Opsins and clusters of sensory G-protein-coupled receptors in the sea urchin genome
Open accessAbstract
Rhodopsin-type G-protein-coupled receptors (GPCRs) contribute the majority of sensory receptors in vertebrates. With 979 members, they form the largest GPCR family in the sequenced sea urchin genome, constituting more than 3% of all predicted genes. The sea urchin genome encodes at least six Opsin proteins. Of these, one rhabdomeric, one ciliary and two G(o)-type Opsins can be assigned to ancient bilaterian Opsin subfamilies. Moreover, we identified four greatly expanded subfamilies of rhodopsin-type GPCRs that we call sea urchin specific rapidly expanded lineages of GPCRs (surreal-GPCRs). Our analysis of two of these groups revealed genomic clustering and single-exon gene structures similar to the most expanded group of vertebrate rhodopsin-type GPCRs, the olfactory receptors. We hypothesize that these genes arose by rapid duplication in the echinoid lineage and act as chemosensory receptors of the animal. In support of this, group B surreal-GPCRs are most prominently expressed in distinct classes of pedicellariae and tube feet of the adult sea urchin, structures that have previously been shown to react to chemical stimuli and to harbor sensory neurons in echinoderms. Notably, these structures also express different opsins, indicating that sea urchins possess an intricate molecular set-up to sense their environment.
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- Findingown data
Sp-Opsin4 is a member of the rhabdomeric Opsin family.
“Sp-Opsin4 is a clear member of the r-Opsin family”
Quote in the paperSubstantial claim - Findingown data
The sea urchin genome contains 979 rhodopsin-type GPCRs.
“With 979 members, they form the largest GPCR family in the sequenced sea urchin genome, constituting more than 3% of all predicted genes.”
- Also stated in: § Rhodopsin-type GPCRs constitute the largest GPCR superfamily in the sea urchin genome · hedged
“we could identify 979 rhodopsin-type GPCRs in the S. purpuratus proteome predicted from the first genome assembly.”
Quote in the paperSubstantial claim - Also stated in: § Rhodopsin-type GPCRs constitute the largest GPCR superfamily in the sea urchin genome · hedged
- Findingown data
Many predicted sea urchin GPCRs show only distant similarity to known bilaterian GPCRs.
“However, we noted that many of the predicted molecules showed only distant similarity to known bilaterian GPCRs.”
Quote in the paperSubstantial claim - Findingown data
Sp-Opsin2 and Sp-Opsin5 show no clear affiliation to any larger Opsin subfamily in phylogenetic and sequence analyses.
“The two remaining sea urchin opsins, Sp-Opsin2 and Sp-Opsin5, show no clear affiliation to any of the larger Opsin subfamilies in phylogenetic and sequence analyses (Figs.1A, B).”
Quote in the paperSubstantial claim - Findingown data
Pedicellariae and tube feet also express different opsins.
“Notably, these structures also express different opsins”
Quote in the paperSubstantial claim
- Findingown data
Rhodopsin-type GPCRs are the largest GPCR superfamily in the sea urchin genome.
“they form the largest GPCR family in the sequenced sea urchin genome”
- Also stated in: § Rhodopsin-type GPCRs constitute the largest GPCR superfamily in the sea urchin genome · boosted
“Rhodopsin-type GPCRs are by far the largest GPCR superfamily in the sea urchin genome”
Quote in the paperSubstantial claim - Also stated in: § Rhodopsin-type GPCRs constitute the largest GPCR superfamily in the sea urchin genome · boosted
- Findingown data
Transcripts of Sp-opsin1, Sp-opsin2, Sp-opsin3.1, and Sp-opsin4 are also present at high levels in tube feet.
“Besides their prominent expression in pedicellariae, transcripts of Sp-opsin1, Sp-opsin2, Sp-opsin3.1 and Sp-opsin4 were also present at high levels in the tube feet (Fig. 2A)”
Quote in the paperNot a substantial claim - Findingown data
Sp-opsin4 is specifically expressed in 1-2 cells close to the ciliated band at the tip of the postoral arms of two-week-old sea urchin larvae.
“This analysis revealed that Sp-opsin4 is specifically expressed in 1-2 cells close to the ciliated band at the tip of the postoral arms of two week old larvae (Figs.2B3, 4, 5).”
- Also stated in: § Sea urchins possess a surprisingly large molecular sensory repertoire · plain
“Our wholemount in situ hybridization analysis revealed that Sp-opsin4 is expressed in a few cells close to the ciliated band on the tip of the postoral arms of two week old sea urchin larvae.”
Quote in the paperSubstantial claim - Also stated in: § Sea urchins possess a surprisingly large molecular sensory repertoire · plain
- Findingown data
Sea urchin GPCRs bear similarity to a broad range of diverse receptor subfamilies.
“This analysis showed that sea urchin GPCRs bear similarity to a broad range of diverse receptor subfamilies (Table S3).”
Quote in the paperSubstantial claim - Findingown data
No opsin transcripts were detected in testis, ovary, ampulla, axial gland, coelomocyte, or gut.
“whereas no opsin transcripts were detected in testis, ovary, ampulla, axial gland, coelomocyte and gut (Fig. 2A).”
Quote in the paperNot a substantial claim - Findingown data
Among sampled adult sea urchin tissues, all tested opsins are most prominently transcribed in pedicellariae.
“Notably, the sampling of different tissues revealed that all opsins are most prominently transcribed in pedicellariae (Fig. 2A).”
Quote in the paperSubstantial claim - Findingown data
Of the sea urchin rhodopsin-type GPCRs that had counterparts in the reference database, only 55 were also best reciprocal hits.
“Of those that had counterparts in the reference database, only 55 were also best reciprocal hits.”
Quote in the paperSubstantial claim - Findingown data
These GPCR domains are more closely related to other sea urchin GPCRs than to those of other organisms.
“Moreover, these GPCR domains are more closely related to other sea urchin GPCRs than to those of other organisms”
Quote in the paperSubstantial claim - Findingown data
Rhodopsin-type GPCRs constitute more than 3% of all predicted genes in the sea urchin genome.
“constituting more than 3% of all predicted genes.”
Quote in the paperNot a substantial claim - Findingown data
Except for Sp-opsin5, transcription of all tested opsins is detectable in one-week-old sea urchin larvae, the earliest developmental stage investigated.
“Except for Sp-opsin5, transcription of all opsins could already be detected in one week old larvae, the earliest developmental stage investigated (Fig. 2A).”
Quote in the paperNot a substantial claim - Findingown data
Of the 979 rhodopsin-type GPCRs, 5 did not match any known protein within the cut-off range (e-value 0.001).
“5 GPCRs did not match any known protein within the cut-off range (e-value of 0.001).”
Quote in the paperNot a substantial claim - Findingown data
Neural ring and tube express low levels of Sp-opsin1.
“In addition, neural ring and tube express low levels of Sp-opsin1”
Quote in the paperNot a substantial claim - Findingown data
At least three pan-bilaterian Opsin families are present in the sea urchin.
“Phylogenetic analysis using neighbor joining and maximum likelihood methods, combined with an examination of the diagnostic amino acid fingerprints (Arendt et al., 2004), reveals that at least three pan-bilaterian Opsin families are present in the sea urchin (Fig. 1A).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Arendt et al., 2004 source
Not checked in the free run.
- Findingnone given
The S. purpuratus genome contains a G_o G-protein (SPU_007553).
“The S. purpuratus genome also contains a G o G-protein (SPU_007553)”
Quote in the paperSubstantial claim - Findingown data
Two surreal-GPCR groups have single-exon gene structures similar to vertebrate olfactory receptors.
“Our analysis of two of these groups revealed genomic clustering and single-exon gene structures similar to the most expanded group of vertebrate rhodopsin-type GPCRs, the olfactory receptors.”
Quote in the paperSubstantial claim - Findingown data
Sp-Opsin3.1 and Sp-Opsin3.2 form a well-supported group with the Patinopecten G_o-Opsin and two Amphioxus Opsins.
“Strikingly, two Opsins, Sp-Opsin3.1 and Sp-Opsin3.2,form one well-supported group with the Patinopecten G o -Opsin (Kojima et al., 1997) and two Opsins from Amphioxus (Koyanagi et al., 2002) (Fig. 1A)”
Quote in the paperSubstantial claim0 of 2 sources checked - Findingown data
Group B surreal-GPCRs are most prominently expressed in distinct classes of pedicellariae and tube feet of the adult sea urchin.
“group B surreal-GPCRs are most prominently expressed in distinct classes of pedicellariae and tube feet of the adult sea urchin”
Quote in the paperSubstantial claim - Findingown data
The sea urchin genome contains four greatly expanded subfamilies of rhodopsin-type GPCRs called surreal-GPCRs.
“we identified four greatly expanded subfamilies of rhodopsin-type GPCRs that we call sea urchin specific rapidly expanded lineages of GPCRs (surreal-GPCRs).”
Quote in the paperSubstantial claim - Findingown data
All-against-all comparisons of sea urchin rhodopsin-type GPCR domains revealed several groups with high mutual similarity.
“This analysis revealed several groups of sea urchin GPCR domains with high mutual similarity (Fig. 3A).”
Quote in the paperSubstantial claim - Findingown data
All five tested sea urchin opsins (Sp-opsin1, Sp-opsin2, Sp-opsin4, Sp-opsin5, and Sp-opsin3.1) are expressed in adult sea urchins.
“In adult sea urchins, all five tested opsins are expressed.”
Quote in the paperNot a substantial claim - Findingown data
Sp-Opsin1 belongs to the ciliary Opsin family.
“Sp-Opsin1 belongs to the c-Opsin family”
Quote in the paperSubstantial claim - Findingown data
The three sampled major pedicellaria classes express different opsin subsets at different levels.
“This analysis revealed that all three classes express different opsin subsets at different levels (Fig. 2A).”
Quote in the paperSubstantial claim - Findingcitation
Sp-opsin4 is orthologous to the rhabdomeric opsin/GPCR type present in the eyes of primary ciliated larvae.
“As a representative of the rhabdomeric opsin family, Sp-opsin4 is orthologous to the GPCR type present in the eyes of primary ciliated larvae (Arendt et al., 2002).”
- Also stated in: § Sea urchins possess a surprisingly large molecular sensory repertoire · hedged
“As indicated by our phylogenetic analysis, Sp-opsin4 is orthologous to the r-opsin expressed in the rhabdomeric photoreceptor cells (PRCs) of the larval and adult eyes of primary ciliated protostome larvae, such as the trochophora larvae of mollusks and polychaetes.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Arendt et al., 2002 source
Not checked in the free run.
- Also stated in: § Sea urchins possess a surprisingly large molecular sensory repertoire · hedged
- Findingown data
Two surreal-GPCR groups show genomic clustering similar to vertebrate olfactory receptors.
“Our analysis of two of these groups revealed genomic clustering and single-exon gene structures similar to the most expanded group of vertebrate rhodopsin-type GPCRs, the olfactory receptors.”
Quote in the paperSubstantial claim - Findingnone given
No vertebrate, insect or nematode G_o-Opsin family member is present in sequence databases to date.
“Remarkably, no vertebrate, insect or nematode G o -Opsin family member is present in the sequence databases to date”
Quote in the paperSubstantial claim - Findingown data
The sea urchin genome encodes at least six Opsin proteins.
“The sea urchin genome encodes at least six Opsin proteins.”
- Also stated in: § Rhodopsin-type GPCRs constitute the largest GPCR superfamily in the sea urchin genome · plain
“Among the predicted sea urchin genes, six encode bona fide Opsins (Fig. 1A).”
Quote in the paperSubstantial claim - Also stated in: § Rhodopsin-type GPCRs constitute the largest GPCR superfamily in the sea urchin genome · plain
- Findingown data
One group of surreal-GPCR-B27 stained cells locates at the tip of the postoral arm, in a position comparable to the Sp-opsin4-positive cells.
“Interestingly, one group of stained cells locates at the tip of the postoral arm (Fig. 5B3), in a position comparable to the Sp-opsin4-positive cells (Figs. 2B3-5)”
Quote in the paperSubstantial claim - Findingown data
Fast-evolving intronless genes are also found outside the major clusters, for instance in the minor classes e and f.
“The fast-evolving intronless genes are also found outside of the major clusters, for instance in the minor classes e and f (Fig. 3A).”
Quote in the paperNot a substantial claim - Findingown data
Surreal-GPCR-B27 is present in isolated single cells in the ectoderm at different positions in two-week-old larvae.
“We find that surreal-GPCR-B27 is present in isolated single cells in the ectoderm at different positions (Figs.5B2-5).”
Quote in the paperSubstantial claim - Findingcitation
Only limited expression was found for surreal-GPCR genes in the transcriptome map of Samanta et al. (2006).
“In contrast, only limited expression was found for surreal-GPCR genes in the transcriptome map of Samanta et al. (2006).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Samanta et al., 2006 source
Not checked in the free run.
- Findingown data
In both cases, these clusters are located at one end of the scaffold.
“In both cases, these clusters are located at one end of the scaffold.”
Quote in the paperSubstantial claim - Findingown data
Another scaffold (Scaffold_v2_66648) harbored 9 genes related to group C surreal-GPCRs in tandem duplication, in addition to one reversed gene.
“Similarly, another scaffold (Scaffold_v2_66648) harbored 9 genes related to group C surreal-GPCRs in tandem duplication, in addition to one reversed gene (Fig. 3D).”
Quote in the paperNot a substantial claim - Findingown data
For group A surreal-GPCRs, the profile HMM search produced weak, similarly distant relationships to both muscarinic acetylcholine receptors and serotonin receptors.
“For group A surreal-GPCRs, the profile HMM search produced weak, yet similarly distant relationships to both muscarinic acetylcholine receptors and serotonin receptors.”
Quote in the paperNot a substantial claim - Findingown data
Based on reciprocal similarity profiles, four major groups of surreal-GPCRs (A-D) and multiple small ones (e and f) can be distinguished in the sea urchin genome.
“Based on their reciprocal similarity profiles, we can distinguish four major groups of surreal-GPCRs in the sea urchin genome (labeled A-D in Fig. 3A), and multiple small ones (exemplified as e and f).”
Quote in the paperSubstantial claim - Findingown data
The few representatives of the group B surreal-GPCRs tested so far show distinct expression profiles in tube feet and different pedicellariae classes of the adult.
“Another strong line of evidence is provided by the fact that the few representatives of the group B surreal-GPCRs that we have tested so far show distinct expression profiles in tube feet and different pedicellariae classes of the adult”
Quote in the paperSubstantial claim - Findingown data
Group C surreal-GPCRs show overall higher divergence than group B surreal-GPCRs.
“This group shows an overall higher divergence than the group B surreal-GPCRs (Fig. 4B).”
Quote in the paperNot a substantial claim - Findingcitation
In both group B and group C surreal-GPCRs, the third intracellular loop (ic3) and the C-terminal tail show high divergence, as is common for GPCRs in general.
“In both group B and group C surreal-GPCRs, the third intracellular loop (ic3) as well as the C-terminal tail show a high divergence, as common for GPCRs in general (Mathi et al., 1997).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Mathi et al., 1997 source
Not checked in the free run.
- Findingown data
In group B surreal-GPCRs, variable positions (residues not shared among 60% of sequences) are distributed across all seven-transmembrane domains.
“This comparison revealed that variable positions (defined as residues not shared among 60% of the sequences) were distributed across all seven-transmembrane domains (Fig. 4A).”
Quote in the paperNot a substantial claim - Findingown data
Group B surreal-GPCRs are highly expressed in globiferous pedicellariae.
“The high expression of group B surreal-GPCR members in globiferous pedicellariae (Fig. 5A) is particularly intriguing”
Quote in the paperNot a substantial claim - Findingown data
In the second genome assembly, one scaffold (Scaffold_v2_21524) contained 10 group B surreal-GPCR genes in a tandem array of single-exon genes, covering a locus of around 100 kb.
“One scaffold (Scaffold_v2_21524) was found to contain 10 group B surreal-GPCRgenes in a tandem array of single-exon genes, covering a locus of around 100 kb (Fig. 3C).”
Quote in the paperNot a substantial claim - Findingnone given
Go-Opsins appear to be absent from other deuterostome groups, including vertebrates.
“Interestingly, G o -Opsins appear to be absent from the other deuterostome groups, including vertebrates”
Quote in the paperSubstantial claim - Findingown data
A DRIP78 binding sequence (FxxxFxxxF) is conserved in the C-termini of a subset of group B and group C surreal-GPCR sequences.
“In line with such a role, we find a dopamine receptor interacting protein 78 (DRIP78) binding sequence (FxxxFxxxF) (Bermak et al., 2001) conserved in the C-termini of a subset of sequences ( SPU_006834, SPU_008884, SPU_010318, SPU_010318; SPU_000576, SPU_015119, SPU_006431).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Bermak et al., 2001 source
Not checked in the free run.
- Findingown data
The distribution of amino acid variance in sea urchin group B surreal-GPCRs is different from both mouse and Drosophila olfactory GPCRs.
“This is different from both mouse and Drosophila olfactory GPCRs.”
Quote in the paperSubstantial claim - Findingown data
All three approaches (EST resources, QPCR, in situ hybridizations) independently revealed that different members of the group B surreal-GPCR family are present at larval and adult stages.
“All three approaches independently revealed that different members of the group B surreal-GPCR family are present at larval and adult stages (Figs.5A, B and Table S4).”
Quote in the paperSubstantial claim - Findingnone given
Many group B surreal-GPCRs are encoded by single exons, including the members in the genomic cluster mentioned above.
“A characteristic feature of group B surreal-GPCRs that is possibly linked to their expansion is that many of them are encoded by single exons, including the members in the genomic cluster mentioned above.”
Quote in the paperNot a substantial claim - Findingown data
The four main surreal-GPCR expansions together account for a total of around 450 rhodopsin-type GPCRs in the sea urchin genome.
“The four main surreal-GPCR expansions together account for a total of around 450 rhodopsin-type GPCRs in the sea urchin genome (Table S3).”
Quote in the paperSubstantial claim - Findingcitation
The unequal distribution of variable amino acids in the group C surreal-GPCRs, and in particular their strongest variability in the fourth and fifth TM helix, is reminiscent of the classical vertebrate olfactory receptors.
“In contrast, the unequal distribution of variable amino acids in the group C surreal-GPCRs, and in particular their strongest variability in the fourth and fifth TM helix, is reminiscent of the classical vertebrate olfactory receptors (Buck and Axel, 1991;Pilpel and Lancet, 1999).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Buck and Axel, 1991
Not checked in the free run.
- cites Pilpel and Lancet, 1999
Not checked in the free run.
- Findingown data
Additional stained cells were observable between the anterior lateral arm and the apical plate.
“Additional stained cells were observable between the anterior lateral arm and the apical plate (Figs.2B2, 4, 5).”
Quote in the paperNot a substantial claim - Findingown data
In group C surreal-GPCRs, rapidly changing amino acid residues preferentially cluster in the third to sixth transmembrane helix, with TM IV and TM V being the most variable helices.
“Moreover, the rapidly changing amino acid residues are not distributed equally across the seven transmembrane domains (as is the case for the group B surreal-GPCRs), but preferentially cluster in the third to sixth transmembrane helix, with the TM IV and TM V being the most variable helices (Fig. 4B).”
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No expression of group B surreal-GPCRs is detected in testis, ovary, coelomocyte, axial gland, and gut tissue.
“No expression is detected in testis, ovary, coelomocyte, axial gland and gut tissue”
Quote in the paperSubstantial claim - Findingown data
The largest cluster in the first genome assembly contained 12 genes in tandem orientation on scaffold 548.
“The largest cluster in the first genome assembly was found to contain 12 genes in tandem orientation on scaffold 548 (highlighted by red circles in Fig. 3B).”
Quote in the paperNot a substantial claim - Findingown data
The 8 remaining non-clustered group B genes were either located on small genomic scaffolds or at the border of medium-sized scaffolds containing a maximum of 3 other genes.
“Without exception, the 8 remaining, non-clustered genes were either located on small genomic scaffolds, or they were lying at the border of medium-sized scaffolds containing a maximum of 3 other genes.”
Quote in the paperSubstantial claim - Findingown data
Low additional expression (at least 10 times lower than in tube feet and globiferous and trifoliate pedicellariae) can only be detected in three additional adult tissues: the neural tube and ring, as well as the ampulla.
“Low additional expression (at least 10 times lower than in tube feet and globiferous and trifoliate pedicellariae) can only be detected in three additional adults tissues; the neural tube and ring, as well as in the ampulla (Fig. 5A).”
Quote in the paperSubstantial claim - Findingown data
In a maximum likelihood analysis, representative bilaterian opsins fall into three subfamilies with significant support.
“Representative bilaterian Opsin members cluster into three significantly supported subfamilies in a maximum likelihood analysis.”
Quote in the paperSubstantial claim - Findingown data
The genomic average of single-exon genes is 15% of all predicted gene models.
“This proportion is in stark contrast to the genomic average of single-exon genes that we calculate to be 15% of all predicted gene models.”
Quote in the paperSubstantial claim - Findingown data
Systematic analysis indicated multiple small expansions in distinct rhodopsin GPCR subfamilies.
“In addition, our systematic analysis indicated multiple small expansions in distinct rhodopsin GPCR subfamilies.”
Quote in the paperSubstantial claim - Findingown data
Profile HMM search provided evidence that group C surreal-GPCRs are most similar to the opioid receptor subfamily (IPR001418).
“A more sensitive search, using profile hidden Markov models (profile HMMs), provided evidence that group C is most similar to the opioid receptor subfamily (IPR001418)”
Quote in the paperSubstantial claim - Findingnone given
The currently predicted S. purpuratus proteome does not contain any clear retinochrome (RGR Opsins).
“For instance, the currently predicted S. purpuratus proteome does not contain any clear retinochrome (RGR Opsins)”
Quote in the paperSubstantial claim - Findingown data
The genome of S. purpuratus includes rapidly expanded, sea urchin-specific GPCR lineages known as surreal-GPCRs.
“The S. purpuratus genome contains sea urchin specific rapidly expanded lineages of GPCRs (surreal-GPCRs).”
Quote in the paperSubstantial claim - Findingnone given
Group D surreal-GPCRs possess the highest similarity to GPCRs of other organisms.
“We noted that group D surreal-GPCRs also possess the highest similarity to GPCRs of other organisms.”
Quote in the paperNot a substantial claim - Findingown data
Most group C surreal-GPCRs have an intronless gene structure.
“The intronless gene structure is also found in most group C surreal-GPCRs (see Table S3).”
Quote in the paperNot a substantial claim - Findingown data
The sea urchin has three ancestral opsin types.
“It is noteworthy that the sea urchin does not only possess three ancestral Opsin types, as we describe in our study”
Quote in the paperSubstantial claim - Findingown data
Analysis of opsin genes revealed that three ancestral bilaterian classes, the rhabdomeric, ciliary, and Go class, are present in the sea urchin.
“Our analysis of opsin genes has revealed that three ancestral bilaterian classes, the rhabdomeric, ciliary and G o class are present in the sea urchin.”
Quote in the paperSubstantial claim - Findingown data
Four out of six tested group B surreal-GPCR genes are clearly expressed in larvae.
“Based on QPCR data, four out of six tested genes are clearly expressed at this stage (Fig. 5A).”
Quote in the paperSubstantial claim - Findingown data
Out of 43 core group B surreal-GPCR sequences predicted from the first genome assembly, all but 8 were found in direct neighborhood to at least one other surreal-GPCR gene.
“Out of 43 core group B surreal-GPCR sequences predicted from the first genome assembly, all but 8 were found in direct neighborhood to at least one other surreal-GPCR gene (Fig. 3B).”
Quote in the paperSubstantial claim - Findingown data
BLAST significance values for many protein hits in the reference database are comparably weak, consistent with fast evolution of these receptor groups.
“Consistent with a fast evolution of these receptor groups, the BLAST significance values for many protein hits in the reference database are comparably weak.”
Quote in the paperNot a substantial claim - Findingnone given
The amino acid variance of the sea urchin group B surreal-GPCRs is distributed relatively evenly among the seven transmembrane domains.
“Interestingly, the amino acid variance of the sea urchin group B surreal-GPCRs is distributed relatively evenly among the seven transmembrane domains.”
Quote in the paperSubstantial claim - Findingown data
Preliminary sequence analysis of expressed group B surreal-GPCR sequences isolated from single individuals reveals several cases where more than two variants exist for a single gene present in the genome.
“preliminary sequence analysis of expressed group B surreal-GPCR sequences isolated from single individuals reveals several cases where more than two variants exist for a single gene present in the genome”
Quote in the paperSubstantial claim - Findingown data
Sp-opsin4 expression is detected in two-week-old larvae.
“Expression of Sp-opsin4 as detected by whole-mount in situ hybridization in two-week-old larvae.”
Quote in the paperNot a substantial claim - Findingown data
Scaffold_v2_66648 contains ten genes related to group C surreal-GPCRs.
“10 genes related to group C surreal-GPCRs on Scaffold_v2_66648.”
Quote in the paperNot a substantial claim - Findingown data
Surreal-GPCR-B27 expression is detected in two-week-old larvae.
“Expression of surreal-GPCR-B27 as detected by whole-mount in situ hybridization in twoweek-old larvae.”
Quote in the paperNot a substantial claim - Findingown data
Different opsin classes are expressed in both pedicellariae and tube feet.
“the expression of distinct opsin classes in pedicellariae and tube feet is particularly remarkable.”
Quote in the paperSubstantial claim - Findingown data
Selected surreal-GPCRs are expressed in both larval and adult sea urchins.
“Expression of representative surreal-GPCRs in the sea urchin larva and adult.”
Quote in the paperSubstantial claim - Findingown data
Amino acid residues differ among group B and group C surreal-GPCRs.
“Variability of amino acid residues in group B and group C surreal-GPCRs.”
Quote in the paperNot a substantial claim - Findingown data
Scaffold_v2_21524 contains ten group B surreal-GPCR representatives.
“Ten representatives of group B surreal-GPCRs are shown in their genomic location on Scaffold_v2_21524.”
Quote in the paperNot a substantial claim - Findingown data
The study examined 979 rhodopsin-type GPCR domains in the S. purpuratus genome.
“Clustered matrix showing the pairwise similarities between all 979 investigated rhodopsin-type sea urchin GPCR domains”
Quote in the paperSubstantial claim - Interpretationreasoning
The G_o-Opsin family might be connected to cell types present in marine species with primary ciliated larvae.
“indicating that the G o -Opsin family might be connected to cell types present in marine species with primary ciliated larvae.”
Quote in the paperSubstantial claim - Premisecitation
The sea urchin occupies a phylogenetic position at the base of deuterostomes.
“Given the phylogenetic position of the sea urchin at the base of deuterostomes (Delsuc et al., 2006)”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Delsuc et al., 2006 source
Not checked in the free run.
- Interpretationreasoning
The link between G_o-Opsin and the G_o G-protein may be evolutionarily conserved.
“consistent with the possibility that the link between G o -Opsin and the G o G-protein is evolutionarily conserved.”
Quote in the paperNot a substantial claim - Interpretationown data
The absolute number and frequency of sea urchin rhodopsin-type GPCRs are comparable to several mammalian genomes.
“The absolute number and the frequency of sea urchin rhodopsin-type GPCRs is thus comparable to several mammalian genomes”
Quote in the paperSubstantial claim - Interpretationreasoning
Surreal-GPCRs act as chemosensory receptors of the animal.
“and act as chemosensory receptors of the animal.”
- Also stated in: § Introduction · hedged
“We argue that these receptors are likely to function as chemoreceptors of the sea urchin.”
Quote in the paperSubstantial claim - Also stated in: § Introduction · hedged
- Interpretationown data
The existence of an ancestral bilaterian G_o-Opsin family is supported.
“thus supporting the existence of an ancestral bilaterian G o -Opsin family.”
Quote in the paperSubstantial claim - Interpretationreasoning
The sea urchin has more than one family of chemoreceptors.
“we suggest that the sea urchin has more than one family of chemoreceptors.”
Quote in the paperSubstantial claim - Premisecitation
Sea urchin rhodopsin-type GPCRs contain clear counterparts of functionally characterized GPCRs.
“Besides the Opsins, the rhodopsin-type sea urchin GPCRs contain other clear counterparts of functionally characterized GPCRs (Burke et al., 2006;Whittaker et al., 2006, and see below).”
Quote in the paperNot a substantial claim0 of 2 sources checked - Interpretationreasoning
Sp-Opsin2 and Sp-Opsin5 are likely highly derived members of one of the known Opsin groups.
“Therefore, these proteins are likely to be highly derived members of one of the known groups.”
Quote in the paperSubstantial claim - Premisecitation
Experiments using antibodies against bovine rhodopsin indicated the presence of a ciliary-type Opsin in asteroids and ophiuroids.
“experiments using antibodies directed against bovine rhodopsin indicated the presence of a ciliary-type Opsin in asteroids and ophiuroids (Johnsen, 1997).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Johnsen, 1997 source
Not checked in the free run.
- Interpretationreasoning
The existence of two G_o-Opsin family members in the sea urchin is likely due to a recent duplication.
“The existence of two G o -Opsin family members in the sea urchin is likely to be due to a recent duplication.”
Quote in the paperNot a substantial claim - Interpretationreasoning
Sea urchins possess an intricate molecular set-up to sense their environment.
“indicating that sea urchins possess an intricate molecular set-up to sense their environment.”
Quote in the paperSubstantial claim - Interpretationown data
Both ciliary and rhabdomeric Opsin subfamilies were present at the beginning of Bilateria.
“confirming that both Opsin subfamilies were present at the beginning of Bilateria.”
Quote in the paperSubstantial claim - Study design and conductown data
The accuracy of whole-mount in situ hybridization results was confirmed by control experiments using sense probes.
“The accuracy of the whole-mount in situ hybridization results was confirmed by control experiments using sense probes.”
Quote in the paperNot a substantial claim - Interpretationown data
A major proportion of sea urchin rhodopsin-type GPCRs are not part of 1:1 orthologous relationships.
“This already indicated that a major proportion of sea urchin rhodopsin-type GPCRs was not part of 1:1 orthologous relationships.”
Quote in the paperSubstantial claim - Interpretationown data
The sea urchin uses a complex sensory receptor repertoire localized in decentralized appendages all over its body to analyze its environment.
“We thus provide evidence that the sea urchin uses a complex sensory receptor repertoire localized in decentralized appendages all over its body to analyze its environment.”
Quote in the paperSubstantial claim - Interpretationreasoning
The dissimilarity between the distribution of divergent residues in vertebrates, sea urchin, and insects suggests that the chemosensory receptors in these groups evolved independently.
“The dissimilarity between the distribution of divergent residues in vertebrates, sea urchin and insects suggests that the chemosensory receptors in these groups evolved independently.”
Quote in the paperSubstantial claim - Interpretationown data
The sea urchin genome has partially retained ancestral genomic complexity that has been secondarily reduced in other evolutionary lineages.
“First, there are cases of ancestral rhodopsin-type GPCR ortholog genes that have been retained in the sea urchin genome, whereas they have been lost from other animals, indicating that the sea urchin has partially retained ancestral genomic complexity that has been secondarily reduced in other evolutionary lineages.”
Quote in the paperSubstantial claim - Interpretationown data
The sea urchin rhodopsin-type GPCR repertoire contains several largely expanded subfamilies that appear specific to the sea urchin lineage.
“Together, these findings show that the sea urchin rhodopsin-type GPCR repertoire contains several largely expanded subfamilies that appear to be specific to the sea urchin lineage.”
Quote in the paperSubstantial claim - Interpretationown data
The genome predictions for surreal-GPCR diversity reflect a lower bound estimate of the real surreal-GPCR diversity.
“indicating that the genome predictions rather reflect a lower bound estimate of the real surreal-GPCR diversity (data not shown).”
Quote in the paperSubstantial claim - Interpretationreasoning
Some additional genomic clustering of surreal-GPCRs was not detectable due to incomplete assembly of the genome.
“This suggested that some additional genomic clustering of surreal-GPCRs was not detectable due to incomplete assembly of the genome.”
Quote in the paperSubstantial claim - Study design and conductown data
Embryos stained with sense control RNA never exhibited any such single cell staining.
“Embryos stained with sense control RNA were never exhibiting any such single cell staining (Fig. 5B1).”
Quote in the paperSubstantial claim - Interpretationown data
At least the members of the current group B surreal-GPCR sampling have a clear tendency to be almost exclusively present in sensory appendages, as well as in the nervous system of the sea urchin.
“indicating that at least the members of our current group B surreal-GPCR sampling have a clear tendency to be almost exclusively present in sensory appendages, as well as in the nervous system of the sea urchin.”
Quote in the paperSubstantial claim - Interpretationown data
Sea urchin rhodopsin-type GPCRs have undergone multiple independent expansions, causing a considerable gain of receptors in the sea urchin lineage.
“Second, as our discovery of surreal-GPCRs indicates, sea urchin rhodopsin-type GPCRs have undergone multiple independent expansions, causing a considerable gain of receptors in the sea urchin lineage.”
Quote in the paperSubstantial claim - Interpretationreasoning
The sea urchin appears to retain two full sets of ancestral light-sensitive molecules.
“With two apparently non-reduced sets of ancestral light-sensitive molecules”
Quote in the paperSubstantial claim - Interpretationreasoning
The unexpectedly high abundance of rhodopsin-type GPCRs in the sea urchin is consistent with a rich molecular repertoire of sensory molecules in this species.
“This unexpectedly high abundance of rhodopsin-type GPCRs in the sea urchin is consistent with a rich molecular repertoire of sensory molecules in this species.”
Quote in the paperSubstantial claim - Premisecitation
In Drosophila olfactory GPCRs, the most divergent regions are the first extracellular loop, the third transmembrane helix, as well as the first and second transmembrane domains.
“whereas in Drosophila the most divergent regions are the first extracellular loop, the third transmembrane helix, as well as the first and second transmembrane domains (Clyne et al., 1999).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Clyne et al., 1999 source
Not checked in the free run.
- Interpretationown data
A sensory cell cluster might be located at the tip of these arms.
“indicating that a sensory cell cluster might be located at the tip of these arms.”
Quote in the paperSubstantial claim - Premisecitation
Three ancient bilaterian cryptochrome families have been reported in the sea urchin.
“has also been reported to contain three ancient bilaterian families of Cryptochromes (Rubin et al., 2006).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Rubin et al., 2006 source
Not checked in the free run.
- Interpretationown data
These GPCR domains belong to paralogous groups.
“indicating that they belong to paralogous groups.”
Quote in the paperSubstantial claim - Interpretationown data
The ancestor of group C surreal-GPCRs lay within the opioid receptor subfamily.
“suggesting that the ancestor of this group lay within this subfamily.”
Quote in the paperSubstantial claim - Interpretationreasoning
Group B and C surreal-GPCRs encode putative chemoreceptors of the sea urchin.
“Group B and C surreal-GPCRs encode putative chemoreceptors of the sea urchin”
- Also stated in: § Sea urchins possess a surprisingly large molecular sensory repertoire · hedged
“Our argument that the members of the group B/C surreal-GPCRs that we investigated more closely are likely to be chemosensory receptors therefore does not build on sequence similarities to other known chemosensory receptor families.”
Quote in the paperSubstantial claim- Rests on an assumption: Specific rapid and large expansions of transmembrane domain genes are common to molecules that animals use to detect a large variety of divergent molecules, as in the immune system and chemoreception.
- Also stated in: § Sea urchins possess a surprisingly large molecular sensory repertoire · hedged
- Premisecitation
Some pedicellariae types continue to respond after being isolated from the animal.
“at least some types of pedicellariae have been shown to retain their responsive behavior after isolation from the animal (Campbell, 1974).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Campbell, 1974 source
Not checked in the free run.
- Interpretationreasoning
The sea urchin is a suitable model for investigating interactions between its opsin and cryptochrome systems.
“the sea urchin will be a well-suited model to study the cross-talk between these systems.”
Quote in the paperNot a substantial claim - Interpretationown data
The data indicate that pedicellariae are probably light-sensitive.
“So far, it had not been found that pedicellariae are themselves light-sensitive, but our data suggest that this is likely.”
Quote in the paperSubstantial claim - Interpretationcitation
In adult urchins, integrative circuits are thought to control local pedicellariae responses.
“In the adult urchin, such integrative circuits are expected to govern local responses of the pedicellariae”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Campbell, 1974 source
Not checked in the free run.
- Interpretationown data
Putative photo- and chemosensory receptors identified in the S. purpuratus genome offer candidates for characterizing the sea urchin's sensory input.
“the identification of putative photo-and chemosensory receptors in the S. purpuratus genome provides candidates to characterize the sensory input of the sea urchin”
Quote in the paperSubstantial claim
- May combine two claims. 32
- The quote only partly states this claim. 2
- Classified as probably not a substantial claim. 18
May combine two claims.
Findingown dataThe six predicted sea urchin Opsins are bona fide Opsins because they contain the key lysine residue for Schiff base formation, share Opsin-specific amino acids, and cluster in phylogenetic trees.
“These rhodopsin-type GPCRs are Opsins based on three characteristics: (1) the presence of the key lysine residue necessary for the Schiff base formation with retinal (Fig. 1B); (2) amino acids specifically shared among Opsin (Fig. 1B and data not shown); and (3) their well-supported clustering in phylogenetic trees (Fig. 1A and data not shown).”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataRepresentatives of both opsins and presumed chemosensory receptors are specifically and differentially expressed in tube feet and pedicellariae, but not in other tissues such as testes, ovaries, axial gland, coelomocytes or gut.
“In agreement with their presumed function, we find representatives of both opsins and the presumed chemosensory receptors specifically and differentially expressed in prominent sensory structures of the adult sea urchin, the tube feet and pedicellariae, but not in other tissues, such as testes, ovaries, axial gland, coelomocytes or gut.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataSp-Opsin3.1 and Sp-Opsin3.2 are closely similar and their genes are adjacent in the sea urchin genome.
“This is suggested by the close similarity of the Sp-Opsin3.1 and Sp-Opsin3.2sequences and is further supported by the adjacent position of these genes in the sea urchin genome.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataIn group B surreal-GPCRs, additional strong variability is seen in the N- and C-terminal regions and in the third intracellular loop between TM V and VI, all of which also carry length polymorphisms.
“Additional strong variability was seen in the Nand C-terminal regions as well as in the third intracellular loop between TM V and VI, all of which also carried length polymorphisms.”
Quote in the paperNot a substantial claimMay combine two claims.
Findingown dataThe number of rhodopsin-type GPCRs in Strongylocentrotus is similar to that found in mammals and exceeds that of many other invertebrates.
“From this perspective, it is remarkable to note that the number of rhodopsin-type GPCRs in Strongylocentrotus is similar to that found in mammals and exceeds that of many other invertebrates.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataGroup A surreal-GPCRs display a lower proportion of single-exon genes, whereas group D has many genes with a high number of exons.
“Group A surreal-GPCRs display a lower proportion of single-exon genes, whereas group D has many genes with a high number of exons.”
Quote in the paperNot a substantial claimMay combine two claims.
Findingown dataIn both minor groups e and f, genomic clustering was detected, in one case including a total of twelve closely related sequences.
“In both groups, we also detected genomic clustering, in one case (e) including a total of twelve closely related sequences.”
Quote in the paperNot a substantial claimMay combine two claims.
FindingcitationThe fraction of rhodopsin-type GPCRs in the sea urchin genome (3.4%) is comparable to that reported for mouse and human (3.4% and 2.8%, respectively) and exceeds that observed in pufferfish (1.6%), nematode (2%), or fruit fly (0.6%).
“The latter fraction is well comparable to the abundance of rhodopsin-type GPCRs reported for the mouse and human genomes (3.4% and 2.8%, respectively) and exceeds that observed in the pufferfish (1.6%), nematode (2%) or fruit fly genomes (0.6%) (Waterston et al., 2002).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Waterston et al., 2002
Not checked in the free run.
May combine two claims.
Findingown dataThe model derived from group D surreal-GPCRs showed highest similarity to a pond snail protein assigned as glycoprotein hormone receptor (GR101_LYMST), and lower significance to various vertebrate receptors of the relaxin receptor group (IPR008112).
“The model derived from group D surreal-GPCRs showed highest similarity to a pond snail protein assigned as glycoprotein hormone receptor (GR101_LYMST), and, although with lower significance, to various vertebrate receptors of the relaxin receptor group (IPR008112).”
Quote in the paperNot a substantial claimMay combine two claims.
Findingown dataSix opsins are present in the sea urchin, and two of them are opsin3 paralogs.
“The magnified insert shows the six sea urchin opsins, with the two opsin3 paralogs visible as bright quadruplet in the right lower corner.”
Quote in the paperNot a substantial claimMay combine two claims.
Findingown dataThe presence of Go-Opsin orthologs in sea urchin helped confirm Go-Opsins as an ancestral group, improving bootstrap support for the monophyletic origin of Go-Opsins prior to bilaterian radiation.
“The presence of G o -Opsin orthologs is particularly revealing because the sea urchin sequences have helped to confirm G o -Opsins as an ancestral group, improving bootstrap support for the monophyletic origin of G o -Opsins prior to bilaterian radiation.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataMore than 60% of all sea urchin rhodopsin-type GPCRs are single-exon genes, many of which are only remotely related to receptors of other animals.
“A survey among all rhodopsin-type GPCRs revealed that more than 60% of all sea urchin rhodopsin-GPCRs are single-exon genes, many of which are only remotely related to receptors of other animals (Table S3).”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataRhodopsin-type GPCRs are the largest GPCR superfamily encoded by the sequenced sea urchin genome, constituting 3.4% of all currently predicted genes.
“Despite these changes, it is clear that rhodopsin-type GPCRs are the largest GPCR superfamily encoded by the sequenced sea urchin genome, constituting 3.4% of all the currently predicted genes.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataQPCR analysis revealed that within adult urchins the clearly strongest level of expression of group B surreal-GPCRs is present in pedicellariae and tube feet, which express different sets of surreal-GPCRs at different levels.
“QPCR analysis revealed that within adult urchins the clearly strongest level of the expression is present in pedicellariae and tube feet, whichsimilar to the opsinsexpress different sets of surreal-GPCRs at different levels (Fig. 5A).”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataOpsin expression occurs in one-week-old larvae and in adult tissues.
“Expression of opsins in one-week-old larvae and adult tissues as detected by QPCR.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataAmong sea urchin GPCRs, there are four major expanded lineages (groups A-D) and two smaller lineages (e and f).
“Four major expanded lineages (group A-D surreal-GPCRs) are indicated with colored bars, and two smaller ones (e, f) are marked with small letters.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown dataGroup B and group C surreal-GPCRs show length polymorphisms.
“Length polymorphisms are also indicated: Hatched lines mark positions at which other members of the group differ by the insertion of 5 or more amino acids”
Quote in the paperNot a substantial claimThe quote only partly states this claim.
Findingown dataThe group B surreal-GPCR genes on Scaffold_v2_21524 are arranged in tandem orientation.
“Note that all genes are oriented in tandem orientation.”
Quote in the paperNot a substantial claimMay combine two claims.
InterpretationreasoningMany sea urchin GPCRs display remote similarity to their cognates in other animals, and it appeared likely that they have also lost or modified important characteristics of these groups.
“However, regarding the remote similarity that many sea urchin GPCRs display to their cognates in other animals, it appeared likely that they have also lost or modified important characteristics of these groups.”
Quote in the paperSubstantial claimMay combine two claims.
Study design and conductcitationUbiquitin and 18S ribosomal RNA are expressed at constant levels during development and were used as internal controls, with 18S used to normalize all data.
“Ubiquitin and 18S ribosomal RNA, which are known to be expressed at constant levels during development (Nemer et al., 1991;Ransick et al., 2002), were used as internal controls, and 18S Ribosomal RNA was used to normalize all data.”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Nemer et al., 1991
Not checked in the free run.
- cites Ransick et al., 2002 source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationRhodopsin-type GPCRs make up the majority of sensory receptors in vertebrates.
“Rhodopsin-type G-protein-coupled receptors (GPCRs) contribute the majority of sensory receptors in vertebrates.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Ache and Young, 2005
Not checked in the free run.
May combine two claims.
PremisecitationPedicellariae and tube feet have previously been shown to react to chemical stimuli and to harbor sensory neurons in echinoderms.
“structures that have previously been shown to react to chemical stimuli and to harbor sensory neurons in echinoderms.”
Quote in the paperSubstantial claim0 of 3 sources checked- cites Campbell, 1973
Not checked in the free run.
- cites Snyder and Snyder, 1970
Not checked in the free run.
- cites Peters and Campbell, 1987
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationVertebrate olfactory receptors arose by multiple gene duplication events.
“Olfactory receptors of vertebrates arose by multiple gene duplication events (reviewed, e.g., in Ache and Young, 2005).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Ache and Young, 2005
Not checked in the free run.
May combine two claims.
PremisecitationThe secretin-type GPCR superfamily is the second largest GPCR superfamily in the sea urchin genome, with about one sixth the number of rhodopsin-type GPCRs.
“followed second by the secretintype GPCR superfamily with only one sixth of the number (Cameron et al., 2006).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Cameron et al., 2006
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationGene duplications have long been implied as a major mechanism generating evolutionary innovations.
“Gene duplications have long been implied as a major mechanism generating evolutionary innovations (Kimura, 1983;Ohno, 1970).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Kimura, 1983
Not checked in the free run.
- cites Ohno, 1970
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationFour of the sea urchin Opsins are also reported in Burke et al. (2006).
“Four of these are also reported in the independent publication by Burke et al. (2006).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Burke et al. (2006)
Not checked in the free run.
Classified as probably not a substantial claim.
PremisecitationAmong the five main GPCR superfamilies in the GRAFS classification, two encode photosensory and chemosensory functions of vertebrates: the glutamate-receptor superfamily (pheromone and taste) and the rhodopsin-type superfamily (light, olfaction and possibly pheromones).
“Among the five main superfamilies of GPCRs distinguished by the GRAFS classification scheme (Schiöth and Fredriksson, 2005), two encode photosensory and chemosensory functions of vertebrates, namely the glutamate-receptor superfamily (pheromone and taste) and the rhodopsin-type superfamily (light, olfaction and possibly pheromones) (Bjarnadóttir et al., 2005;Liberles and Buck, 2006).”
Quote in the paperSubstantial claim0 of 3 sources checked- cites Schiöth and Fredriksson, 2005
Not checked in the free run.
- cites Bjarnadóttir et al., 2005
Not checked in the free run.
- cites Liberles and Buck, 2006
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationIncreasing organismal complexity has been correlated with higher rates of duplicate retention.
“Moreover, increasing organismal complexity has been correlated with higher rates of duplicate retention (Yang et al., 2003).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Yang et al., 2003
Not checked in the free run.
Probably not a substantial claim: may not pass S1 (asserted).
InterpretationreasoningSurreal-GPCRs arose by rapid duplication in the echinoid lineage.
“We hypothesize that these genes arose by rapid duplication in the echinoid lineage”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Study design and conductown dataPrimer efficiencies were found to exceed 1.9.
“Primer efficiencies (i.e., the amplification factor for each cycle) were found to exceed 1.9.”
Quote in the paperNot a substantial claimClassified as probably not a substantial claim.
PremisecitationChemoreception and light reception are mediated by GPCR superfamilies in vertebrates and invertebrates, with the main exceptions of Trp channels and cryptochrome molecules.
“Chemoreception and light reception arewith the main exception of the Trp channels and cryptochrome moleculesmediated by members of the G-protein-coupled receptor superfamilies in vertebrates and invertebrates (Ache and Young, 2005).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Ache and Young, 2005
Not checked in the free run.
Classified as probably not a substantial claim. The quote only partly states this claim.
Study design and conductown dataOnly proteins containing full, non-overlapping rhodopsin-type GPCR domains with an e-value cut-off of 0.001 were included in the final list of 979 peptides.
“Candidates were compared against the full PFAM database (Finn et al., 2006), and only proteins containing full, non-overlapping domains (e-value cut-off: 0.001) were included in the final list of 979 peptides.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Finn et al., 2006
Not checked in the free run.
May combine two claims.
PremisecitationSeveral distinct Opsin families were present at the base of Bilateria: rhabdomeric and ciliary Opsins, as well as RGR Opsins, with only one clear protostome member so far.
“In contrast to the uncertain evolutionary origin of the olfactory receptors, it is clear that several distinct Opsin families were present at the base of Bilateria, namely the rhabdomeric and the ciliary Opsins (Arendt et al., 2004), as well as the RGR Opsins, with only one clear protostome member so far (Hara-Nishimura et al., 1990;Terakita, 2005).”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites Arendt et al., 2004
Not checked in the free run.
- cites Hara-Nishimura et al., 1990
Not checked in the free run.
- cites Terakita, 2005
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationPhotosensitivity is a common phenomenon in sea urchins.
“These reports suggest that photosensitivity is a common phenomenon in sea urchins.”
Quote in the paperNot a substantial claim0 of 3 sources checked- cites Millott, 1975
Not checked in the free run.
- cites Smith, 1965
Not checked in the free run.
- cites Barnes and Cook, 2001
Not checked in the free run.
May combine two claims.
Interpretationown dataThese data suggest that the four main surreal-GPCR groups have arisen independently and from different members of the ancestral bilaterian GPCR repertoire.
“These data suggest that the four main surreal-GPCR groups have arisen independently and from different members of the ancestral bilaterian GPCR repertoire.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
PremisecitationCryptochromes originate from photolyases and typically respond to UV light.
“Cryptochromes are molecules derived from photolyases, and usually light sensitive in the UV range (Gehring and Rosbash, 2003).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Gehring and Rosbash, 2003 source
Not checked in the free run.
May combine two claims.
InterpretationreasoningThe ancestral molecules from which the different surreal-GPCRs arose are difficult to determine, resulting in distinct GPCR variants with little similarity to any existing classification scheme.
“However, similar to the olfactory receptors of vertebrates, the ancestral molecules from which the different surreal-GPCRs arose are difficult to determine, resulting in distinct GPCR variants with little similarity to any existing classification scheme.”
Quote in the paperNot a substantial claimClassified as probably not a substantial claim.
InterpretationreasoningThe abundance of rhodopsin-type GPCRs in a given genome might be a good indicator of the overall versatility of cell types and functions within a species.
“Due to this versatility, the abundance of rhodopsin-type GPCRs in a given genome might be a good indicator of the overall versatility of cell types and functions within a species.”
Quote in the paperSubstantial claimClassified as probably not a substantial claim.
PremisecitationThe rhodopsin-type GPCR superfamily is a highly versatile substrate for evolution, adapted to functions as diverse as light reception, cell-cell signaling, and chemoreception.
“The rhodopsin-type GPCR superfamily appears to be a highly versatile substrate for evolution, as it has been adapted to functions as diverse as light reception, cell-cell signaling and chemoreception (Bockaert and Pin, 1999).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Bockaert and Pin, 1999
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
InterpretationreasoningThe absolute number of rhodopsin-type GPCRs in the sea urchin genome reported in this study (979) might still change upon completion of the genome.
“The absolute number of rhodopsin-type GPCRs in the sea urchin genome reported in this study (979) might still change upon completion of the genome”
Quote in the paperNot a substantial claimMay combine two claims.
PremisecitationTube feet and different pedicellariae classes have long been proposed or, in the case of globiferous pedicellariae, demonstrated to contain chemosensory neurons.
“that have long been proposed or, in the case of globiferous pedicellariae, even demonstrated to contain chemosensory neurons.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Campbell, 1976
Not checked in the free run.
May combine two claims.
PremisecitationRetinochrome (RGR Opsins) is a group of Opsins found in vertebrates and squid, and possibly (with low branch point support) in the tunicate Ciona intestinalis.
“a group of Opsins that has been found in vertebrates and squid, and possibly (with low branch point support) in the tunicate Ciona intestinalis (Hara-Nishimura et al., 1990;Nakashima et al., 2003;Terakita, 2005).”
Quote in the paperSubstantial claim0 of 3 sources checked- cites Hara-Nishimura et al., 1990
Not checked in the free run.
- cites Nakashima et al., 2003
Not checked in the free run.
- cites Terakita, 2005
Not checked in the free run.
May combine two claims.
InterpretationcitationThe limited expression in the transcriptome map probably reflects that these genes are expressed only at late larval stages, whereas the tiling expression data were generated from four embryonic stages (egg to prism, 65 hpf).
“This probably reflects the fact that these genes are expressed only at late larval stages, whereas the tiling expression data were generated based on poly-A RNA purified from four embryonic stages (from egg to prism, 65 hpf) (Samanta et al., 2006).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Samanta et al., 2006
Not checked in the free run.
May combine two claims.
PremisecitationIn mouse olfactory GPCRs, transmembrane domains 3-5 contain most of the variable residues, with TM IV and TMV being most divergent.
“In mouse, the transmembrane domains 3-5 contain most of the variable residues, with TM IV and TMV being most divergent (Buck and Axel, 1991;Pilpel and Lancet, 1999)”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Buck and Axel, 1991
Not checked in the free run.
- cites Pilpel and Lancet, 1999
Not checked in the free run.
May combine two claims.
InterpretationreasoningThe finding of no retinochrome in sea urchin, together with the observed loss of Go-type Opsins in all sequenced vertebrates, insects, and nematodes, emphasizes that at least the molecular set-up of the ancient bilaterian light sensory system was highly complex and that gene loss has frequently occurred during the evolution of these systems in Bilateria.
“This finding, together with the observed loss of G o -type Opsins in all sequenced vertebrates, insects and nematodes emphasizes that at least the molecular set-up of the ancient bilaterian light sensory system was highly complex and that gene loss has frequently occurred during the evolution of the these systems in Bilateria.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
PremisecitationGlobiferous pedicellariae have been described to bear chemoreceptor cells on their sensory hillock.
“given the fact that these pedicellariae have been described to bear chemoreceptor cells on their sensory hillock (Campbell, 1976).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Campbell, 1976
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
PremisecitationVertebrate olfactory receptors are characteristically single-exon genes that locate to large genomic clusters.
“Notably, the vertebrate olfactory receptors are also characteristically single-exon genes that locate to large genomic clusters (reviewed in Dryer, 2000).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Dryer, 2000
Not checked in the free run.
Probably not a substantial claim: may not pass S2 (checkable).
InterpretationreasoningTargeting specific sensory cells through their molecular repertoire should aid dissection of sensory output circuits in the animal.
“the possibility to address specific sensory cells by virtue of their molecular repertoire will help to dissect sensoryoutput circuits in the animal.”
Quote in the paperNot a substantial claimMay combine two claims.
InterpretationreasoningDespite apparently lacking central sensory organs, the sea urchin has a diverse set of sensory molecules and structures.
“In summary, it appears as if the sea urchin, despite its apparent lack of central sensory organs, possesses a rich repertoire of sensory molecules and sensory structures”
Quote in the paperSubstantial claim- Rests on an assumption: The sea urchin lacks central sensory organs.
The paper relies on this without support.
The molecular basis of chemoreception and photosensation in sea urchins is not yet understood.
“Despite the rich amount of stimuli-dependent behaviors that have been described, and some basic knowledge of putative sensory structures in sea urchins that has been gathered, neither chemo-nor photosensation of the animal are understood at the molecular level.”
The paper relies on this without support.
The sea urchin pluteus larva is a primary ciliated larva.
“Because the sea urchin pluteus larva represents such a primary ciliated larva”
The paper relies on this without support.
The sea urchin pluteus larva has not been described to possess eyes.
“but has not been described to possess eyes”
The paper relies on this without support.
Dedicated complex photosensory organs have not yet been described in the sea urchin.
“As mentioned, dedicated complex photosensory organs have not yet been described in the sea urchin.”
The paper relies on this without support.
Specific rapid and large expansions of transmembrane domain genes are common to molecules that animals use to detect a large variety of divergent molecules, as in the immune system and chemoreception.
“Specific rapid and large expansions of transmembrane domain genes are common to molecules that are utilized by the animal to detect a large variety of divergent molecules, as it is the case in the immune system and in chemoreception.”
The paper relies on this without support.
The sea urchin lacks central sensory organs.
“despite its apparent lack of central sensory organs”
The paper relies on this without support.
Understanding of the molecular machinery behind sea urchin sensory responses remains very limited.
“but our understanding of the underlying molecular machineries is still very limited.”
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
These GPCRs originated by lineage-specific expansion, and many are single-exon genes forming clusters in the genome, reminiscent of lineage-specific putative chemosensory GPCRs in other animal groups.
“These GPCRs originated by lineage-specific expansion, and many are single-exon genes forming clusters in the genome [1], reminiscent of lineage-specific putative chemosensory GPCRs in other animal groups [43].”
From Origin and early evolution of neural circuits for the control of ciliary locomotion (2011) · cites this paper as [43]
Self-check
Percentages, sums and reported differences re-computed from the paper’s own numbers.
Each contradiction between two passages listed here was confirmed by an independent check.
QPCR results are assigned to a sequence-variability figure
MinorThe methods identify Fig. 4A as showing QPCR results, but the Fig. 4 caption identifies panel A as an analysis of amino-acid variability. The QPCR expression results are identified in the Fig. 5 caption.
“QPCR results reported in Figs.2A and 4A are presented as percent of the maximum value.”
“Figure: 4 Fig. 4 . Fig. 4. Variability of amino acid residues in group B and group C surreal-GPCRs.(A) Diversity of group B surreal-GPCRs; reference: SPU_013027 (surreal-GPCR-B27).”
Claim of G o-Opsin absence conflicts with Amphioxus sequences
MinorThe discussion says G o-Opsins appear absent from other deuterostome groups, but the results place two Amphioxus opsins in the G o-Opsin group. Amphioxus is a cephalochordate and therefore another deuterostome.
“Interestingly, G o -Opsins appear to be absent from the other deuterostome groups, including vertebrates, contrasting with the general trend that vertebrates have retained many ancestral proteins (reviewed in Raible and Arendt, 2004).”
“Sp-Opsin3.1 and Sp-Opsin3.2,form one well-supported group with the Patinopecten G o -Opsin (Kojima et al., 1997) and two Opsins from Amphioxus (Koyanagi et al., 2002) (Fig. 1A), thus supporting the existence of an ancestral bilaterian G o -Opsin family.”
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- Authors
- Raible, F., Tessmar-Raible, K., Arboleda, E., Kaller, T., Bork, P., Arendt, D., & Arnone, M. I.
- Affiliations
- European Molecular Biology Laboratory (DE)
- Stazione Zoologica Anton Dohrn (IT)
- Corresponding author
- Raible, Florian
- Arnone, Maria I.
- Venue
- Developmental Biology
- Publisher
- Elsevier BV
- Published
- 2006-12
- Volume
- 300
- Issue
- 1
- Pages
- 461-475
- ISSN
- 0012-1606 (ISSN-L)
- OpenAlex
- W2015311975
- Semantic Scholar
- 43763491a203173b16bacd905876f685c3259875
- Cited by
- 170
- References
- 87
- Citations by year
- 2026: 2 · 2025: 3 · 2024: 4 · 2023: 10 · 2022: 5 · 2021: 12 · 2020: 8 · 2019: 6 · 2018: 8 · 2017: 9 · 2016: 8 · 2015: 6 · 2014: 11 · 2013: 8 · 2012: 4
- FWCI
- 6.79
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Licence
- Locations
- Developmental Biology (publishedVersion)
- PubMed (publishedVersion)
- Funders
- Deutsche Forschungsgemeinschaft
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Receptor Mechanisms and Signaling
- Life Sciences › Neuroscience › Sensory Systems › Olfactory and Sensory Function Studies
- Physical Sciences › Earth and Planetary Sciences › Paleontology › Marine Invertebrate Physiology and Ecology
- Keywords
- sea urchin genome, opsin, genome clustering, olfactory receptors, chemosensory receptors, pedicellariae, tube feet, sensory neurons, echinoderm evolution, gene duplication
- MeSH terms
- Amino Acid Sequence, Animals, Models, Molecular, Molecular Sequence Data, Protein Conformation, Receptors, Cell Surface, RNA, Sea Urchins, Polymerase Chain Reaction, Consensus Sequence, Sequence Alignment, Genome, Rod Opsins, Sequence Homology, Amino Acid, In Situ Hybridization, Receptors, G-Protein-Coupled
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