulted in a shorter body length relative to the wild type , and that a slight elongation of the pharynx was present only in mig-22 ( k141 ) animals ( Figure 9 , A and B ) . [Field: results, subscore: 2.00]
SECTION: results. Thus , it is likely that , although let-2 , fbl-1 and nid-1 act to control the length of the pharynx in the MIG-17 pathway , their roles are not always the same as their roles in the control of gonadal DTC migration . [Field: results, subscore: 2.00]
SECTION: results. Therefore , either a reduction or increase in the NID-1 dosage appears to suppress the mig-17 pharynx phenotype . [Field: results, subscore: 2.00]
SECTION: results. We examined pharynx length in a mig-17 ( k174 ) mutant that overexpresses NID-1-HA ( Figure 8 , A and B ) and observed partial suppression in the transgenic line . [Field: results, subscore: 2.00]
SECTION: results. When nid-1 mutants were then combined with mig-17 ( k174 ) ; fbl-1 ( k201 ) or mig-17 ; fbl-1 ( k206 ) , the mig-17 pharynx length defect was still suppressed . [Field: results, subscore: 2.00]
SECTION: results. These results suggest that MIG-18 is not required for the regulation of pharynx length , although it has a substantial function in DTC migration ( Kim et al . 2014 ) . [Field: results, subscore: 2.00]
SECTION: results. Volume 6 May 2016 | ADAMTS Control of Organ Length | 1453 of the pharynx in mig-17 ( k174 ) ; mig-18 ( k140 ) double mutants was elongated and comparable to that of the mig-17 ( k174 ) single mutants ( Figure 6 , D and E ) . [Field: results, subscore: 2.00]
SECTION: results. Pharynx length ( D ) and body length ( E ) of young adult hermaphrodites were analyzed . [Field: results, subscore: 2.00]
SECTION: results. ( D ) , ( E ) Effects of mig-18 mutations on pharynx length . [Field: results, subscore: 2.00]
SECTION: results. Figure 6 Localization of MIG-17 , FBL-1C and MIG-18 , and effects of mig-18 mutations on pharynx length . [Field: results, subscore: 2.00]
SECTION: results. ( A ) Pharynx length . [Field: results, subscore: 2.00]
SECTION: results. The length Figure 5 Transgenic rescue experiments of mig-17 pharynx and gonadal defects . [Field: results, subscore: 2.00]
SECTION: results. Thus , MIG-18 localized to the pharynx similar to MIG-17 . [Field: results, subscore: 2.00]
SECTION: results. MIG-18 , a cofactor of MIG-17 , is not required for the control of pharynx length MIG-18 is a novel secreted protein whose loss-of-function mutations result in DTC migration defects similar to those in the mig-17 mutations . [Field: results, subscore: 2.00]
SECTION: results. These results indicate that the catalytic activity , the DI domain , and N-glycan modifications are essential for regulating pharynx length . [Field: results, subscore: 2.00]
SECTION: results. Thus , mig-17 appears to control the length of the pharynx specifically . [Field: results, subscore: 2.00]
SECTION: results. We then examined whether mig-17 mutants affect the length of amphid sensory dendrites surrounding the pharynx . [Field: results, subscore: 2.00]
SECTION: results. Thus , it is likely that the length of cells constituting the pharynx is affected in the mutants . [Field: results, subscore: 2.00]
SECTION: results. We examined the number of nuclei in the pharynx in young adult animals expressing SUR-5-GFP and EMB-9-mCherry , markers for somatic nuclei and the basement membrane , respectively . [Field: results, subscore: 2.00]
SECTION: results. Nuclear numbers , pumping rates , and length of the amphid sensory dendrites are not affected in mig- 17 mutants The difference in pharynx length between the wild type and mutants could reflect a difference either in cell number or cell length . [Field: results, subscore: 2.00]
SECTION: results. Thus , it is possible that mig-17 function is required for pharynx length regulation after the L2 stage ( Figure 3C ) . [Field: results, subscore: 2.00]
SECTION: results. In both these alleles , pharynx elongation was observed later than the third larval ( L3 ) stage but not during the L2 stage . [Field: results, subscore: 2.00]
SECTION: results. To understand developmental stages when MIG-17 is required for normal pharynx length , we examined when the pharyngeal abnormality becomes evident in k135 and k174 mutant animals . [Field: results, subscore: 2.00]
SECTION: results. These results suggest that the pharynx in mig-17 mutants extends along the anteriorposterior axis , and decreases in width without affecting its volume . [Field: results, subscore: 2.00]
SECTION: results. We assessed pharynx volumes by examining the area of the sagittal optical section of wild type and mutant pharynges , and found no significant difference between them : 2354 6 93 mm2 ( n = 20 ) for wild type and 2377 6 129 mm2 ( n = 20 ) for mig-17 ( k174 ) ( P = 0 . 528 ) . [Field: results, subscore: 2.00]
SECTION: results. Pharynx length ( C ) and body length ( D ) . [Field: results, subscore: 2.00]
SECTION: results. ( A ) The regions of the pharynx are indicated . [Field: results, subscore: 2.00]
SECTION: results. ( AC ) Pharynx lengths ( n = 20 ) ( A ) , % DTC migration defects ( n = 60 ) ( B ) , and body length ( n = 20 ) ( C ) for each allele are shown . [Field: results, subscore: 2.00]
SECTION: results. The pharynx can be divided into three parts , the corpus , isthmus and terminal bulb , from anterior to posterior ( Figure 3A ) . [Field: results, subscore: 2.00]
SECTION: results. The putative null allele , k174 , which had the strongest effect on DTC migration , also exhibited the strongest phenotype with respect to pharynx length . [Field: results, subscore: 2.00]
SECTION: results. Dashed lines depict anterior and posterior ends of the pharynx . [Field: results, subscore: 2.00]
SECTION: results. mig-17 is required to achieve a pharynx of normal length mig-17 was originally identified through mutations affecting directional migration of gonadal leader cells , the DTCs , in C . elegans ( Nishiwaki et al . 2000 ) . [Field: results, subscore: 2.00]
SECTION: abstract. In contrast to the control of DTC migration , MIG-18 , a secreted cofactor of MIG-17 , was not essential for pharynx length regulation . [Field: abstract, subscore: 1.00]
SECTION: abstract. We found that the pharynx was elongated in mig-17 mutants compared with wild type . [Field: abstract, subscore: 1.00]
SECTION: abstract. Here we show that MIG-17 is also required for the control of pharynx elongation during animal growth . [Field: abstract, subscore: 1.00]
SECTION: introduction. Genetic analyses of genes that act in the MIG-17 pathway to regulate DTC migration , most of which are basement membrane proteins , revealed shared and organ-specific aspects of the molecular mechanisms regulating DTC migration and pharynx length . [Field: introduction, subscore: 1.00]
SECTION: introduction. We showed that the function of MIG-17 was as important for the control of pharynx length as for its function in DTC migration . [Field: introduction, subscore: 1.00]
SECTION: introduction. The pharynx is an epithelial organ surrounded by a basement membrane . [Field: introduction, subscore: 1.00]
SECTION: introduction. The pharynx is a complex organ made up of 62 cells , which includes epithelial cells , muscle cells , marginal cells , grand cells , and neurons ( Altun and Hall 2009 ) . [Field: introduction, subscore: 1.00]
SECTION: introduction. Here we report that mig-17 functions in the control of pharynx length . [Field: introduction, subscore: 1.00]
SECTION: introduction. E-mail : nishiwaki @ kwansei . ac . jp INVESTIGATION Department of MIG-17 localized to the mig-17 mutants , suggesting that MIG-17 , was not essential for MIG-17 involving LET-2 / type IV colla- KEYWORDS organ elongation ADAMTS protease basement membrane pharynx ADAMTS family proteins have received considerable attention because defects in many of these proteins have been linked to hereditary diseases related to disorders affecting the extracellular matrix ( ECM ) ( Dubail and Apte 2015 ; Hubmacher and Apte 2015 ) . [Field: introduction, subscore: 1.00]
SECTION: introduction. In contrast to the control of DTC migration , MIG-18 , a secreted cofactor of pharynx length regulation . [Field: introduction, subscore: 1.00]
SECTION: introduction. We found that the pharynx was elongated in mig-17 mutants compared with wild type . pharyngeal basement membrane as well as to the gonadal basement membrane . [Field: introduction, subscore: 1.00]
SECTION: introduction. Here , we show that MIG-17 is also required for the control of pharynx elongation during animal growth . [Field: introduction, subscore: 1.00]
SECTION: materials. Pharynx length was assessed by measuring the length of the pharyngeal region excepting the buccal cavity . [Field: materials, subscore: 1.00]
SECTION: materials. Pharynx and body length were analyzed using Image J software . [Field: materials, subscore: 1.00]
SECTION: materials. Microscopy Pharynx length , body length and gonad migration phenotypes were determined using a Nomarski microscope ( Axioplan 2 ; Zeiss ) . [Field: materials, subscore: 1.00]
Supplemental links/files: reference in endnote reference in xml Wormbase reference
Score: 193.00
Title: The twisted pharynx phenotype in C . elegans .
Authors: Axang C ; Rauthan M ; Hall DH ; Pilon M
Journal: BMC Dev Biol
Year: 2007
Doc ID: WBPaper00030736
Bibliographic Information
Abstract
Matching Sentences
SECTION: discussion. This result also suggests that the pharyngeal cuticle is not directly restraining an intrinsic twist . We also found that the attachmentof the pharynx to the bodywall at the lips is not preventing the pharynx from twisting in L1 larvae ( Fig . 8 ) , and that the pharyngeal tendons that connect the procorpus to the bodywall appear intact , albeit spirally oriented , in animals with a twisted pharynx ( Fig . 4 ) . [Field: discussion, subscore: 6.00]
SECTION: discussion. Anterior detachment of the pharynx does not result in increased or earlier pharyngeal twisting Figure 8 Anterior detachment of the pharynx does not result in increased or earlier pharyngeal twisting . [Field: discussion, subscore: 4.00]
SECTION: results. Neither of these phenotypes was associated with pharyngeal twist- The anterior part of the intestine is twisted in a direction opposite to that of the pharynx Figure 6 The anterior part of the intestine is twisted in a direction opposite to that of the pharynx . [Field: results, subscore: 4.00]
SECTION: results. ( F ) and ( H ) : geometry of the tendons ( brown ) and pharynx ( blue circle ) viewed in cross sections if the pharynx is not twisted ( F ) or if it were twisted as a whole while held by the tendons ( H ) . [Field: results, subscore: 4.00]
SECTION: results. Of particular importance is that the pharynx seems to float in pseudocoelomic fluid and to make almost no contact with the worm body along its entire length : except for the tendons , the pharynx is secured only at its anterior and posterior ends , where it is connected to the mouth and intestine , respectively . [Field: results, subscore: 4.00]
SECTION: results. Anatomical features of the head region and comparison of the hemicentinrich pharyngeal tendons between wild-type and a mutant with the twisted pharynx phenotype Figure 4 Anatomical features of the head region and comparison of the hemicentin-rich pharyngeal tendons between wild-type and a mutant with the twisted pharynx phenotype . [Field: results, subscore: 4.00]
SECTION: results. BMC Developmental Biology 2007 , 7 : 61 http : / / www . biomedcentral . com / 1471-213X / 7 / 61 Page 6 of 13 ( page number not for citation purposes ) The anterior pharynx is firmly anchored to the lips by intercellular junctions , and more tenuously , via buccal cuticle Besides the tendon contacts along its anterior length , the pharynx is alsoconnected directly to the body-wall at its extreme anterior end , at the lips [ 1 ] . [Field: results, subscore: 4.00]
SECTION: results. him-4 mutants , with defective hemicentin and hence lacking these tendons [ 2 ] do not twist , indicating that lack of the support provided by the tendons is not enough to cause a twist . Also , these tendons do not seem to be involved in transferring a twisting force from outside the pharynx since mnm-4 ; him-4 double mutants still have a twisted pharynx ( data not shown ) . [Field: results, subscore: 4.00]
SECTION: results. BMC Developmental Biology 2007 , 7 : 61 http : / / www . biomedcentral . com / 1471-213X / 7 / 61 Page 5 of 13 ( page number not for citation purposes ) The pharyngeal tendons are pulled by the twisted pharynges The basal surface of the pharynx is covered by a thick basal lamina ( See Fig . 4AC ) , and four rows of 1520 acellular tendinous organs composed of hemicentin and fibulin that anchor the anterior end of the pharynx to the basal lamina that covers the bodywall muscles . [Field: results, subscore: 4.00]
SECTION: results. The twisted pharynx phenotype in mnm-4 ; etIs2 worms of different stages , and in adult mnm-4 ; unc-61 Figure 1 The twisted pharynx phenotype in mnm-4 ; etIs2 worms of different stages , and in adult mnm-4 ; unc-61 . [Field: results, subscore: 4.00]
SECTION: results. Note that the control pharynx of wild-type worms carrying the etIs2 integrated sequence is straight , and that the isolated pharynges of mnm-4 ; etIs2 , mnm-4 , mig-4 and unc-61 worms are twisted whereas the pharynx of the dig-1 animal is untwisted . [Field: results, subscore: 4.00]
SECTION: results. The twisted pharynx is fully functional Twisted pharynges performed as well as wild-type pharynges in bead uptake assays designed to measure feeding : there was a similar steady state number of beads within the pharynx ( i . . beads did not become stuck in twisted pharynges ) and a similar rate of transfer to the intestine ( Fig . 3 ) . [Field: results, subscore: 4.00]
SECTION: results. Quantitation of the twisted pharynx phenotype By measuring the torsion lines seen in a twisted pharynx , one can calculate the degree of twist ( Fig . 1A ) . [Field: results, subscore: 4.00]
SECTION: conclusion. The twisted pharynx phenotype is not a result of pumping activity nor the consequence of an external twisting force applied to the pharynx : it results from a twisting force intrinsic to the pharynx . [Field: conclusion, subscore: 3.00]
SECTION: introduction. The pharynx is sometimes considered to be evolutionarily related to the heart because : ( i ) like the heart , the pharynx is a rhythmically contracting muscular pump [ 3 ] ; ( ii ) the muscle cells of the pharynx have autonomous contractile activity reminiscent of cardiac myocytes [ 4 ] ; and ( iii ) ceh-22 , the C . elegans homolog to the homeobox gene Nk2x . [Field: introduction, subscore: 3.00]
SECTION: discussion. The twisted pharynx is a useful and easy-to-score phenotype , allowing one to screen for new genes involved in extracellular adhesion or organ cohesion . [Field: discussion, subscore: 2.00]
SECTION: discussion. The unc-61 ; mnm-4 double mutant did not show any increase in twisting of the pharynx nor any additive effect on the gonad phenotype ( unpublished observations ) : it is possible that unc-61 and mnm-4 may function in the same pathway . [Field: discussion, subscore: 2.00]
SECTION: discussion. However , only unc-61 displays the twisted pharynx phenotype . [Field: discussion, subscore: 2.00]
SECTION: discussion. Septins are conserved GTP-binding proteins that polymerize to form filamentous structures that act as scaffolds for membrane- and cytoskeleton-binding proteins [ 31 ] , a function consistent with our hypothesis that defects in BMC Developmental Biology 2007 , 7 : 61 http : / / www . biomedcentral . com / 1471-213X / 7 / 61 Page 10 of 13 ( page number not for citation purposes ) cytoskeleton anchorage points is responsible for the twisted pharynx phenotype . [Field: discussion, subscore: 2.00]
SECTION: discussion. It is likely that its other morphological defects , such as the twisted pharynx , also are caused by impaired adhesion . [Field: discussion, subscore: 2.00]
SECTION: discussion. Importantly , all three molecules are expressed in the pharynx [ 9 , 26 , 27 ] . [Field: discussion, subscore: 2.00]
SECTION: discussion. Interference with normal regulation of actin filament length could produce the observed abnormally long filaments ( Fig . 5 ) and also explain a steadily increasing force that could be relieved by gradualtwisting of the whole pharynx to lengthen each sarcomere . [Field: discussion, subscore: 2.00]
SECTION: discussion. This action normally should not introduce any twisting forces , and our observations that the pharynges of 6-day old adult mnm-4 mutants twist no more than that of 1-day old adults , and that mnm-4 L1 larvae kept in M9 for 72 hours show no twist indicate that pumping , at least if not accompanied by growth , plays a minor role , if any , in the development of the pharyngeal twist . Twisting occurs very gradually as the animal and pharynx grows . [Field: discussion, subscore: 2.00]
SECTION: discussion. As the pharynx rhythmically contracts , the actomyosin array foreshortens radially to expand and contract the pharyngeal lumen . [Field: discussion, subscore: 2.00]
SECTION: discussion. The pharynx features two prominent cytoskeletal filamentous arrays , both of which are oriented in a radial fashion . [Field: discussion, subscore: 2.00]
SECTION: discussion. Therefore the phenotype likely originates from a relatively minor intrinsic defect within the mutant pharynx . [Field: discussion, subscore: 2.00]
SECTION: discussion. Thus it seems that the twisted pharynx of the studied mutants is functional and properly anchored to the rest of the worm body . [Field: discussion, subscore: 2.00]
SECTION: discussion. We examined several parameters relevant to understanding the twisted pharynx phenotype . [Field: discussion, subscore: 2.00]
SECTION: discussion. Arrows point to the anterior end of the pharynx . [Field: discussion, subscore: 2.00]
SECTION: discussion. Shown are DIC images of L1 larvae of the indicated genotypes and in which the pharynx has detached from the lips and moved posteriorly . [Field: discussion, subscore: 2.00]
SECTION: discussion. Pharynx regions are as follows ( metacorpus in A ; isthmus in B ; isthmus in C and D ) . [Field: discussion, subscore: 2.00]
SECTION: discussion. The basal lamina of the ectodermal cells can not be easily distinguished from this flocculent matrix , whereas the very thick basal lamina of the pharynx is now visible as a prominent electron dense line . [Field: discussion, subscore: 2.00]
SECTION: discussion. Radial muscle filaments have developed and the pharynx is now contractile . [Field: discussion, subscore: 2.00]
SECTION: discussion. Transverse thin section through the head indicates the region between the pharynx ( Ph ) and bodywall where basal lamina deposition is taking place ( arrowheads point to plasma membrane of body-wall ectodermal cells in each panel ) . [Field: discussion, subscore: 2.00]
SECTION: discussion. Discussion The twisted pharynx is a viable phenotype that worsens during post-embryonic development . [Field: discussion, subscore: 2.00]
SECTION: results. The dashed line indicates the border between the pharynx ( to the left ) and intestine . [Field: results, subscore: 2.00]
SECTION: results. Wild-type ( N2 ) and mnm-4 mutant adults expressing ajm-1 : : gfp reporter were photographed to reveal not only the twisting within the pharynx of the mutant , but also the opposite twist in its anterior intestine . [Field: results, subscore: 2.00]
SECTION: results. One hypothesis is that the twisting force is already present in early larval stages but that its effect is hampered by attachment of the pharynx to surrounding it issue ( see previous section ) . [Field: results, subscore: 2.00]
SECTION: results. It is uncertain whether this accessory pseudocoelom is continuous with the pseudocoelom proper , due to close approach to the pharynx by GLR processes beneath the nerve ring [ 1 , 18 ] . [Field: results, subscore: 2.00]
SECTION: results. Only in larvae and adults can a distinct " accessory pseudocoelomic space " be seen that separates the thin ectodermal basal laminae from the thick basal lamina of the pharynx ( Fig . 7CD ) . [Field: results, subscore: 2.00]
SECTION: results. This suggests that appreciable friction between bodywall and pharyngeal it issues may hold the pharynx in place prior to the onset of pumping in the L1 , and may thus restrict onset of the twisting phenotype . [Field: results, subscore: 2.00]
SECTION: results. Compared to more posterior regions of the body cavity , this tightness remains a feature of the head throughout development , but some minimal separation does develop to accommodate vigorous motions of the pharynx during feeding . [Field: results, subscore: 2.00]
SECTION: results. Direct contact between the basal laminae of the pharynx and body-wall is very tight in the embryo Multiple studies of C . elegans embryos , including our own , have shown practically no available space between the mesodermal layers and ectodermal layers prior to hatching ( Fig . 7 ) . [Field: results, subscore: 2.00]
SECTION: results. More tenuously , the cuticle lining of the pharynx is also continuous with the bodywall cuticle in the region of the arcade cells ( buccal cavity ) . [Field: results, subscore: 2.00]
SECTION: results. BMC Developmental Biology 2007 , 7 : 61 http : / / www . biomedcentral . com / 1471-213X / 7 / 61 Page 7 of 13 ( page number not for citation purposes ) that the foremost intestinal cells twist in a direction opposite to the pharynx , hence relieving the tension induced by the pharyngeal twist ( Fig . 6 ) . [Field: results, subscore: 2.00]
SECTION: results. Because it is tilted with respect to the body axis , the tendon is better seen close to the pharynx in this image , but goes out of the plane of section as it passes between the muscle cells . [Field: results, subscore: 2.00]
SECTION: results. In ( B ) , note that strong degrees of twist can be detected in both the procorpus ( leftmost portion ) and at the isthmus between the two bulbs in the mutant pharynx , but only subtle twisting within each bulb . [Field: results, subscore: 2.00]
SECTION: results. Note that in all the specimens , the actin filaments are evenly spaced and generally aligned near-perpendicularly to the longitudinal axis of the pharynx if it were untwisted . [Field: results, subscore: 2.00]
SECTION: results. Indeed , when the twist is visualized using ajm1 : gfp in the mnm-4 background , the pharynx seems to have rotated at the posterior end around the terminal bulb passively dragging the intestine with it , such The actin cytoskeleton is twisted and lengthened in twisted pharynges Figure 5 The actin cytoskeleton is twisted and lengthened in twisted pharynges . [Field: results, subscore: 2.00]
SECTION: results. Number of beads in the intestine : 27 . 1 7 . 4 for N2 , and 30 . 8 7 . 7 ( n = 35 ) , indicating a similar transfer rate of beads from the pharynx to the intestine in the two genotypes . [Field: results, subscore: 2.00]
SECTION: results. Number of beads counted in the pharynx : 64 . 6 9 . 3 for N2 , and 78 . 1 11 . 4 ( n = 19 ) , indicating a similar steady-state number of beads in the two genotypes ; ( B ) Overlays of a DIC image and an epifluorescence image of thefluorescent beads in the intestine swallowed during a brief 2 min period . [Field: results, subscore: 2.00]
SECTION: results. ( A ) Overlays of a DIC image and an epifluorescence image of the fluorescent beads present in the pharynx following a feeding period of 30 minutes . [Field: results, subscore: 2.00]
SECTION: results. This is interesting since misregulation of filament length in growing pharyngeal muscles may be a cause of twisting : long actin filaments may be accommodated by twisting the pharynx . [Field: results, subscore: 2.00]
SECTION: results. Thus the whole pharynx , including its basal lamina to which the tendons are attached , is twisted in mutants . [Field: results, subscore: 2.00]
SECTION: results. As shown in Fig . 4C , the pharyngeal tendons are attached to the basal lamina that surrounds the pharynx and project between the bodywall muscle cells in each muscle quadrant , then connect on their opposite ends to the basement membranes running between bodywall muscles and thin extensions of the hypodermal cells , near the cuticle . [Field: results, subscore: 2.00]
SECTION: results. ( A ) An example of a twisted pharynx and the three measurements that can be obtained by DIC microscopy to estimate the actual degree of twist within the isthmus using the formula shown on the right side ( D is diameter ; L is isthmus length ; and is the angle between the torsion lines and the pharyngeal axis ) . [Field: results, subscore: 2.00]
SECTION: results. Panels G-J show the pharyngeal twist in mnm-4 animals carrying mutations that cause abnormal pharynx morphologies ( pha-2 , sma-1 ) or reduced pumping rate ( eat-3 ) . [Field: results, subscore: 2.00]
SECTION: results. However , eat-3 ; mnm-4 double mutants also showed a 10 % reduction in pharyngeal length ( " L " in Fig . 1A ) compared with mnm-4 mutants , and this may have contributed to the decrease in twist if growth of the pharynx is important . [Field: results, subscore: 2.00]
SECTION: results. * Only those dig- 1 and mig-4 worms with a pharyngeal were used to calculate the average twist of these mutants . http : / / www . biomedcentral . com / 1471-213X / 7 / 61 Page 3 of 13 ( page number not for citation purposes ) severely twisted pharynx ( etIs2 ; mnm-4 ; 3 . 7 0 . 3 pumps per second ) . [Field: results, subscore: 2.00]
SECTION: results. The pharynx of dig-1 and unc-61 mutants occasionally straightens ( 2 of 15 and 3 of 16 dissected pharynges , respectively ) when separated from the worm . [Field: results, subscore: 2.00]
SECTION: results. The result is also consistent with the twisting force being intrinsic to the pharynx itself . [Field: results, subscore: 2.00]
SECTION: results. This result does not exclude the possibility that the twist originates from outside the pharynx , but it shows that the twist has become irreversible by the time of dissection . [Field: results, subscore: 2.00]
SECTION: results. Conversely , the pharynx of unc-61 , mig-4 , dig-1 and mnm-4 mutant worms mostly keep their twisted shape after being dissected ( Fig . 2AF ) . [Field: results, subscore: 2.00]
SECTION: results. They were then scored for the presence and extent of twist . We found that the pharynx of wild-type worms does not twist when dissected away from the body . [Field: results, subscore: 2.00]
SECTION: results. Pharyngeal twist is retained in isolated pharynges , and thickened pharyngeal parts resist twisting We were interested to test whether the force causing twisting is an intrinsic property of the organ or originates from outside the pharynx . [Field: results, subscore: 2.00]
SECTION: results. This result suggests that these three genes may either interact directly or are involved in multiple steps in a single developmental process during larval growth of the pharynx . [Field: results, subscore: 2.00]
SECTION: results. Worms heterozygous for the mnm-4 mutation also exhibited a twisted pharynx , although the twist was less pronounced than in age-matched homozygous mutants ( Table 1 ) . [Field: results, subscore: 2.00]
SECTION: results. Properties of the twisted pharynx phenotype We began by studying the mnm-4 mutant and found that its pharyngeal twist is first visible post-embryonically and increases during development throughout the larval stages , rather than only in sudden jumps during molting , to reach its maximum during the fourth larval stage ( Fig . 1 ; Table 1 ) . [Field: results, subscore: 2.00]
SECTION: title. The twisted pharynx phenotype in C . elegans . [Field: title, subscore: 2.00]
SECTION: abstract. The twisted pharynx is a useful and easy-to-score phenotype for genes required in extracellular adhesion or organ attachment , and perhaps for genes required for cytoskeleton regulation . [Field: abstract, subscore: 1.00]
SECTION: abstract. In a mini screen of adhesion molecule mutants , we also identified one more twisting pharynx mutant , sax-7 . [Field: abstract, subscore: 1.00]
SECTION: abstract. : We find that the twisting phenotype worsens throughout larval development , that in most mutants the pharynx retains its twist when dissected away from the worm body , and that double mutants between mnm-4 and mutants with thickened pharyngeal domains ( pha-2 and sma-1 ) have less twisting in these regions . [Field: abstract, subscore: 1.00]
SECTION: abstract. Several C . elegans mutants have been reported to exhibit a twisted pharynx phenotype but no careful studies have been made to directly address this phenomenon . [Field: abstract, subscore: 1.00]
SECTION: abstract. : BACKGROUND : The pharynx of C . elegans is an epithelial tube whose development has been compared to that of the embryonic heart and the kidney and hence serves as an interesting model for organ development . [Field: abstract, subscore: 1.00]
SECTION: conclusion. The twisted pharynx phenotype is found in several C . elegans mutants that have defects in adhesion molecules or molecules that regulate attachment of the actin cytoskeleton to the cell cortex . 2 . [Field: conclusion, subscore: 1.00]
SECTION: introduction. The aim of the present study was to better understand the twisted pharynx phenotype and its relationship to extracellular matrix components . [Field: introduction, subscore: 1.00]
SECTION: introduction. Finally , the mnm-4 mutant was isolated in a screen for abnormal morphology of the M2 pharyngeal neurons , and also has a twisted pharynx [ 12 ] . [Field: introduction, subscore: 1.00]
SECTION: introduction. Interestingly , a mutation in the other C . elegans septin , unc-59 , does not cause a twisted pharynx ; this is thus one of thefew phenotypes for which unc-59 and unc-61 differ [ 11 ] . [Field: introduction, subscore: 1.00]
SECTION: introduction. The uncoordinated mutant unc-61 also has a twisted pharynx . [Field: introduction, subscore: 1.00]
SECTION: introduction. Several C . elegans mutants have been reported to exhibit a twisted pharynx phenotype . [Field: introduction, subscore: 1.00]
SECTION: introduction. There are 80 cell nuclei from five cell types present in the pharynx : muscle cells , nerve cells , marginal cells , epithelial cells and gland cells [ 1 , 2 ] . [Field: introduction, subscore: 1.00]
SECTION: introduction. Background The pharynx is a simple muscular epithelial tube responsible for the ingestion and maceration of food in C . elegans . [Field: introduction, subscore: 1.00]
SECTION: introduction. The twisted pharynx is a useful and easy-to-score phenotype for genes required in extracellular adhesion or organ attachment , and perhaps forgenes required for cytoskeleton regulation . [Field: introduction, subscore: 1.00]
SECTION: introduction. In a mini screen of adhesionmolecule mutants , we also identified one more twisting pharynx mutant , sax-7 . [Field: introduction, subscore: 1.00]
SECTION: introduction. : We find that the twisting phenotype worsens throughout larval development , that in most mutants the pharynx retains its twist when dissected away from the worm body , and that double mutants between mnm-4 and mutants with thickened pharyngeal domains ( pha-2 and sma- 1 ) have less twisting in these regions . [Field: introduction, subscore: 1.00]
SECTION: introduction. Several C . elegans mutants have been reported to exhibit a twisted pharynx phenotype but no careful studies have been made to directly address this phenomenon . [Field: introduction, subscore: 1.00]
SECTION: introduction. Background : The pharynx of C . elegans is an epithelial tube whose development has been compared to that of the embryonic heart and the kidney and hence serves as an interesting model for organ development . [Field: introduction, subscore: 1.00]
SECTION: introduction. BMC Developmental Biology Research article The twisted pharynx phenotype in C . elegans Claes Axng1 , Manish Rauthan1 , David H Hall Address : 1Dept . of Chemical and Biological Engineering , Lundberg Laboratory , Chalmers University , Box 462 , S-405 30 , Gteborg , Sweden , Neuroscience , Albert Einstein College of Medicine , Bronx , New York , NY 10461 , USA and Box 462 , S-405 30 , Gteborg , Sweden Email : Claes Axng - claes . axang @ molbiotech . chalmers . se ; Manish Rauthan - manish . rauthan @ cmb . gu . se ; David H Hall - hall @ aecom . yu . edu ; Marc Pilon * - marc . pilon @ cmb . gu . se * Corresponding author Abstract [Field: introduction, subscore: 1.00]
SECTION: materials. The worms were thereafter mounted on agarose pad ( 2 % agarose in M9 buffer ) in a drop of 100 mM levamisol , covered with a cover slip and scored for the presence beads in the intestine and the pharynx using epifluorescence microscopy at 400 magnification . [Field: materials, subscore: 1.00]
SECTION: references. Raizen DM , Avery L : Electrical activity and behavior in the pharynx of Caenorhabditis elegans . [Field: references, subscore: 1.00]
SECTION: references. Mrck C , Axng C , Pilon M : A genetic analysis of axon guidance in the C . elegans pharynx . [Field: references, subscore: 1.00]
SECTION: references. Portereiko MF , Mango S : Early morphogenesis of the Caenorhabditis elegans pharynx . [Field: references, subscore: 1.00]
SECTION: references. Avery L , Shtonda BS : Food transport in the C . elegans pharynx . [Field: references, subscore: 1.00]
SECTION: references. Albertson DG , Thomson JN : The pharynx of Caenorhabditis elegans . [Field: references, subscore: 1.00]
Supplemental links/files: reference in endnote reference in xml Wormbase reference
Score: 191.00
Title: Coordination of ges-1 expression between the Caenorhabditis pharynx and intestine .
Authors: Marshall SDG ; McGhee JD
Journal: Dev Biol
Year: 2001-11-15
Doc ID: WBPaper00004994
Bibliographic Information
Abstract
Matching Sentences
SECTION: discussion. Two differences in the behavior of the two ges-1 genes were also described : ( 1 ) the endogenous Cb-ges-1 gene shows a low level of activity in the pharynx ( and possibly in the rectum ) , whereas the Ce-ges-1 counterpart does not ; this difference is carried over into transgenic nematodes , where the exogenous Cb-ges-1 gene produces a significantly greater level of pharynx expression , whether the transformation host is C . elegans or C . briggsae ; and ( 2 ) deletion of either one of the two C . elegans WGATAR sites or deletion of an immediately adjacent region activates Ce-ges-1 expression only in the anterior gut ( Egan et al . , 1995 ; Schroeder and McGhee , 1998 ) ; in contrast , no evidence could be found for anterior gut expression produced by any of the variously modified Cb-ges-1 constructs . [Field: discussion, subscore: 4.00]
SECTION: discussion. The main features of ges-1 control also appear to be conserved between the two species : ( 1 ) expression of both genes in the intestine lineage depends on a tandem pair of WGATAR sites located in the 5 -flanking region ; ( 2 ) deletion of these WGATAR sites abolishes expression in the gut lineage but concomitantly activates expression in cells of the pharynx and rectum ; and ( 3 ) the cryptic pharynx enhancer , revealed when the WGATAR sites are deleted , lies in a conserved sequence at the 3 -end of ges-1 in both species and contains a conserved PHA-4 binding site . [Field: discussion, subscore: 4.00]
SECTION: results. In a total of 19 transformed strains ( 11 and 8 strains with the Ce-ges-1 and Cb-ges-1 enhancers , respectively ) , all but one strain expressed the GFP reporter strongly in the pharynx and only the pharynx ( easily detectable , even at the level of the fluorescent dissecting microscope ) . [Field: results, subscore: 4.00]
SECTION: results. The C . elegans ( and presumably C . briggsae ) ortholog of HNF3 is the product of the pha-4 gene , which is expressed in all cells of the pharynx and in many cells of the rectum ; moreover , pha-4 is crucial for development of the pharynx as an organ ( Mango et al . , 1994 ; Azzaria et al . , 1996 ; Horner et al . , 1998 ; Kalb et al . , 1998 ) . [Field: results, subscore: 4.00]
SECTION: results. The ges-1 Pharynx Enhancer Sequence Interacts with PHA-4 To identify binding factors potentially responsible for the ges-1 pharynx enhancer activity , the two 300-bp regions were aligned as shown in Fig . 6B and analyzed by the TRANSFAC program ( Heinemeyer et al . , 1998 ) . [Field: results, subscore: 4.00]
SECTION: results. Identification of the Pharynx Activator in the Conserved 3 Region of the C . briggsae ges-1 Gene The results shown in Fig . 6A identify the pharynx activator within the Cb-ges-1 gene . [Field: results, subscore: 4.00]
SECTION: results. First , deletion of either the upstream or the downstream WGATAR site from Cb-ges-1 produces the complete switch in expression pattern from gut to pharynx ( Fig . 4C ) ; in contrast , deletion of either the upstream or the downstream WGATAR sites from Ce-ges-1 abolishes expression in the posterior gut , maintains expression in the anterior gut , but does not activate expression in the pharynx rectum ( Egan et al . , 1995 ) . [Field: results, subscore: 4.00]
SECTION: results. Although the majority of transgenic esterase activity is indeed in the gut , both embryos show low but significant levels of staining in the pharynx , considerably more than is observed when the Ce-ges-1 gene is transformed back into C . elegans ( where pharynx rectum staining is low though not zero ; data not shown and see also Egan et al . , 1995 ) . [Field: results, subscore: 4.00]
SECTION: introduction. While the ELT-2 GATA factor is currently the best ( but not the only ) candidate for binding the WGATAR sites and activating ges-1 expression in the gut ( Hawkins and McGhee , 1995 ; Fukushige et al . , 1998 ) , the identity of the pharynx / rectum repressor , the pharynx / rectum enhancer , and the pharynx / rectum enhancer binding factor are presently unknown . [Field: introduction, subscore: 3.00]
SECTION: introduction. Coordination of ges-1 Expression Between the Caenorhabditis Pharynx and Intestine Sean D . G . Marshall1 and James D . McGhee2 Department of Biochemistry and Molecular Biology , Genes Development Research Group , University of Calgary , Calgary , Alberta , Canada T2N 4N1 We have previously shown that the Caenorhabditis elegans gut-specific esterase gene ( Ce-ges-1 ) has the unusual ability to be expressed in different modules of the embryonic digestive tract ( anterior pharynx , posterior pharynx , and rectum ) depending on sequence elements within the Ce-ges-1 promoter . [Field: introduction, subscore: 3.00]
SECTION: abstract. We have previously shown that the Caenorhabditis elegans gut-specific esterase gene ( Ce-ges-1 ) has the unusual ability to be expressed in different modules of the embryonic digestive tract ( anterior pharynx , posterior pharynx , and rectum ) depending on sequence elements within the Ce-ges-1 promoter . [Field: abstract, subscore: 2.00]
SECTION: discussion. Perhaps ges-1 is expressed in the pharynx only under certain environmental conditions yet to be defined . [Field: discussion, subscore: 2.00]
SECTION: discussion. A further possibility is that ges-1 is expressed in the pharynx of dauer larvae . [Field: discussion, subscore: 2.00]
SECTION: discussion. However , the pharynx enhancer is expressed weakly if at all in pharyngeal gland cells ( Fig . 7C ) . [Field: discussion, subscore: 2.00]
SECTION: discussion. Under normal conditions , ges-1 expression in the pharynx would be detrimental and is kept repressed , but the worm nonetheless retains the latent ability to activate ges-1 when needed . [Field: discussion, subscore: 2.00]
SECTION: discussion. ( 3 ) The third model , which presently seems the most reasonable , proposes that ges-1 serves some function in the pharynx but only under particular environmental conditions or at particular developmental stages . [Field: discussion, subscore: 2.00]
SECTION: discussion. 361 ges-1 Expression in the Caenorhabditis Pharynx Copyright 2001 by Academic Press . [Field: discussion, subscore: 2.00]
SECTION: discussion. ( C ) A projection of a focal series of digitally deconvolved images taken through the pharynx of an L4 larva from strain JM97 ; expressing nuclei are visualized by GFP fluorescence . [Field: discussion, subscore: 2.00]
SECTION: discussion. ( A , B ) A pretzel stage embryo and an L2 larva stained for -galactosidase activity ; note strong pharynx staining but absence of staining in the rectum . [Field: discussion, subscore: 2.00]
SECTION: discussion. ( 2 ) Perhaps the selection pressure maintaing the ges-1 expression switch lies not in the pharynx but rather involves a role for pha-4 in the gut . [Field: discussion, subscore: 2.00]
SECTION: discussion. However , if silencing of ges-1 expression in the pharynx were the only driving force for maintenance of the ges-1 expression switch , it would seem much more likely that the pharyngeal enhancer would simply be disabled by mutation , rather than by the elaboration and maintenance of a complex repression mechanism . [Field: discussion, subscore: 2.00]
SECTION: discussion. At some point in evolution , esterase expression in the pharynx ( rectum ) might have become detrimental to the worm and thus ges-1 repression would provide an advantage . [Field: discussion, subscore: 2.00]
SECTION: discussion. PHA-4 protein is present in all cells of the embryonic pharynx ( Horner et al . , 1998 ; Kalb et al . , 1998 ) and yet the WGATAR-deleted ges-1 genes are only expressed in a subset of these cells ( Egan et al . , 1995 ; Fukushige et al . , 1996 ) , as are the reporter constructs driven by the ges-1 3 -enhancer ( see Fig . 7C ) . [Field: discussion, subscore: 2.00]
SECTION: discussion. Indeed , one possible explanation for the observation that mutation and deletion of the WGATAR sites produce somewhat different effects ( Fig . 5C above , and Fig . 3 of Egan et al . , 1995 ) is that the factors controlling the gut activation and the pharynx rectum repression have different and separable sequence requirements . [Field: discussion, subscore: 2.00]
SECTION: discussion. Of course , the pharynx repressor need not be a GATA factor but need only bind in the immediate vicinity of the WGATAR sites . [Field: discussion, subscore: 2.00]
SECTION: discussion. All rights of reproduction in any form reserved . these factors are present either in the ( ABa-derived ) anterior pharynx or in the ( ABp-derived ) rectum , nor whether they are present when the altered ges-1 genes are first expressed . [Field: discussion, subscore: 2.00]
SECTION: discussion. Although the C . elegans genomic sequence reveals 11 proteins with significant similarity to GATA factors , none appear good candidates for the ges-1 pharynx rectum repressor . [Field: discussion, subscore: 2.00]
SECTION: discussion. Do our present results provide any insight into the nature of the pharynx repressor ? [Field: discussion, subscore: 2.00]
SECTION: discussion. However , in the case of the ges-1 gene in the pharynx , activation by PHA-4 must be repressed by a factor that binds in or around the 5 - WGATAR region . [Field: discussion, subscore: 2.00]
SECTION: discussion. We have suggested that PHA-4 participates in all acts of transcription in the developing pharynx ( Kalb et al . , 1998 ) . [Field: discussion, subscore: 2.00]
SECTION: discussion. PHA-4 is the C . elegans homolog of Drosophila forkhead and the vertebrate HNF-3 , , transcription factors , is present in all committed pharyngeal precursor cells , and functions as a " pharynx identity factor " ( Horner et al . , 1998 ; Kalb et al . , 1998 ) . [Field: discussion, subscore: 2.00]
SECTION: discussion. In cells of the pharynx ( and perhaps rectum as well ) , we propose that PHA-4 protein is bound to the conserved enhancer at the ges-1 3 -end . [Field: discussion, subscore: 2.00]
SECTION: introduction. As illustrated in Fig . 1 , the four distinct modules of the C . elegans digestive tract are derived from four distinct cell lineages : anterior pharynx , posterior pharynx , and rectum are derived from the ABa , MS , and ABp blastomeres , respectively , and the gut proper is clonally derived from the single E blastomere of the eight-cell embryo ( Sulston et al . , 1983 ) . [Field: introduction, subscore: 2.00]
SECTION: results. All rights of reproduction in any form reserved . marginal cells ; staining in other classes of pharynx cells ( e . . , epithelial , gland or nerve cells , as well as cells of the pharyngealintestinal valve ) was much weaker and more variable . [Field: results, subscore: 2.00]
SECTION: results. 359 ges-1 Expression in the Caenorhabditis Pharynx Copyright 2001 by Academic Press . [Field: results, subscore: 2.00]
SECTION: results. Deletion of this site in construct pJM102 . 25 abolishes the pharynx expression from Cb-ges-1 . [Field: results, subscore: 2.00]
SECTION: results. ( D ) A more detailed deletion analysis demonstrating the bipartite nature of the C . briggsae 3 -pharynx enhancer . [Field: results, subscore: 2.00]
SECTION: results. ( B ) Sequence alignment of the proposed pharynx enhancer from the C . briggsae and C . elegans ges-1 genes ( region # 2 from Fig . 2 ) . [Field: results, subscore: 2.00]
SECTION: results. The region suggested to contain the pharynx enhancer is indicated by the pair of vertical dashed lines . [Field: results, subscore: 2.00]
SECTION: results. Deletion constructs depicted below the gene line produce a strong gut staining pattern but pharynx staining is abolished ; precise coordinates of the various deletions are provided in Marshall ( 1998 ) . [Field: results, subscore: 2.00]
SECTION: results. Clearly , cells in both the anterior and posterior pharynx express the reporter gene . [Field: results, subscore: 2.00]
SECTION: results. However , pharynx activity is also abolished by deletions that extend from 3791 to 3805 bp , suggesting that at least one other nearby site must also be involved ; this latter region is not easily matched with a conserved sequence in the alignment shown in Fig . 6B . [Field: results, subscore: 2.00]
SECTION: results. As predicted , a deletion that removes the conserved PHA-4 binding site ( together with neighboring conserved sequences as shown in Fig . 6B ) does indeed abolish pharynx activity . [Field: results, subscore: 2.00]
SECTION: results. Does this conserved PHA-4 binding site influence pharynx enhancer activity within the context of the ges-1 gene ? [Field: results, subscore: 2.00]
SECTION: results. Electrophoretic mobility shift assays ( Fig . 6C ) demonstrate that this conserved site from the C . elegans ges-1 3 pharynx enhancer is indeed a binding site for pure PHA-4 protein ( isoform C , produced in baculovirus ) ; similar results were obtained by using the corresponding site from the C . briggsae gene ( data not shown ) . [Field: results, subscore: 2.00]
SECTION: results. However , when two copies of this 68-bp Ce-ges-1 sequence were inserted at the 3 -end of Cb-ges-1 ( within deletion pJM102 . 21 ; see Fig . 6A ) , pharynx expression was not restored ( data not shown ) , suggesting that any enhancer activity that this region might possess is minor . [Field: results, subscore: 2.00]
SECTION: results. A potential pharynx rectum activator element had been tentatively identified immediately upstream of the Ce-ges-1 ATG ( Aamodt et al . , 1991 ; Egan et al . , 1995 ) . [Field: results, subscore: 2.00]
SECTION: results. We conclude that the conserved 3 - region of the ges-1 genes contains the major pharynx activator whose activity is revealed by deletion of the 5 WGATAR sites . [Field: results, subscore: 2.00]
SECTION: results. The Conserved 3 Region Is the Major Pharynx Activator in the ges-1 Genes We deleted the 3 -conserved regions from Ce-ges-1 and Cb-ges-1 constructs that were also deleted for the 5 tandem pair of WGATAR sites . [Field: results, subscore: 2.00]
SECTION: results. Overall , the results demonstrate that the pharynx activator must lie within several hundred base pairs downstream of the poly ( A ) addition site , within the conserved # 2 region identified in Fig . 2 . [Field: results, subscore: 2.00]
SECTION: results. In contrast , deletions depicted below the gene line abolish this pharynx activity ( but retain gut activity ) . [Field: results, subscore: 2.00]
SECTION: results. Deletions depicted above the line representing the Cb-ges-1 gene have no effect on the normal low level of pharynx staining observed with the intact Cb-ges-1 transgene construct , pJM102 . [Field: results, subscore: 2.00]
SECTION: results. However , the upstream Cbges-1 WGATAR site shows a different behavior : whereas deletion of the upstream site abolishes gut expression and activates pharynx rectum expression ( Fig . 4C , above ) , mutation of the same site has little effect ( Fig . 5C ) . [Field: results, subscore: 2.00]
SECTION: results. 357 ges-1 Expression in the Caenorhabditis Pharynx Copyright 2001 by Academic Press . [Field: results, subscore: 2.00]
SECTION: results. Mutation of both WGATAR sites or mutation of only the downstream WGATAR site has the same effect as site deletion , i . . , gut activity is abolished and pharynx ( rectum ) staining is activated in both C . elegans and C . briggsae hosts . [Field: results, subscore: 2.00]
SECTION: results. All transformations are done in a C . elegans host . Moving the tandem pair of WGATAR sites 300 bp downstream from their normal location reestablishes wild-type Cb-ges-1 staining ( strong gut , weak pharynx rectum ) . [Field: results, subscore: 2.00]
SECTION: results. All rights of reproduction in any form reserved . paired WGATAR sites were moved ( in the reverse orientation ) upstream 1 kb from their normal location , wild-type Cb-ges-1 expression was only partially reconstituted , i . . , expression in the gut was weak and pharynx ( rectum ) staining persisted . [Field: results, subscore: 2.00]
SECTION: results. ( C ) Deletion of either or both of the WGATAR sites from the Cb-ges-1 5 -flanking region is sufficient to abolish Cb-ges-1 staining in the gut and activate expression in the pharynx . [Field: results, subscore: 2.00]
SECTION: results. Transformed deletion constructs depicted above the gene line produce a wild-type Cb-ges-1 staining pattern , both in a C . elegans and a C . briggsae host . Deletion constructs depicted below the gene line do not produce gut staining but rather produce staining in the pharynx and rectum . [Field: results, subscore: 2.00]
SECTION: results. 355 ges-1 Expression in the Caenorhabditis Pharynx Copyright 2001 by Academic Press . [Field: results, subscore: 2.00]
SECTION: results. Esterase staining is intense in all cells of the gut ; weaker staining is apparent in the pharynx . [Field: results, subscore: 2.00]
SECTION: results. The arrow in the 4-h C . briggsae sample points to weak endogenous ges-1 staining in the C . briggsae embryonic pharynx . [Field: results, subscore: 2.00]
SECTION: results. As shown in Fig . 5B , reinsertion of the paired WGATAR sites in the forward orientation at a site 300 bp downstream from their normal location is sufficient to reconstitute normal expression patterns , i . . , strong gut , weak pharynx . [Field: results, subscore: 2.00]
SECTION: results. Can the WGATAR sites function as portable " gutactivators / pharynx repressors " within the context of the Cb-ges-1 gene ? [Field: results, subscore: 2.00]
SECTION: results. One possible explanation for the unusual expression switches of the ges-1 genes is that their expression is influenced by elements controlling nearby genes ; for example , if the neighboring genes were expressed in pharynx or rectum . [Field: results, subscore: 2.00]
SECTION: results. Although the overall features of this intradigestive tract switch in expression pattern seem to be maintained be- 353 ges-1 Expression in the Caenorhabditis Pharynx Copyright 2001 by Academic Press . [Field: results, subscore: 2.00]
SECTION: results. In general , rectum staining is weaker and more variable than observed with corresponding Ce-ges-1 deletions and from this point on , we focus primarily on pharynx staining . [Field: results, subscore: 2.00]
SECTION: results. Deletion of these two sites ( a 35-bp deletion within the otherwise intact construct pJM102 ) does indeed cause the Cb-ges-1 expression pattern to switch from the gut into the pharynx ( Fig . 4C ) . [Field: results, subscore: 2.00]
SECTION: results. The Gut-Activation / Pharynx Rectum Repression of the C . briggsae ges-1 Gene Center on a Tandem Pair of WGATAR Sites The dot-matrix comparison of Fig . 2 shows no obvious sequence conservation between the 5 WGATAR region of Ce-ges-1 and the 100-bp region controlling the gut-topharynx rectum switch in Cb-ges-1 expression , nor could significant alignments be produced by the programs GAP or BESTFIT [ Wisconsin Package Version 9 . 1 , Genetics Computer Group ( GCG ) , Madison , WI ] . [Field: results, subscore: 2.00]
SECTION: results. In contrast , for the two deletions represented below the gene line ( in both C . briggsae and C . elegans hosts ) , expression in the gut is extinguished essentially completely ( best demonstrated in the C . elegans ges-1 null mutant ) but concomitantly activated in cells of the pharynx . [Field: results, subscore: 2.00]
SECTION: results. The two deletions shown above the central line produce essentially the same expression pattern as does the undeleted parent construct pJM102 , i . . , intense gut staining and weak staining in the pharynx ( and perhaps rectum ) . [Field: results, subscore: 2.00]
SECTION: results. We must assume that this also holds true for C . briggsae embryos and that the endogenous activity in the C . briggsae pharynx is indeed caused by Cb-ges-1 and not by some other esterase . [Field: results, subscore: 2.00]
SECTION: results. However , a weak activity can be detected in the C . briggsae pharynx after 4 h staining that is not detected in C . elegans under the same condition ( see also Fukushige et al . , 1996 ) . [Field: results, subscore: 2.00]
SECTION: results. The Endogenous Cb-ges-1 Gene Shows a Low Level of Pharyngeal Activity Figure 3A compares the esterase staining patterns observed with wild-type C . elegans ( strain N2 ) and wild-type C . briggsae ( strain AF16 ) at three different staining periods : 3 min ( appropriate for assaying multicopy transgenes ) , 1 h ( appropriate for detecting the single copy endogenous ges-1 gene ) , and 4 h ( used to detect the low level of pharynx rectum staining produced by chromosomal deletions of the endogenous Ce-ges-1 promoter ; Fukushige et al . , 1996 ) . [Field: results, subscore: 2.00]
SECTION: results. This region will be shown below to contain the cryptic pharynx enhancer . [Field: results, subscore: 2.00]
SECTION: title. Coordination of ges-1 expression between the Caenorhabditis pharynx and intestine . [Field: title, subscore: 2.00]
SECTION: abstract. We propose that , in the pharynx ( and rectum ) , PHA-4 is normally bound to the ges-1 3 ' -enhancer sequence but that the activation function of PHA-4 is kept repressed by a ( presently unknown ) factor binding in the vicinity of the 5 ' WGATAR sites . [Field: abstract, subscore: 1.00]
SECTION: abstract. This region contains a conserved binding site for PHA-4 ( the C . elegans ortholog of forkhead / HNF3 alpha , beta , gamma factors ) , which is expressed in all cells of the developing pharynx and a subset of cells of the developing rectum . [Field: abstract, subscore: 1.00]
SECTION: abstract. We use sequence alignments and subsequent deletions to identify a region at the 3 ' -end of both Ce-ges-1 and Ce-ges-1 that acts as the ges-1 cryptic pharynx enhancer whose activity is revealed by removal of the 5 ' WGATAR sites . [Field: abstract, subscore: 1.00]
SECTION: abstract. In the present paper , we analyze the expression of the ges-1 homolog ( Cb-ges-1 ) from the related nematode Caenorhabditis briggsae and show that Cb-ges-1 also has the ability to switch expression between gut and pharynx + rectum . [Field: abstract, subscore: 1.00]
SECTION: introduction. 351 ges-1 Expression in the Caenorhabditis Pharynx Copyright 2001 by Academic Press . [Field: introduction, subscore: 1.00]
SECTION: introduction. All nuclei in the pharynx and a subset of nuclei in the rectum ( red ) are visualized by using an antibody against PHA-4 ( Kalb et al . , 1998 ) . [Field: introduction, subscore: 1.00]
SECTION: introduction. Two of the principal factors that we propose participate in this intermodule regulatory network are shown in Fig . 1 : PHA-4 in the pharynx and the rectum ( red ) and ELT-2 in the intestine proper ( green ) . [Field: introduction, subscore: 1.00]
SECTION: introduction. The pharynx ( rectum ) repressor proposed normally to silence the PHA-4 transactivation function is as yet unknown . [Field: introduction, subscore: 1.00]
SECTION: introduction. The pha-4 gene is the worm homolog of Drosophila forkhead and vertebrate HNF-3 , , genes and has been shown to act as an " organ-identity-factor " during pharynx ( and rectum ) development ( Horner et al . , 1998 ; Kalb et al . , 1998 ) . [Field: introduction, subscore: 1.00]
SECTION: introduction. We use sequence comparisons and subsequent gene deletions to identify the major pharynx enhancer , which lies at the 3 -end of both C . elegans and C . briggsae ges-1 genes and whose activity is revealed by deletion of the WGATAR sequences . [Field: introduction, subscore: 1.00]
SECTION: introduction. We show that the central feature of ges-1 regulation is retained in C . briggsae , namely a WGATAR site-dependent switch in expression between gut and pharynx / rectum . [Field: introduction, subscore: 1.00]
SECTION: introduction. Our present model is that ges-1 harbors a pharynx / rectum enhancer sequence somewhere within the gene but that the activity of this enhancer is normally kept repressed by a factor that binds to , or in the vicinity , of these 5 WGATAR sites . [Field: introduction, subscore: 1.00]
SECTION: introduction. ges-1 shows unusual and interesting behavior that suggests that gene expression may indeed be coordinated throughout the entire digestive tract : deletion of a tandem pair of WGATAR sites ( where W A or T and R A or G ) in the 5 -flanking region abolishes ges-1 expression in the gut but simultaneously activates ges-1 expression in both anterior and posterior pharynx as well as the rectum , i . . , the three other modules of the digestive tract ( Aamodt et al . , 1991 ; Egan et al . , 1995 ; Fukushige et 1 Present address : HortResearch , Private Bag 92169 , Auckland , New Zealand . [Field: introduction, subscore: 1.00]
SECTION: introduction. 2001 Academic Press Key Words : Caenorhabditis elegans ; Caenorhabditis briggsae ; ges-1 ; gene expression ; intestine ; pharynx ; elt-2 ; pha-4 . [Field: introduction, subscore: 1.00]
SECTION: introduction. We propose that , in the pharynx ( and rectum ) , PHA-4 is normally bound to the ges-1 3 -enhancer sequence but that the activation function of PHA-4 is kept repressed by a ( presently unknown ) factor binding in the vicinity of the 5 WGATAR sites . [Field: introduction, subscore: 1.00]
SECTION: introduction. This region contains a conserved binding site for PHA-4 ( the C . elegans ortholog of forkhead / HNF3 , , factors ) , which is expressed in all cells of the developing pharynx and a subset of cells of the developing rectum . [Field: introduction, subscore: 1.00]
SECTION: introduction. We use sequence alignments and subsequent deletions to identify a region at the 3 -end of both Ce-ges-1 and Ce-ges-1 that acts as the ges-1 cryptic pharynx enhancer whose activity is revealed by removal of the 5 WGATAR sites . [Field: introduction, subscore: 1.00]
SECTION: introduction. In the present paper , we analyze the expression of the ges-1 homolog ( Cb-ges-1 ) from the related nematode Caenorhabditis briggsae and show that Cb-ges-1 also has the ability to switch expression between gut and pharynx rectum . [Field: introduction, subscore: 1.00]
SECTION: references. Received for publication May 8 , 2001 Revised August 22 , 2001 Accepted August 22 , 2001 Published online October 11 , 2001 363 ges-1 Expression in the Caenorhabditis Pharynx Copyright 2001 by Academic Press . [Field: references, subscore: 1.00]
SECTION: references. pha-4 is Ce-fkh-1 , a fork head / HNF-3 , , homolog that functions in organogenesis of the C . elegans pharynx . [Field: references, subscore: 1.00]
SECTION: references. A fork head / HNF-3 homolog expressed in the pharynx and intestine of the Caenorhabditis elegans embryo . [Field: references, subscore: 1.00]
Supplemental links/files: reference in endnote reference in xml Wormbase reference
Score: 183.00
Title: TGF- signaling can act from multiple it issues to regulate C . elegans body size .
Authors: Dineen A ; Gaudet J
Journal: BMC Dev Biol
Year: 2014-12-06
Doc ID: WBPaper00046090
Bibliographic Information
Abstract
Matching Sentences
SECTION: discussion. The pharynges of Dpy mutants are also significantly smaller than wild type suggesting that growth of this organ is limited by the length of the entire animal . Thus , pharynx length appears to be determined partly by positive TGF- / Sma / Mab signaling but is also dependent on the overall body length of the animal . It was interesting that signaling in the pharynx never resulted in full rescue of pharynx length even when body length was completely rescued . [Field: discussion, subscore: 6.00]
SECTION: discussion. Discussion The TGF- / Sma / Mab pathway regulates pharynx length In a previous study on the morphology of the pharynx , we identified a number of Sma mutants with decreased pharynx lengths compared to wild type , including members of the TGF- / Sma / Mab signaling pathway sma-2 and sma-3 [ 20 ] . [Field: discussion, subscore: 6.00]
SECTION: discussion. It is unclear if Twp animals have reduced pharynx lengths but if so it would be a noteworthy parallel between pharynx growth and overall body growth . [Field: discussion, subscore: 4.00]
SECTION: discussion. Previous study of mutations in extracellular matrix ( ECM ) components ( including the cuticle collagen dpy-7 ) and membrane proteins revealed a role in pharynx morphology and a twisted pharynx ( Twp ) phenotype [ 39 , 40 ] . [Field: discussion, subscore: 4.00]
SECTION: discussion. This result implies that TGF- / Sma / Mab signaling from outside the pharynx must also play a role in determining pharynx length . [Field: discussion, subscore: 4.00]
SECTION: results. We conclude from this experiment that pharynx length is regulated by TGF- / Sma / Mab pathway signaling from both within and outside the pharynx . [Field: results, subscore: 4.00]
SECTION: results. The simple interpretation of these results is that TGF- / Sma / Mab signaling acts within the pharynx to control pharynx length , as it also does in the hypodermis . [Field: results, subscore: 4.00]
SECTION: results. We also found that pharynx length of sma-3 ( e491 ) mutants was significantly restored by the sma-3p : : sma-3 transgene ( 119 6 % compared to 100 4 % for non-transgenic siblings , p < 0 . 001 ) , consistent with the expectation that pharynx length is regulated by sma-3 . [Field: results, subscore: 4.00]
SECTION: results. Given the expression pattern of sma-3 in pharyngeal cells and the decreased pharynx length of sma-3 mutants , we next asked whether pharynx length could be rescued by pharyngeal expression of sma-3 . [Field: results, subscore: 4.00]
SECTION: results. S . The TGF- / Sma / Mab effector SMA-3 can act in the pharynx to regulate body length sma-3 is reported to be expressed in the pharynx ( as are other components of the pathway ) [ 16 , 17 ] , yet the individual cells in which it is expressed have not been described . [Field: results, subscore: 4.00]
SECTION: results. However , another possibility is that pharynx length is reduced as a consequence of reduced body length , specifically , that growth of the pharynx might be constrained by the smaller hypodermis . [Field: results, subscore: 4.00]
SECTION: results. The reduced pharynx length in these TGF- / Sma / Mab pathway mutants suggests that pathway activity is required for pharyngeal growth , consistent with expression of pathway components in the pharynx . [Field: results, subscore: 4.00]
SECTION: results. These two pieces of evidence suggest first , that TGF- / Sma / Mab pathway signaling may regulate pharynx length and second , that signaling within the pharynx may contribute significantly to body length regulation . [Field: results, subscore: 4.00]
SECTION: results. Furthermore , pharynx lengths of Sma mutants at the L3 stage were found to be slightly but significantly smaller than the pharynx length of wild type N2 . [Field: results, subscore: 4.00]
SECTION: conclusion. TGF- / Sma / Mab signaling in the pharynx is capable of contributing to pharynx length and body length regulation but this signaling is not sufficient or necessary to facilitate wild type pharynx length or body length . [Field: conclusion, subscore: 3.00]
SECTION: conclusion. This suggests that TGF- / Sma / Mab signaling in the pharynx and hypodermis may be coordinating growth of many tissues within the animal . Figure 6 Linear relationship between pharynx length and body length in mutants of the TGF- / Sma / Mab pathway and strains carrying various sma-3 rescuing extrachromosomal arrays in the sma-3 ( e491 ) mutant background . [Field: conclusion, subscore: 2.00]
SECTION: discussion. Interestingly , the K07C11 . 4 promoter , which drives expression in the pharynx and late stage somatic gonad did not rescue by itself but in combination with the rol-6p : : sma-3 transgene almost completely rescued the sma-3 ( wk30 ) body length phenotype to N2 levels . [Field: discussion, subscore: 2.00]
SECTION: discussion. Interestingly , at least one other TGF- / Sma / Mab component has been demonstrated to act in the pharynx . [Field: discussion, subscore: 2.00]
SECTION: discussion. Finally , components of the TGF- / Sma / Mab pathway , including SMA-3 , are expressed in the pharynx and hypodermis , consistent with the proposed function of this pathway in these tissues . [Field: discussion, subscore: 2.00]
SECTION: discussion. Second , we find that simultaneous expression of sma-3 in both the hypodermis and pharynx provides stronger rescue of body length than when sma-3 is expressed in either it issue alone suggesting that both can contribute to normal growth in an additive manner . [Field: discussion, subscore: 2.00]
SECTION: discussion. Activity of the TGF- / Sma / Mab pathway in multiple it issues can control body length The results described here support a model in which downstream effectors of TGF- / Sma / Mab signaling can act in both the pharynx and hypodermis to influence overall body length in C . elegans . [Field: discussion, subscore: 2.00]
SECTION: discussion. It would be interesting to see if this pathway mediates cell and it issue growth via different downstream targets in the pharynx vs the hypodermis . [Field: discussion, subscore: 2.00]
SECTION: discussion. In contrast , the pharynx does not undergo any cell number or ploidy changes during post embryonic development [ 4 ] . [Field: discussion, subscore: 2.00]
SECTION: discussion. Furthermore , pharynx and body length can be restricted by morphology defects in components of the surrounding extracellular cuticle . [Field: discussion, subscore: 2.00]
SECTION: discussion. It is important to note the similarity in growth control of the pharynx and the hypodermis , where both organs utilize positive TGF- / Sma / Mab signaling to regulate organ and overall body length . [Field: discussion, subscore: 2.00]
SECTION: discussion. Here we report that mutants of TGF- / Sma / Mab pathway components have significantly smaller pharynx lengths compared to Dpy mutants of similar body length . [Field: discussion, subscore: 2.00]
SECTION: results. The increase in rescue that is observed with the elevated dose of dbl-1 ligand validates our conclusion it is TGF- / Sma / Mab pathway signaling in the pharynx that is rescuing body length in these animals and not an independent function of SMA-3 . [Field: results, subscore: 2.00]
SECTION: results. Surprisingly , the animals that carried the extra chromosomal pharyngeal sma-3 rescue array and dbl-1 ( + ) array insertion had fully rescued body length compared to wild type ( p = 0 . 132 ) but not pharynx length ( p < 0 . 001 ) ( Figure 5 , Additional file 5 ) . [Field: results, subscore: 2.00]
SECTION: results. As expected , all worms with the dbl-1 ( + ) array that did not get the extra chromosomal pharyngeal sma-3 rescue array ( as determined by the lack of extra chromosomal array reporter expression ) had small mean pharynx length and body length ( Figure 5 , Additional file 5 ) . [Field: results, subscore: 2.00]
SECTION: results. Figure 5 Mean pharynx and body length measurements standard deviation of sodium azide anesthetized wild type ( Wt ) and sma-3 mutant animals at 120 1 hrs AEL from various sma-3 minigene rescue experiments . [Field: results, subscore: 2.00]
SECTION: results. These results confirm that pharyngeal signaling is partially rescuing pharynx length and body length in sma-3 ( e491 ) mutants . [Field: results, subscore: 2.00]
SECTION: results. Furthermore , a statistical significant difference is observed between sma-3 ( e491 ) mutants at 120 1 hours and pharyngeal rescued transgenic animals at 96 1 hours for both pharynx length ( 110 5 % compared to 100 3 % , p < 0 . 001 ) and body length ( 122 11 % compared to 100 8 % , p < 0 . 001 ) ( Additional file 5 ) . [Field: results, subscore: 2.00]
SECTION: results. Table 2 Percentage of Wt and sma-3 ( e491 ) animals ( + or - pharyngeal sma-3 minigene constructs ) that had laid an egg by the time point indicated ( hrs after initial egg laid ) Strain / Time Point 48 hrs 66 hrs 72 hrs 90 hrs 96 hrs 114 hrs 120 hrs n Wt 0 % 8 . 7 % 34 . 8 % 100 % 100 % 100 % 100 % 23 sma-3 ( e491 ) 0 % 0 % 0 % 20 % 68 % 100 % 100 % 25 sma-3 ( e491 ) 0 % 0 % 0 % 42 . 9 % 66 . 7 % 100 % 100 % 21 myo-2p : : sma-3 + marg-1p : : sma-3 ( - ) sma-3 ( e491 ) 0 % 0 % 0 % 88 . 2 % 91 . 1 % 100 % 100 % 34 myo-2p : : sma-3 + marg-1p : : sma-3 ( + ) We observed partial rescue of both pharynx length and body length with the pharyngeal promoters driving the sma-3 minigene ( sma-3 ( e491 ) background ) as well as the elt- 3 : : gfp : : sma-3 transgene ( sma-3 ( wk30 ) background ) ( Figure 5 , Additional file 5 ) which is consistent with our previous results . [Field: results, subscore: 2.00]
SECTION: results. This K07C11 . 4p : : sma-3 and rol-6p : : sma-3 combination had significantly greater pharynx and body length rescue than that observed with the combination of myo-2p : : sma-3 , marg- 1p : : sma-3 , and rol-6p : : sma-3 ( p < 0 . 001 ) . [Field: results, subscore: 2.00]
SECTION: results. However , the combination of K07C11 . 4p : : sma-3 and rol-6p : : sma-3 showed significant rescue of both pharynx length ( 120 5 % compared to 100 3 % for nontransgenic siblings , p < 0 . 001 ) and body length ( 150 15 % of N2 length compared to 100 7 % for non-transgenic siblings , p < 0 . 001 ) ( Figure 3C , D , Additional file 5 ) . [Field: results, subscore: 2.00]
SECTION: results. Again , we found that the combination of the pharyngeal ( myo-2p : : sma-3 , marg- 1p : : sma-3 ) and hypodermal ( rol-6p : : sma-3 ) transgenes provided significantly greater rescue of sma-3 ( wk30 ) mutants by 96 1 hours after egg laying compared to the pharyngeal transgenes alone ( 114 5 % of non-transgenic sibling pharynx length compared to 107 6 % for pharyngeal transgenes alone , p < 0 . 001 and 137 12 % of non-transgenic sibling body length compared to 113 8 % for pharyngeal transgenes alone , p < 0 . 001 ) ( Figure 3C , D , Additional file 5 ) . [Field: results, subscore: 2.00]
SECTION: results. However , hypodermal expression using the elt-3p : : gfp : : sma-3 transgene did significantly rescue pharynx length ( 107 5 % compared to 100 3 % for non-transgenic siblings , p < 0 . 001 ) and body length ( 122 12 % compared to 100 5 % for non-transgenic siblings , p < 0 . 001 ) . [Field: results, subscore: 2.00]
SECTION: results. Similar to the results observed for the sma- 3 ( e491 ) background , we did not see significant rescue of the pharynx or body length phenotypes of sma-3 ( wk30 ) with the rol-6p : : sma-3 transgene ( Figure 3C , D , Additional file 5 ) . [Field: results, subscore: 2.00]
SECTION: results. Furthermore , the combination of pharyngeal ( myo-2p : : sma- 3 and marg-1p : : sma-3 ) and hypodermal ( rol-6p : : sma-3 ) transgenes was able to rescue body length of sma-3 ( e491 ) mutants to the same extent as the native sma-3p : : sma-3 transgene ( 154 18 % and 155 16 % respectively of their non-transgenic siblings , p = 0 . 173 ) , though the difference in pharynx lengths was still significant ( p < 0 . 001 ) . [Field: results, subscore: 2.00]
SECTION: results. This pharyngeal and hypodermal combination of transgenes resulted in significantly greater rescue of both pharynx and body length in sma-3 ( e491 ) mutants than seen with the pharyngeal transgenes alone ( p < 0 . 001 ) ( Figure 3A , B , Additional file 5 ) . [Field: results, subscore: 2.00]
SECTION: results. We found that this combination of pharyngeal and hypodermal transgenes leads to an average pharynx length of 118 4 % compared to 100 2 % for nontransgenic siblings , and an average body length of 154 18 % compared to 100 7 % for nontransgenic siblings ( Figure 3A , B , Additional file 5 ) . [Field: results, subscore: 2.00]
SECTION: results. Quantitating the intensity in a 16bit black and white file we detect a 25-30 fold difference between GFP reporter expression intensity in the pharynx and the faint adjacent cells , which was barely above background intensity . [Field: results, subscore: 2.00]
SECTION: results. As outlined above , in overexposed images we observed variable weak GFP expression in a few cells adjacent to the pharynx ( Figure 2D , Additional file 4 ) . [Field: results, subscore: 2.00]
SECTION: results. Strong expression was observed in the pharynx however , the exposure was also significantly increased to rule out low levels of ectopic expression . [Field: results, subscore: 2.00]
SECTION: results. These results also indicate significant but incomplete rescue of body length by pharynx specific sma-3 minigene constructs . [Field: results, subscore: 2.00]
SECTION: results. As with rescue of pharynx length , both the myo-2p : : sma-3 and marg-1p : : sma-3 transgenes showed some rescue of body length individually , while the combination of transgenes ( 130 16 % compared with 100 7 % for non-transgenic siblings , 20 ng / L injection mix ) exhibited a greater rescue than either transgene alone ( p < 0 . 01 ) . [Field: results, subscore: 2.00]
SECTION: results. The combination of myo-2p : : sma-3 and marg-1p : : sma-3 transgenes ( 20 ng / L injection mix ) produced animals with an average pharynx length of 113 5 % compared to 100 3 % for non-transgenic sibling controls , a significantly greater degree of rescue compared to either alone ( p < 0 . 001 ) . [Field: results, subscore: 2.00]
SECTION: results. The relative small effect of rescue on pharynx length by each of these pharyngeal transgenes was nonetheless statistically significant compared to their non-transgenic siblings ( p < 0 . 001 , MannWhitney Rank Sum Test on raw data ) . [Field: results, subscore: 2.00]
SECTION: results. myo-2p : : sma-3 rescued animals had pharynx lengths that were 110 5 % compared to 100 3 % for non-transgenic siblings and marg-1p : : sma-3 rescued animals were 107 3 % of N2 length compared to 100 2 % for non-transgenic siblings . [Field: results, subscore: 2.00]
SECTION: results. We next tested whether the myo-2p : : sma-3 and marg-1p : : sma-3 transgenes could rescue sma-3 ( e491 ) pharynx length to N2 levels , either alone or in combination ( Figures 3A , 4 , Additional file 5 ) . [Field: results, subscore: 2.00]
SECTION: results. Figure 3 Mean pharynx and body length measurements standard deviation of sma- 3 ( e491 ) ( A , B ) and sma-3 ( wk30 ) ( C , D ) animals from various sma-3 minigene rescue experiments at 96 hrs AEL . [Field: results, subscore: 2.00]
SECTION: results. Extremely weak expression was also observed in one to four cells just outside of the pharynx in half of the animals ( n = 140 , Additional file 4 ) but only when the exposure was increased dramatically . [Field: results, subscore: 2.00]
SECTION: results. As expected , in all stages we observed strong expression of these translational fusions in the pharynx with notable lack of expression in the pharyngeal glands ( Figure 2D ) . [Field: results, subscore: 2.00]
SECTION: results. We attempted to rescue pharynx length by expressing a sma-3 ( + ) minigene ' under the control of different pharyngeal promoters ( see Additional file 3 for a list of all promoters used in rescue experiments and their it issue specificity ) : myo-2 , which is expressed solely in pharyngeal muscles [ 29 ] and K07C11 . 4 , which is expressed in pharyngeal muscle , marginal cells and epithelia , the intestine , and in late stage somatic gonad [ 22 ] . [Field: results, subscore: 2.00]
SECTION: results. Expression is absent from pharyngeal gland cells ( arrowhead ) and weak expression is occasionally observed outside of the pharynx ( arrow ) . [Field: results, subscore: 2.00]
SECTION: results. However , the sma-3 ( e491 ) allele may have dominant neomorphic properties as both mean pharynx and body length are significantly smaller in heterozygous ( p < 0 . 001 ) and homozygous ( p < 0 . 001 ) animals compared to the sma-3 ( wk30 ) nulls ( Additional files 1 , 2 ) . [Field: results, subscore: 2.00]
SECTION: results. Within the pharynx , we observe expression in most or all pharyngeal muscles and marginal cells ( Figure 2A ) . [Field: results, subscore: 2.00]
SECTION: results. As previously described [ 16 ] , expression of the reporter was observed in hypodermis , intestine and pharynx . [Field: results, subscore: 2.00]
SECTION: results. All other differences in pharynx length between strains but not directly indicated on the graph are significant ( p < 0 . 001 ) with the exception of N . [Field: results, subscore: 2.00]
SECTION: results. Figure 1 Mean pharynx and body length measurements standard deviation of wild type ( Wt ) N2 and body size mutants at 96 1 hrs AEL . [Field: results, subscore: 2.00]
SECTION: results. These measurements imply that pharynx length may be partially reduced in response to smaller body length but is also positively influenced by TGF- / Sma / Mab signaling . [Field: results, subscore: 2.00]
SECTION: results. To test this latter possibility , we measured pharynx length in two hypodermal collagen mutants with reduced body length , dpy-5 ( e61 ) and dpy-10 ( e128 ) [ 26 , 27 ] . [Field: results, subscore: 2.00]
SECTION: results. The TGF- / Sma / Mab pathway regulates pharynx length Previous reports indicated that TGF- / Sma / Mab signaling in the hypodermis controls body length [ 16 , 17 , 25 ] . [Field: results, subscore: 2.00]
SECTION: abstract. Specifically , we find that pharyngeal expression of the SMAD protein SMA-3 partially rescues both pharynx length and body length of sma-3 mutants . [Field: abstract, subscore: 1.00]
SECTION: abstract. : Here we show that TGF- / Sma / Mab signaling is required for the normal growth of the pharynx . [Field: abstract, subscore: 1.00]
SECTION: abstract. However , components of this signaling pathway are expressed in other organs , such as the intestine and pharynx , raising the question of what the function of the pathway is in these organs . [Field: abstract, subscore: 1.00]
SECTION: conclusion. Furthermore , organs of Dpy mutants such as the pharynx and gonad often appear compressed compared to those of TGF- / Sma / Mab pathway mutants which appear more proportional to the overall body length of the animal [ 11 ] . [Field: conclusion, subscore: 1.00]
SECTION: conclusion. Consistent with signaling between it issues , animals in which manipulation of TGF- / Sma / Mab signaling results in uncoupled hypodermal and pharynx lengths were not observed ( i . . we have not seen small animals with big pharynges or vice versa ) . [Field: conclusion, subscore: 1.00]
SECTION: conclusion. We note a strong linear correlation between pharynx and body length , except in Dpy mutants , in which signaling is presumably normal but body length is reduced due to defects in hypodermal collagen ( Figure 6 ) . [Field: conclusion, subscore: 1.00]
SECTION: conclusion. Taken together , using our results presented here and those of previous studies on it issue specific regulation of body length [ 16 , 17 , 42 ] , we make the following conclusions about pharynx length and body length regulation by TGF- / Sma / Mab signaling . [Field: conclusion, subscore: 1.00]
SECTION: conclusion. Given the developmental delay observed in animals homozygous for sma-3 mutant alleles and the artificial nature of C . elegans extrachromosomal arrays it is difficult to quantitate exactly how much contribution TGF- / Sma / Mab signaling in the pharynx makes to regulation of body length . [Field: conclusion, subscore: 1.00]
SECTION: introduction. However , components of this signaling pathway are expressed in other organs , such as the intestine and pharynx , raising the question of what the function of the pathway is in these organs . [Field: introduction, subscore: 1.00]
SECTION: materials. Pharynx length was measured as the distance from the posterior of the buccal cavity to the pharyngeal-intestinal valve . [Field: materials, subscore: 1.00]
SECTION: materials. Pharynx and body lengths were measured in ImageJ using segmented lines [ 24 ] . [Field: materials, subscore: 1.00]
SECTION: references. All differences in pharynx length not directly indicated on the graph are significant ( p < 0 . 001 ) , except where indicated by N . [Field: references, subscore: 1.00]
SECTION: references. Graphs of Pharynx and Body Lengths of Various Strains in the Absence of Anesthetic . [Field: references, subscore: 1.00]
SECTION: references. Measurements of pharynx and body length of sma-3 ( e491 ) and sma-3 ( wk30 ) animals carrying different sma-3 minigene constructs ( mean standard deviation ) . [Field: references, subscore: 1.00]
SECTION: references. Pharynx and Body Lengths of Strains Imaged Using Various Anesthetics at Different Time Points After Egg Laying During Development . [Field: references, subscore: 1.00]
SECTION: references. Very weak expression is occasionally observed outside of the pharynx in some animals ( arrowheads ) ( 6373k ) http : / / www . biomedcentral . com / content / supplementary / s12861-014-0043-8-s4 . tiff Additional file 5 : Table S3 . [Field: references, subscore: 1.00]
SECTION: references. All other differences in pharynx and body lengths between strains not directly indicated on the graphs are significant ( p < 0 . 001 ) ( 493k ) http : / / www . biomedcentral . com / content / supplementary / s12861-014-0043-8-s2 . tiff Additional file 3 : Table S2 . it issue Specificity of Promoters Used to Drive Rescue Constructs . [Field: references, subscore: 1.00]
SECTION: references. Mean pharynx and body length measurements standard deviation of Wild type ( Wt ) N2 , sma-3 mutants and sma-3 heterozygotes . [Field: references, subscore: 1.00]
SECTION: references. Graphs of Pharynx and Body Lengths of Wt and Various sma-3 Mutant Strains . [Field: references, subscore: 1.00]
SECTION: references. Measurements of pharynx and body length of wild type ( Wt ) N2 and various mutant strains , expressed as mean ( standard deviation ) . [Field: references, subscore: 1.00]
SECTION: references. Pharynx and Body Lengths of Various Strains . [Field: references, subscore: 1.00]
SECTION: references. Jafari G , Burghoorn J , Kawano T , Mathew M , Morck C , Axang C , Ailion M , Thomas JH , Culotti JG , Swoboda P : Genetics of extracellular matrix remodeling during organ growth using the Caenorhabditis elegans pharynx model . [Field: references, subscore: 1.00]
SECTION: references. Axang C , Rauthan M , Hall DH , Pilon M : The twisted pharynx phenotype in C . elegans . [Field: references, subscore: 1.00]
SECTION: references. Raharjo WH , Ghai V , Dineen A , Bastiani M , Gaudet J : Cell architecture : surrounding muscle cells shape gland cell morphology in the Caenorhabditis elegans pharynx . [Field: references, subscore: 1.00]
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