WormMine

WS296

Intermine data mining platform for C. elegans and related nematodes

Gene :

WormBase Gene ID  ? WBGene00003401 Gene Name  mpk-1
Sequence Name  ? F43C1.2 Brief Description  mpk-1 encodes a mitogen-activated protein (MAP) kinase an ERK ortholog functioning in vulval cell fate specification, cell migration/guidance, defense against bacterial infection, and other processes; in proximal germline, sperm-dependent physiological MPK-1 activation results in phosphorylation based inactivation of NOS-3, FEM-CUL-2-mediated degradation of TRA-1 and the promotion of membrane organization during oogenesis; mpk-1 affect LET-60(Ras)-mediated induction of vulval cell fates, larval viability, morphology of the male spicules; both LIN-1 and LIN-31 act genetically downstream of mpk-1 with respect to vulval induction; MPK-1 is necessary for pachytene cell organization; mpk-1 mutants display germ cells arrested in pachytene; MPK-1 is necessary for the progression from distal to proximal pachytene; MPK-1 promotes pachytene progression, with the rise in dpMPK-1 triggering a transition from a distal pachytene to a proximal pachytene subtype; MPK-1 functions in the germline for meiotic prophase progression and gametogenesis; MPK-1 ERK signaling is necessary for the male germ cell fate; MPK-1 ERK has the nonessential function of promoting the proliferative germ cell fate; mpk-1 acts in combination with mek-2 to permit germ cell exit from the pachytene stage of first meiotic prophase; MPK-1 activation is temporally/spatially dynamic compared to relatively constant levels of total MPK-1.
Organism  Caenorhabditis elegans Automated Description  Enables protein serine/threonine kinase activity. Involved in several processes, including intracellular signaling cassette; oocyte maturation; and positive regulation of macromolecule metabolic process. Located in cytoplasm and nucleus. Expressed in F cell; U cell; germ line; gonad; and oocyte. Used to study cancer. Human ortholog(s) of this gene implicated in several diseases, including Alzheimer's disease; Noonan syndrome 13; carcinoma (multiple); and childhood pilocytic astrocytoma. Is an ortholog of human MAPK1 (mitogen-activated protein kinase 1).
Biotype  SO:0001217 Genetic Position  III :-3.65654 ±0.031007
Length (nt)  ? 11346
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1 Organism

Name Taxon Id
Caenorhabditis elegans 6239

1 Synonyms

Value
WBGene00003401

Genomics

3 Transcripts

WormMine ID Sequence Name Length (nt) Chromosome Location
Transcript:F43C1.2a.2 F43C1.2a.2 1764   III: 4216763-4219643
Transcript:F43C1.2a.1 F43C1.2a.1 1981   III: 4216764-4228108
Transcript:F43C1.2b.1 F43C1.2b.1 1335   III: 4217396-4228094
 

Other

2 CDSs

WormMine ID Sequence Name Length (nt) Chromosome Location
CDS:F43C1.2b F43C1.2b 1335   III: 4217396-4217615
CDS:F43C1.2a F43C1.2a 1131   III: 4217396-4217615

115 RNAi Result

WormBase ID
WBRNAi00101975
WBRNAi00101976
WBRNAi00110543
WBRNAi00113183
WBRNAi00067095
WBRNAi00067367
WBRNAi00067538
WBRNAi00067597
WBRNAi00067959
WBRNAi00068041
WBRNAi00068090
WBRNAi00068205
WBRNAi00047161
WBRNAi00072495
WBRNAi00072496
WBRNAi00027587
WBRNAi00101981
WBRNAi00085349
WBRNAi00072494
WBRNAi00068479
WBRNAi00068478
WBRNAi00068751
WBRNAi00073555
WBRNAi00073557
WBRNAi00073560
WBRNAi00087339
WBRNAi00087338
WBRNAi00108296
WBRNAi00068811
WBRNAi00068923

161 Allele

Public Name
WBVar01692349
WBVar01692350
WBVar02067469
WBVar02067468
WBVar02067470
WBVar01262784
WBVar01262785
WBVar01262786
WBVar01262781
WBVar01262782
WBVar01262787
WBVar01262788
WBVar01607000
WBVar01656519
WBVar01656518
h8506
h3084
gk903091
gk369434
gk365332
gk663140
WBVar02010399
q1190
gk172376
ga111
gk383262
ga117
gk795351
gk172378
gk680920

1 Chromosome

WormBase ID Organism Length (nt)
III Caenorhabditis elegans 13783801  

1 Chromosome Location


Feature . Primary Identifier
Start End Strand
WBGene00003401 4216763 4228108 -1

4 Data Sets

Name URL
WormBaseAcedbConverter  
GO Annotation data set  
C. elegans genomic annotations (GFF3 Gene)  
Panther orthologue and paralogue predictions  

1 Downstream Intergenic Region

WormBase ID Name Sequence Name Length (nt) Chromosome Location Organism
intergenic_region_chrIII_4216750..4216762   13 III: 4216750-4216762 Caenorhabditis elegans

165 Expression Clusters

Regulated By Treatment Description Algorithm Primary Identifier
  Genes with expression altered >= 3-fold in dpy-10(e128) mutants. Data across the wild type series was analyzed using the Significance analysis of Microarrays (SAM) algorithm (to calculate the False Discovery Rate (FDR)). WBPaper00035873:dpy-10_regulated
  Transcripts that showed significantly increased expression in L1 neural cells comparing to in adult neural cells. DESeq2 (v1.18.1) fold change > 2, P-adj<0.05, using BenjaminiHochberg correction. WBPaper00060811:L1_vs_adult_upregulated_neural
  Transcripts expressed in neuronal cells, by analyzingfluorescence-activated cell sorted (FACS) neurons. DESeq. False discovry rate (FDR) < 0.1. WBPaper00048988:neuron_expressed
  Genes significantly enriched (> 2x, FDR < 5%) in a particular cell-type versus a reference sample of all cells at the same stage. A Mann-Whitney U test with an empirical background model and FDR correction for multiple testing was used to detect expressed transcripts (Benjamini and Hochberg 1995). Genes and TARs with an FDR <= 0.05 were reported as expressed above background. Authors detected differentially expressed transcripts using a method based on linear models. Genes and TARs were called differentially expressed if the FDR was <= 0.05 and the fold change (FC) >= 2.0. To more strictly correct for potential false-positives resulting from multiple sample comparisons, authors divided individual FDR estimates by the number of samplesor sample comparisons, respectively. This resulted in an adjusted FDR of 1.3 * 0.0001 for expression above background and of 7.4 * 0.0001 for differential expression. Authors called genes selectively enriched in a given tissue if they met the following requirements: (1) enriched expression in a given tissue (FDR <= 0.05 and FC >= 2.0), (2) fold change versus reference among the upper 40% of the positive FC range observed for this gene across all tissues, and (3) fold-change entropy among the lower 40% of the distribution observed for all genes. WBPaper00037950:A-class-motor-neurons_larva_enriched
  Genes that showed expression levels higher than the corresponding reference sample (embryonic 24hr reference). A Mann-Whitney U test with an empirical background model and FDR correction for multiple testing was used to detect expressed transcripts (Benjamini and Hochberg 1995). Genes and TARs with an FDR <= 0.05 were reported as expressed above background. Authors detected differentially expressed transcripts using a method based on linear models. Genes and TARs were called differentially expressed if the FDR was <= 0.05 and the fold change (FC) >= 2.0. To more strictly correct for potential false-positives resulting from multiple sample comparisons, authors divided individual FDR estimates by the number of samplesor sample comparisons, respectively. This resulted in an adjusted FDR of 1.3 * 0.0001 for expression above background and of 7.4 * 0.0001 for differential expression. Authors called genes selectively enriched in a given tissue if they met the following requirements: (1) enriched expression in a given tissue (FDR <= 0.05 and FC >= 2.0), (2) fold change versus reference among the upper 40% of the positive FC range observed for this gene across all tissues, and (3) fold-change entropy among the lower 40% of the distribution observed for all genes. WBPaper00037950:all-neurons_L1-larva_expressed
adult vs dauer larva Transcripts that showed differential expression in adult vs dauer lava in N2 animals at 20C. N.A. WBPaper00050488:adult_vs_dauer_regulated_N2_20C
  mRNAs that showed decreased expression in 1 cell mebryo comparing to in oocyte, according to RNAseq analysis. Gaussian error propagation. As cutoff for the up-regulated genes authors used log2 fold change > 1 and P < 0.05 and as cutoff for the down-regulated genes authors used log2 fold change < -1 and P < 0.05. WBPaper00045420:fertilization_downregulated_transcript
  Genes that showed expression levels higher than the corresponding reference sample (embryonic 24hr reference). A Mann-Whitney U test with an empirical background model and FDR correction for multiple testing was used to detect expressed transcripts (Benjamini and Hochberg 1995). Genes and TARs with an FDR <= 0.05 were reported as expressed above background. Authors detected differentially expressed transcripts using a method based on linear models. Genes and TARs were called differentially expressed if the FDR was <= 0.05 and the fold change (FC) >= 2.0. To more strictly correct for potential false-positives resulting from multiple sample comparisons, authors divided individual FDR estimates by the number of samplesor sample comparisons, respectively. This resulted in an adjusted FDR of 1.3 * 0.0001 for expression above background and of 7.4 * 0.0001 for differential expression. Authors called genes selectively enriched in a given tissue if they met the following requirements: (1) enriched expression in a given tissue (FDR <= 0.05 and FC >= 2.0), (2) fold change versus reference among the upper 40% of the positive FC range observed for this gene across all tissues, and (3) fold-change entropy among the lower 40% of the distribution observed for all genes. WBPaper00037950:AVE-neuron_L1-larva_expressed
Osmotic stress Transcripts that showed significantly altered expression with 500 mM salt (NaCl) vs 100 mM salt when food was present DESeq(version 1.10.1), FDR < 0.05. WBPaper00050726:OsmoticStress_regulated_Food
Osmotic stress Transcripts that showed significantly altered expression with 500 mM salt (NaCl) vs 100 mM salt when no food was present DESeq(version 1.10.1), FDR < 0.05. WBPaper00050726:OsmoticStress_regulated_NoFood
  Genes that showed expression levels higher than the corresponding reference sample (embryonic 24hr reference). A Mann-Whitney U test with an empirical background model and FDR correction for multiple testing was used to detect expressed transcripts (Benjamini and Hochberg 1995). Genes and TARs with an FDR <= 0.05 were reported as expressed above background. Authors detected differentially expressed transcripts using a method based on linear models. Genes and TARs were called differentially expressed if the FDR was <= 0.05 and the fold change (FC) >= 2.0. To more strictly correct for potential false-positives resulting from multiple sample comparisons, authors divided individual FDR estimates by the number of samplesor sample comparisons, respectively. This resulted in an adjusted FDR of 1.3 * 0.0001 for expression above background and of 7.4 * 0.0001 for differential expression. Authors called genes selectively enriched in a given tissue if they met the following requirements: (1) enriched expression in a given tissue (FDR <= 0.05 and FC >= 2.0), (2) fold change versus reference among the upper 40% of the positive FC range observed for this gene across all tissues, and (3) fold-change entropy among the lower 40% of the distribution observed for all genes. WBPaper00037950:bodywall-muscle_L1-larva_expressed
  Proteins interacting with NHR-49-GFP according to co-IP and LC-MS. N.A. WBPaper00064071:NHR-49_interacting
  Transcripts expressed in the epithelial tissues surrounding the pharynx that includes the arcade and intestinal valve (AIV) cells, according to PAT-Seq analysis using Pbath-15-GFP-3XFLAG mRNA tagging. Cufflinks FPKM value >=1. WBPaper00050990:arcade_intestinal-valve_expressed
  Transcripts expressed in body muscle, according to PAT-Seq analysis using Pmyo-3-GFP-3XFLAG mRNA tagging. Cufflinks FPKM value >=1. WBPaper00050990:body-muscle_expressed
  Transcripts expressed in GABAergic neuron, according to PAT-Seq analysis using Punc-47-GFP-3XFLAG mRNA tagging. Cufflinks FPKM value >=1. WBPaper00050990:GABAergic-neuron_expressed
  Transcripts expressed in hypodermis, according to PAT-Seq analysis using Pdpy-7-GFP-3XFLAG mRNA tagging. Cufflinks FPKM value >=1. WBPaper00050990:hypodermis_expressed
  Transcripts expressed in intestine, according to PAT-Seq analysis using Pges-1-GFP-3XFLAG mRNA tagging. Cufflinks FPKM value >=1. WBPaper00050990:intestine_expressed
  Transcripts expressed in NMDA neuron, according to PAT-Seq analysis using Pnmr-1-GFP-3XFLAG mRNA tagging. Cufflinks FPKM value >=1. WBPaper00050990:NMDA-neuron_expressed
  Transcripts expressed in pharynx, according to PAT-Seq analysis using Pmyo-2-GFP-3XFLAG mRNA tagging. Cufflinks FPKM value >=1. WBPaper00050990:pharynx_expressed
  Genes with expression level regulated by genotype (N2 vs CB4856) and age at L3 larva and Late reproduction stage (96 hours at 24 centigrade). For model 2, authors used 100 permutations to estimate the FDR threshold. Per permutation, genotypes and ages were independently randomly distributed, keeping the among-gene structure intact. Then for each spot (23,232) on the array, model 2 was tested. The obtained P-values were used to estimate a threshold for each of the explanatory factors. Authors also used a genome-wide threshold of -log10 P-value = 2, which resembles an FDR of 0.072 and 0.060 for marker and the interaction age-marker for the developing worms and FDR of 0.050 and 0.065 for marker and age-marker for the aging worms. For the physiological age effect, authors used a log10 P-value = 8 in developing worms (0.012 FDR) and -log10 P-value = 6 (0.032 FDR). WBPaper00040858:eQTL_age_regulated_developing
  Transcripts expressed in seam cells, according to PAT-Seq analysis using Pgrd-10-GFP-3XFLAG mRNA tagging. Cufflinks FPKM value >=1. WBPaper00050990:seam_expressed
  Genes with expression level regulated by genotype (N2 vs CB4856) at L3 larva and Late reproduction stage (96 hours at 24 centigrade). For model 2, authors used 100 permutations to estimate the FDR threshold. Per permutation, genotypes and ages were independently randomly distributed, keeping the among-gene structure intact. Then for each spot (23,232) on the array, model 2 was tested. The obtained P-values were used to estimate a threshold for each of the explanatory factors. Authors also used a genome-wide threshold of -log10 P-value = 2, which resembles an FDR of 0.072 and 0.060 for marker and the interaction age-marker for the developing worms and FDR of 0.050 and 0.065 for marker and age-marker for the aging worms. For the physiological age effect, authors used a log10 P-value = 8 in developing worms (0.012 FDR) and -log10 P-value = 6 (0.032 FDR). WBPaper00040858:eQTL_regulated_developing
  Transcripts detected in germline isolated from day-1 adult hermaphrodite animals. All three experiments have CPM >= 1. WBPaper00067147:germline_expressed
Bacteria infection: Bacillus thuringiensis mRNAs that showed significantly decreased expression after pathogenic bacteria Bacillus thuringiensis infections comparing to non pathogenic BT (BT247(1 to 10 mix) vs BT407 12h), according to RNAseq. Cuffdiff, ajusted p-value < 0.01. WBPaper00046497:B.thuringiensis_0.1mix_downregulated_12h
Bacteria infection: Bacillus thuringiensis mRNAs that showed significantly decreased expression after pathogenic bacteria Bacillus thuringiensis infections comparing to non pathogenic BT (BT247(1 to 2 mix) vs BT407 6h), according to RNAseq. Cuffdiff, ajusted p-value < 0.01. WBPaper00046497:B.thuringiensis_0.5mix_downregulated_6h
Bacteria infection: Bacillus thuringiensis mRNAs that showed significantly decreased expression after pathogenic bacteria Bacillus thuringiensis infections comparing to non pathogenic BT (BT247(1 to 2 mix) vs BT407 12h), according to RNAseq. Cuffdiff, ajusted p-value < 0.01. WBPaper00046497:B.thuringiensis_0.5mix_downregulated_12h
  Transcripts that showed significantly altered expression after 24 hour exposure to stavudine (d4T) starting at L1 lava stage. DESeq WBPaper00053302:stavudine_24h_regulated
  Transcripts that showed significantly increased expression in 10-days post L4 adult hermaphrodite N2 grown at 20C, comparing to in 1-day post L4 adult hermaphrodite N2 animals grown at 20C. CuffDiff, fold change > 2. WBPaper00065096:Day10_vs_Day1_upregulated
  Maternal class (M): genes that are called present in at least one of the three PC6 replicates. A modified Welch F statistic was used for ANOVA. For each gene, regressed error estimates were substituted for observed error estimates. The substitution is justified by the lack of consistency among the most and least variable genes at each time point. Regressed error estimates were abundance-dependent pooled error estimates that represented a median error estimate from a window of genes of similar abundance to the gene of interest. A randomization test was used to compute the probability Pg of the observed F statistic for gene g under the null hypothesis that developmental time had no effect on expression. P-values were not corrected for multiple testing. [cgc5767]:expression_class_M
  Strictly maternal class (SM): genes that are the subset of maternal genes that are not also classified as embryonic. A modified Welch F statistic was used for ANOVA. For each gene, regressed error estimates were substituted for observed error estimates. The substitution is justified by the lack of consistency among the most and least variable genes at each time point. Regressed error estimates were abundance-dependent pooled error estimates that represented a median error estimate from a window of genes of similar abundance to the gene of interest. A randomization test was used to compute the probability Pg of the observed F statistic for gene g under the null hypothesis that developmental time had no effect on expression. P-values were not corrected for multiple testing. [cgc5767]:expression_class_SM

15 Expression Patterns

Remark Reporter Gene Primary Identifier Pattern Subcellular Localization
    Expr1031571 Tiling arrays expression graphs  
Strain: BC14536 [mpk-1::gfp] transcriptional fusion. PCR products were amplified using primer A: 5' [CATTTTACGGGTTCTCTCTCATCT] 3' and primer B 5' [GTATCCACGTTGGGATCATGT] 3'. Expr6056 Adult Expression: Nervous System; head neurons; Larval Expression: pharyngeal-intestinal valve; Nervous System; head neurons;  
Picture: Figure 1A, 2A.   Expr7890 In wild-type hermaphrodites 24 hr post mid-L4, total MPK-1 is found throughout the germline, with slightly lower levels in the distal-most end and slightly higher levels in the proximal region. dpMPK-1(diphosphorylated activated) is undetectable in the mitotic and transition zone regions, peaks in the proximal part of pachytene, becomes low but detectable in the loop region, and peaks again in diakinesis oocytes in the proximal gonad. Total MPK-1 is primarily cytoplasmic except in late-stage oocytes where there is nuclear enrichment.
Picture: Figure 1C.   Expr7881 MPK-1A was present in both GL+ and GL- animals, but MPK-1B protein was found only in GL+ animals. Thus mpk-1b RNA and its MPK-1B protein are predominantly expressed in the germline.  
Picture: Figure 1B.   Expr7880 The mpk-1ab mRNA was abundant in both GL+ and GL- animals, but mpk-1b mRNA was very low or undetectable in GL- animals. Therefore, mpk-1b appears to be enriched in the germline.  
Picture: Fig. 4, A and C.   Expr7879 This antibody stains the two to three most proximal oocytes only in the presence of sperm. The MAPK-YT antibody only recognizes mpk-1 MAP kinase gene products because no staining is observed in gonads from mpk-1(ga117) homozygotes, a likely protein null.  
    Expr1151022 Developmental gene expression time-course. Raw data can be downloaded from ftp://caltech.wormbase.org/pub/wormbase/datasets-published/hashimshony2015  
Picture: N.A. Reporter gene fusion type not specified.   Expr8797 Expression of mpk-1::gfp was observed in the rectal epithelial cells, including F and U, in 100% of wild type L1 (16/16), L2 (19/19), L3 (15/15), L4 (21/21) and adult (16/16)worms.  
Picture: Figure S1.   Expr8056 Within the gonad, activated, di-phosphorylated MAP kinase (MPK-1) is first detectable at the mid-pachytene stage of the meiotic prophase I. The activated form of MAP kinase disappears as germ cells pass the pachytene stage and enter the diplotene/diakinesis phase of meiotic prophase I. Activated, M.A.P kinase become detectable again in the most proximal oocyte(s) that are undergoing maturation, prior to fertilization. This proximal MAP kinase activity is induced by a cue from sperm.  
Curated by authors based on online in-situ hybridization database (Y. Kohara, personal communication; http://nematode.lab.nig.ac.jp).   Expr4016 Broadly expressed in gonad.  
    Expr2013667 Single cell embryonic expression. Only cell types with an expression fraction of greater 0.2 of the maximum expressed fraction are labeled (Full data can be downloaded from http://caltech.wormbase.org/pub/wormbase/datasets-published/packer2019/). The colors represent the broad cell class to which the cell type has been assigned. The size of the point is proportional to the log2 of the numbers of cells in the dataset of that cell type. Interactive visualizations are available as a web app (https://cello.shinyapps.io/celegans/) and can also be installed as an R package (https://github.com/qinzhu/VisCello.celegans).  
    Expr2031901 Single cell embryonic expression. Only cell types with an expression fraction of greater 0.2 of the maximum expressed fraction are labeled (Full data can be downloaded from http://caltech.wormbase.org/pub/wormbase/datasets-published/packer2019/). The colors represent the broad cell class to which the cell type has been assigned. The size of the point is proportional to the log2 of the numbers of cells in the dataset of that cell type. Interactive visualizations are available as a web app (https://cello.shinyapps.io/celegans/) and can also be installed as an R package (https://github.com/qinzhu/VisCello.celegans).  
Original chronogram file: chronogram.785.xml [F43C1.2:gfp] transcriptional fusion. Chronogram1867    
    Expr3589   Diffuse in nerve cords.
    Expr1014050 Developmental gene expression time-course. Raw data can be downloaded from ftp://caltech.wormbase.org/pub/wormbase/datasets-published/levin2012  

40 GO Annotation

Annotation Extension Qualifier
  enables
  enables
has input(WB:WBGene00004804) enables
  enables
  enables
  enables
  enables
  enables
  enables
  enables
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  enables
  enables
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  enables
  involved_in
  involved_in
  involved_in
  involved_in
  involved_in
  involved_in
  enables
  involved_in
  involved_in
  involved_in
  involved_in
  involved_in
  involved_in
  involved_in
  involved_in

14 Homologues

Type
orthologue
orthologue
orthologue
orthologue
orthologue
least diverged orthologue
least diverged orthologue
least diverged orthologue
least diverged orthologue
least diverged orthologue
orthologue
orthologue
orthologue
orthologue

1 Locations


Feature . Primary Identifier
Start End Strand
WBGene00003401 4216763 4228108 -1

40 Ontology Annotations

Annotation Extension Qualifier
  enables
  enables
has input(WB:WBGene00004804) enables
  enables
  enables
  enables
  enables
  enables
  enables
  enables
  enables
  enables
  enables
  enables
  enables
  involved_in
  involved_in
  involved_in
  involved_in
  involved_in
  involved_in
  enables
  involved_in
  involved_in
  involved_in
  involved_in
  involved_in
  involved_in
  involved_in
  involved_in

5 Regulates Expr Cluster

Regulated By Treatment Description Algorithm Primary Identifier
  Genes shown no response to MPK-1 signaling. Hierarchical clustering and the Cluster and Treeview programs were used. WBPaper00028788:No_Response_to_MPK-1
  Genes shown medium response to MPK-1 signaling. Hierarchical clustering and the Cluster and Treeview programs were used. WBPaper00028788:MPK-1_Response_Medium
  Genes shown weak response to MPK-1 signaling. Hierarchical clustering and the Cluster and Treeview programs were used. WBPaper00028788:MPK-1_Response_Weak
  Genes shown strong response to MPK-1 signaling. Hierarchical clustering and the Cluster and Treeview programs were used. WBPaper00028788:MPK-1_Response_Strong
  Genes upregulated by mpk-1(ga111). Hierarchical clustering and the Cluster and Treeview programs were used. WBPaper00028788:Upregulated_in_mpk-1(ga111)

1 Sequence

Length
11346

1 Sequence Ontology Term

Identifier Name Description
gene  

20 Strains

WormBase ID
WBStrain00026496
WBStrain00027313
WBStrain00027327
WBStrain00028924
WBStrain00030717
WBStrain00030719
WBStrain00033908
WBStrain00033906
WBStrain00033914
WBStrain00033911
WBStrain00033912
WBStrain00033910
WBStrain00048933
WBStrain00051720
WBStrain00051721
WBStrain00051722
WBStrain00051646
WBStrain00052064
WBStrain00052063
WBStrain00003817

1 Upstream Intergenic Region

WormBase ID Name Sequence Name Length (nt) Chromosome Location Organism
intergenic_region_chrIII_4228109..4229140   1032 III: 4228109-4229140 Caenorhabditis elegans