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Bernard M. Degnan - One of the best experts on this subject based on the ideXlab platform.
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A Post-Synaptic Scaffold at the Origin of the Animal
2020Co-Authors: Sakarya, Bernard M. Degnan, Maja Adamska, Marcin Adamski, Kathryn A. Armstrong, I-fan Wang, Bruce Tidor, H Todd, Kenneth S. KosikAbstract:Background. The evolution of complex sub-cellular structures such as the synapse requires the assembly of multiple proteins, each conferring added functionality to the integrated structure. Tracking the early evolution of synapses has not been possible without genomic information from the earliest branching animals. As the closest extant relatives to the Eumetazoa, Porifera (sponges) represent a pivotal group for understanding the evolution of nervous systems, because sponges lack neurons with clearly recognizable synapses, in contrast to eumetazoan animals. Methodology/Principal Findings. We show that the genome of the demosponge Amphimedon queenslandica possesses a nearly complete set of post-synaptic protein homologs whose conserved interaction motifs suggest assembly into a complex structure. In the critical synaptic scaffold gene, dlg, residues that make hydrogen bonds and van der Waals interactions with the PDZ ligand are 100% conserved between sponge and human, as is the motif organization of the scaffolds. Expression in Amphimedon of multiple post-synaptic gene homologs in larval flask cells further supports the existence of an assembled structure. Among the few post-synaptic genes absent from Amphimedon, but present in Eumetazoa, are receptor genes including the entire ionotropic glutamate receptor family. Conclusions/Significance. Highly conserved protein interaction motifs and co-expression in sponges of multiple proteins whose homologs interact in eumetazoan synapses indicate that a complex protein scaffold was present at the origin of animals, perhaps predating nervous systems. A relatively small number of crucial innovations to this pre-existing structure may represent the founding changes that led to a post-synaptic element.
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co expression of synaptic genes in the sponge Amphimedon queenslandica uncovers ancient neural submodules
Scientific Reports, 2019Co-Authors: Eunice Wong, Sandie M Degnan, Jan Molter, Victor Anggono, Bernard M. DegnanAbstract:The synapse is a complex cellular module crucial to the functioning of neurons. It evolved largely through the exaptation of pre-existing smaller submodules, each of which are comprised of ancient sets of proteins that are conserved in modern animals and other eukaryotes. Although these ancient submodules themselves have non-neural roles, it has been hypothesized that they may mediate environmental sensing behaviors in aneural animals, such as sponges. Here we identify orthologues in the sponge Amphimedon queenslandica of genes encoding synaptic submodules in neural animals, and analyse their cell-type specific and developmental expression to determine their potential to be co-regulated. We find that genes comprising certain synaptic submodules, including those involved in vesicle trafficking, calcium-regulation and scaffolding of postsynaptic receptor clusters, are co-expressed in adult choanocytes and during metamorphosis. Although these submodules may contribute to sensory roles in this cell type and this life cycle stage, total synaptic gene co-expression profiles do not support the existence of a functional synapse in A. queenslandica. The lack of evidence for the co-regulation of genes necessary for pre- and post-synaptic functioning in A. queenslandica suggests that sponges, and perhaps the last common ancestor of sponges and other extant animals, had the ability to promulgate sensory inputs without complete synapse-like functionalities. The differential co-expression of multiple synaptic submodule genes in sponge choanocytes, which have sensory and feeding roles, however, is consistent with the metazoan ancestor minimally being able to undergo exo- and endocytosis in a controlled and localized manner.
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Sponge Long Non-Coding RNAs Are Expressed in Specific Cell Types and Conserved Networks
Non-Coding RNA, 2018Co-Authors: Federico Gaiti, William L. Hatleberg, Miloš Tanurdžić, Bernard M. DegnanAbstract:Although developmental regulation by long non-coding RNAs (lncRNAs) appears to be a widespread feature amongst animals, the origin and level of evolutionary conservation of this mode of regulation remain unclear. We have previously demonstrated that the sponge Amphimedon queenslandica—a morphologically-simple animal—developmentally expresses an array of lncRNAs in manner akin to more complex bilaterians (insects + vertebrates). Here, we first show that Amphimedon lncRNAs are expressed in specific cell types in larvae, juveniles and adults. Thus, as in bilaterians, sponge developmental regulation involves the dynamic, cell type- and context-specific regulation of specific lncRNAs. Second, by comparing gene co-expression networks between Amphimedon queenslandica and Sycon ciliatum—a distantly-related calcisponge—we identify several putative co-expression modules that appear to be shared in sponges; these network-embedded sponge lncRNAs have no discernable sequence similarity. Together, these results suggest sponge lncRNAs are developmentally regulated and operate in conserved gene regulatory networks, as appears to be the case in more complex bilaterians.
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lipidomics of the sea sponge Amphimedon queenslandica and implication for biomarker geochemistry
Geobiology, 2017Co-Authors: David A Gold, Bernard M. Degnan, Sandie M Degnan, Shane S Oreilly, Jabin R Watson, Jens O Kromer, Roger E SummonsAbstract:Demosponges are a rich natural source of unusual lipids, some of which are of interest as geochemical biomarkers. Although demosponges are animals, they often host dense communities of microbial symbionts, and it is therefore unclear which lipids can be synthesized by the animal de novo, and which require input from the microbial community. To address this uncertainty, we analyzed the lipids of Amphimdeon queenslandica, the only demosponge with a published genome. We correlated the genetic and lipid repertoires of A. queenslandica to identify which biomarkers could potentially be synthesized and/or modified by the sponge. The fatty acid profile of A. queenslandica is dominated by an unusual Δ^(5,9) fatty acid (cis-5,9-hexacosadienoic acid)—similar to what has been found in other members of the Amphimdeon genus—while the sterol profile is dominated by C_(27)-C_(29( derivatives of cholesterol. Based on our analysis of the A. queenslandica genome, we predict that this sponge can synthesize sterols de novo, but it lacks critical genes necessary to synthesize basic saturated and unsaturated fatty acids. However, it does appear to have the genes necessary to modify simpler products into a more complex “algal-like” assemblage of unsaturated fatty acids. Ultimately, our results provide additional support for the poriferan affinity of 24-isopropylcholestanes in Neoproterozoic-age rocks (the “sponge biomarker” hypothesis) and suggest that some algal proxies in the geochemical record could also have animal contributions.
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transcriptomic profiling of the allorecognition response to grafting in the demosponge Amphimedon queenslandica
Marine Drugs, 2017Co-Authors: Laura F. Grice, Bernard M. DegnanAbstract:Sponges, despite their simple body plan, discriminate between self and nonself with remarkable specificity. Sponge grafting experiments simulate the effects of natural self or nonself contact under laboratory conditions. Here we take a transcriptomic approach to investigate the temporal response to self and nonself grafts in the marine demosponge Amphimedon queenslandica. Auto- and allografts were established, observed and sampled over a period of three days, over which time the grafts either rejected or accepted, depending on the identity of the paired individuals, in a replicable and predictable manner. Fourteen transcriptomes were generated that spanned the auto- and allograft responses. Self grafts fuse completely in under three days, and the process appears to be controlled by relatively few genes. In contrast, nonself grafting results in a complete lack of fusion after three days, and appears to involve a broad downregulation of normal biological processes, rather than the mounting of an intense defensive response.
Junichi Kobayashi - One of the best experts on this subject based on the ideXlab platform.
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zamamidine d a manzamine alkaloid from an okinawan Amphimedon sp marine sponge
Journal of Natural Products, 2017Co-Authors: Takaaki Kubota, Jane Fromont, Tohru Gonoi, Kenta Nakamura, Shinichiro Kurimoto, Kanae Sakai, Junichi KobayashiAbstract:A new manzamine alkaloid, zamamidine D (1), was isolated from an Okinawan Amphimedon sp. marine sponge. The structure of zamamidine D (1) including the relative configuration was elucidated on the basis of spectroscopic data. Zamamidine D (1) is the first manzamine alkaloid possessing a 2,2′-methylenebistryptamine unit as the aromatic moiety instead of a β-carboline unit. Zamamidine D (1) showed antimicrobial activity against several bacteria and fungi.
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zamamiphidin a a new manzamine related alkaloid from an okinawan marine sponge Amphimedon sp
Organic Letters, 2013Co-Authors: Takaaki Kubota, Jane Fromont, Yuki Kamijyo, Azusa Takahashinakaguchi, Tohru Gonoi, Junichi KobayashiAbstract:A manzamine related alkaloid, zamamiphidin A (1), consisting of a new heptacyclic ring system has been isolated from an Okinawan marine sponge Amphimedon sp. The structure of 1 including the relative stereochemistry was elucidated on the basis of the spectroscopic data. Compound 1 showed antibacterial activity against Staphylococcus aureus (MIC, 32 μg/mL).
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Pyrinodemins G–I, new bis-3-alkylpyridine alkaloids from a marine sponge Amphimedon sp.
Tetrahedron, 2013Co-Authors: Takaaki Kubota, Jane Fromont, Kenichi Kura, Junichi KobayashiAbstract:Three new bis-3-alkylpyridine alkaloids, pyrinodemins G–I (1–3), were isolated from an Okinawan marine sponge Amphimedon sp. and the structures of 1–3 were elucidated on the basis of spectroscopic data. Pyrinodemins G–I (1–3) were novel bis-3-alkylpyridine alkaloids possessing an N-methoxy amide moiety, an α,β-unsaturated 3,5-disubstituted γ-lactone moiety, and a 2,3-di-substituted acrolein at a junction of two 3-alkylpyridines, respectively. Pyrinodemins G (1) and H (2) showed cytotoxicity against P388 murine leukemia cells.
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pyrinodemins g i new bis 3 alkylpyridine alkaloids from a marine sponge Amphimedon sp
Tetrahedron, 2013Co-Authors: Takaaki Kubota, Jane Fromont, Kenichi Kura, Junichi KobayashiAbstract:Three new bis-3-alkylpyridine alkaloids, pyrinodemins G–I (1–3), were isolated from an Okinawan marine sponge Amphimedon sp. and the structures of 1–3 were elucidated on the basis of spectroscopic data. Pyrinodemins G–I (1–3) were novel bis-3-alkylpyridine alkaloids possessing an N-methoxy amide moiety, an α,β-unsaturated 3,5-disubstituted γ-lactone moiety, and a 2,3-di-substituted acrolein at a junction of two 3-alkylpyridines, respectively. Pyrinodemins G (1) and H (2) showed cytotoxicity against P388 murine leukemia cells.
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pyrinodemins e and f new 3 alkylpyridine alkaloids from sponge Amphimedon sp
Bioorganic & Medicinal Chemistry Letters, 2011Co-Authors: Kenichi Kura, Jane Fromont, Takaaki Kubota, Junichi KobayashiAbstract:Two new 3-alkylpyridine alkaloids, pyrinodemins E (1) and F (2), were isolated from an Okinawan marine sponge Amphimedon sp. and the structures of 1 and 2 were elucidated on the basis of spectroscopic data. Pyrinodemins E (1) and F (2) were novel 3-alkylpyridine alkaloids possessing a 4-(methoxyamino)piperidinone moiety and an indol-3-glyoxylamide moiety, respectively. Pyrinodemin E (1) showed cytotoxicity against P388 and L1210 murine leukemia cells.
Takaaki Kubota - One of the best experts on this subject based on the ideXlab platform.
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zamamidine d a manzamine alkaloid from an okinawan Amphimedon sp marine sponge
Journal of Natural Products, 2017Co-Authors: Takaaki Kubota, Jane Fromont, Tohru Gonoi, Kenta Nakamura, Shinichiro Kurimoto, Kanae Sakai, Junichi KobayashiAbstract:A new manzamine alkaloid, zamamidine D (1), was isolated from an Okinawan Amphimedon sp. marine sponge. The structure of zamamidine D (1) including the relative configuration was elucidated on the basis of spectroscopic data. Zamamidine D (1) is the first manzamine alkaloid possessing a 2,2′-methylenebistryptamine unit as the aromatic moiety instead of a β-carboline unit. Zamamidine D (1) showed antimicrobial activity against several bacteria and fungi.
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zamamiphidin a a new manzamine related alkaloid from an okinawan marine sponge Amphimedon sp
Organic Letters, 2013Co-Authors: Takaaki Kubota, Jane Fromont, Yuki Kamijyo, Azusa Takahashinakaguchi, Tohru Gonoi, Junichi KobayashiAbstract:A manzamine related alkaloid, zamamiphidin A (1), consisting of a new heptacyclic ring system has been isolated from an Okinawan marine sponge Amphimedon sp. The structure of 1 including the relative stereochemistry was elucidated on the basis of the spectroscopic data. Compound 1 showed antibacterial activity against Staphylococcus aureus (MIC, 32 μg/mL).
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pyrinodemins g i new bis 3 alkylpyridine alkaloids from a marine sponge Amphimedon sp
Tetrahedron, 2013Co-Authors: Takaaki Kubota, Jane Fromont, Kenichi Kura, Junichi KobayashiAbstract:Three new bis-3-alkylpyridine alkaloids, pyrinodemins G–I (1–3), were isolated from an Okinawan marine sponge Amphimedon sp. and the structures of 1–3 were elucidated on the basis of spectroscopic data. Pyrinodemins G–I (1–3) were novel bis-3-alkylpyridine alkaloids possessing an N-methoxy amide moiety, an α,β-unsaturated 3,5-disubstituted γ-lactone moiety, and a 2,3-di-substituted acrolein at a junction of two 3-alkylpyridines, respectively. Pyrinodemins G (1) and H (2) showed cytotoxicity against P388 murine leukemia cells.
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Pyrinodemins G–I, new bis-3-alkylpyridine alkaloids from a marine sponge Amphimedon sp.
Tetrahedron, 2013Co-Authors: Takaaki Kubota, Jane Fromont, Kenichi Kura, Junichi KobayashiAbstract:Three new bis-3-alkylpyridine alkaloids, pyrinodemins G–I (1–3), were isolated from an Okinawan marine sponge Amphimedon sp. and the structures of 1–3 were elucidated on the basis of spectroscopic data. Pyrinodemins G–I (1–3) were novel bis-3-alkylpyridine alkaloids possessing an N-methoxy amide moiety, an α,β-unsaturated 3,5-disubstituted γ-lactone moiety, and a 2,3-di-substituted acrolein at a junction of two 3-alkylpyridines, respectively. Pyrinodemins G (1) and H (2) showed cytotoxicity against P388 murine leukemia cells.
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pyrinodemins e and f new 3 alkylpyridine alkaloids from sponge Amphimedon sp
Bioorganic & Medicinal Chemistry Letters, 2011Co-Authors: Kenichi Kura, Jane Fromont, Takaaki Kubota, Junichi KobayashiAbstract:Two new 3-alkylpyridine alkaloids, pyrinodemins E (1) and F (2), were isolated from an Okinawan marine sponge Amphimedon sp. and the structures of 1 and 2 were elucidated on the basis of spectroscopic data. Pyrinodemins E (1) and F (2) were novel 3-alkylpyridine alkaloids possessing a 4-(methoxyamino)piperidinone moiety and an indol-3-glyoxylamide moiety, respectively. Pyrinodemin E (1) showed cytotoxicity against P388 and L1210 murine leukemia cells.
Jane Fromont - One of the best experts on this subject based on the ideXlab platform.
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zamamidine d a manzamine alkaloid from an okinawan Amphimedon sp marine sponge
Journal of Natural Products, 2017Co-Authors: Takaaki Kubota, Jane Fromont, Tohru Gonoi, Kenta Nakamura, Shinichiro Kurimoto, Kanae Sakai, Junichi KobayashiAbstract:A new manzamine alkaloid, zamamidine D (1), was isolated from an Okinawan Amphimedon sp. marine sponge. The structure of zamamidine D (1) including the relative configuration was elucidated on the basis of spectroscopic data. Zamamidine D (1) is the first manzamine alkaloid possessing a 2,2′-methylenebistryptamine unit as the aromatic moiety instead of a β-carboline unit. Zamamidine D (1) showed antimicrobial activity against several bacteria and fungi.
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zamamiphidin a a new manzamine related alkaloid from an okinawan marine sponge Amphimedon sp
Organic Letters, 2013Co-Authors: Takaaki Kubota, Jane Fromont, Yuki Kamijyo, Azusa Takahashinakaguchi, Tohru Gonoi, Junichi KobayashiAbstract:A manzamine related alkaloid, zamamiphidin A (1), consisting of a new heptacyclic ring system has been isolated from an Okinawan marine sponge Amphimedon sp. The structure of 1 including the relative stereochemistry was elucidated on the basis of the spectroscopic data. Compound 1 showed antibacterial activity against Staphylococcus aureus (MIC, 32 μg/mL).
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pyrinodemins g i new bis 3 alkylpyridine alkaloids from a marine sponge Amphimedon sp
Tetrahedron, 2013Co-Authors: Takaaki Kubota, Jane Fromont, Kenichi Kura, Junichi KobayashiAbstract:Three new bis-3-alkylpyridine alkaloids, pyrinodemins G–I (1–3), were isolated from an Okinawan marine sponge Amphimedon sp. and the structures of 1–3 were elucidated on the basis of spectroscopic data. Pyrinodemins G–I (1–3) were novel bis-3-alkylpyridine alkaloids possessing an N-methoxy amide moiety, an α,β-unsaturated 3,5-disubstituted γ-lactone moiety, and a 2,3-di-substituted acrolein at a junction of two 3-alkylpyridines, respectively. Pyrinodemins G (1) and H (2) showed cytotoxicity against P388 murine leukemia cells.
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Pyrinodemins G–I, new bis-3-alkylpyridine alkaloids from a marine sponge Amphimedon sp.
Tetrahedron, 2013Co-Authors: Takaaki Kubota, Jane Fromont, Kenichi Kura, Junichi KobayashiAbstract:Three new bis-3-alkylpyridine alkaloids, pyrinodemins G–I (1–3), were isolated from an Okinawan marine sponge Amphimedon sp. and the structures of 1–3 were elucidated on the basis of spectroscopic data. Pyrinodemins G–I (1–3) were novel bis-3-alkylpyridine alkaloids possessing an N-methoxy amide moiety, an α,β-unsaturated 3,5-disubstituted γ-lactone moiety, and a 2,3-di-substituted acrolein at a junction of two 3-alkylpyridines, respectively. Pyrinodemins G (1) and H (2) showed cytotoxicity against P388 murine leukemia cells.
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pyrinodemins e and f new 3 alkylpyridine alkaloids from sponge Amphimedon sp
Bioorganic & Medicinal Chemistry Letters, 2011Co-Authors: Kenichi Kura, Jane Fromont, Takaaki Kubota, Junichi KobayashiAbstract:Two new 3-alkylpyridine alkaloids, pyrinodemins E (1) and F (2), were isolated from an Okinawan marine sponge Amphimedon sp. and the structures of 1 and 2 were elucidated on the basis of spectroscopic data. Pyrinodemins E (1) and F (2) were novel 3-alkylpyridine alkaloids possessing a 4-(methoxyamino)piperidinone moiety and an indol-3-glyoxylamide moiety, respectively. Pyrinodemin E (1) showed cytotoxicity against P388 and L1210 murine leukemia cells.
Rob W. M. Van Soest - One of the best experts on this subject based on the ideXlab platform.
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Perplexing distribution of 3-alkylpyridines in haplosclerid sponges
2020Co-Authors: Leontine E. Becking, Yoichi Nakao, Rob W. M. Van Soest, Nobuhiro Fusetani, Nicole J. De Voogd, Shigeki MatsunagaAbstract:Abstract: In this study we reviewed the natural product literature for the distribution of 3-alkylpyridines among sponge taxa. In parallel, we traced selected 3-alkylpyridines, amphitoxins, in three haplosclerid genera (Amphimedon, Callyspongia, Haliclona) in order to establish the utility of such compounds as genuine chemotaxonomic markers. We confirmed that this group of compounds had been almost solely extracted from sponges of the order Haplosclerida. Three groups of compounds within the 3-alkylpyridine derivatives were noteworthy, as they appear to be concentrated in restricted taxonomic units within the order Haplosclerida: 1) polymers, 2) cyclic dimers, and 3) bicyclic dimers. There was a concentration of the polymer amphitoxin in the families Niphatidae and Callyspongiidae of the suborder Haplosclerina, and more particularly in the genera Amphimedon and Callyspongia. Our experimental results reconfirmed the presence of amphitoxin in Callyspongia (Euplacella) biru, but we were unable to trace any amphitoxins or 3-alkylpyridines of any kind in Callyspongia (Callyspongia) truncata, Haliclona sp., and Amphimedon aff. queenslandica. Assuming that these classifications are correct, our results diminish the value of both amphitoxins and 3-alkylpyridines as monophyletic markers.
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phylogenetic relationships of the marine haplosclerida phylum porifera employing ribosomal 28s rrna and mitochondrial cox1 nad1 gene sequence data
PLOS ONE, 2011Co-Authors: J Raleigh, Simon A Travers, Brian Bradshaw, Salla Vartia, Kelly M Stephens, Niamh E. Redmond, Michelle Kelly, Rob W. M. Van Soest, Grace P MccormackAbstract:The systematics of the poriferan Order Haplosclerida (Class Demospongiae) has been under scrutiny for a number of years without resolution. Molecular data suggests that the order needs revision at all taxonomic levels. Here, we provide a comprehensive view of the phylogenetic relationships of the marine Haplosclerida using many species from across the order, and three gene regions. Gene trees generated using 28S rRNA, nad1 and cox1 gene data, under maximum likelihood and Bayesian approaches, are highly congruent and suggest the presence of four clades. Clade A is comprised primarily of species of Haliclona and Callyspongia, and clade B is comprised of H. simulans and H. vansoesti (Family Chalinidae), Amphimedon queenslandica (Family Niphatidae) and Tabulocalyx (Family Phloeodictyidae), Clade C is comprised primarily of members of the Families Petrosiidae and Niphatidae, while Clade D is comprised of Aka species. The polyphletic nature of the suborders, families and genera described in other studies is also found here.
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amphimedosides 3 alkylpyridine glycosides from a marine sponge Amphimedon sp
Journal of Natural Products, 2006Co-Authors: Yoshihiko Takekawa, Rob W. M. Van Soest, Shigeki Matsunaga, Nobuhiro FusetaniAbstract:Five 3-alkylpyridine alkaloids, amphimedosides A−E (1−5), have been isolated from a marine sponge Amphimedon sp. The structures of 1−5 have been determined by analysis of NMR and FABMS data and chiral GC analyses of the acid hyrolyzates. In particular, the site of glycosylation in 1 was confirmed by the 1H−15N HMBC experiment, and the location of the double bond in 5 was assigned on the basis of tandem FABMS data. Amphimedosides are the first examples of β-d-glucosylated 3-alkylpyridine alkaloids and exhibited cytotoxic activities comparable with those of the nonglycosylated congeners.
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haplosclerida porifera demospongiae from the coast of maranhao state brazil southwestern atlantic
Zootaxa, 2005Co-Authors: Mauricio Campos, Rafael Eckert, Beatriz Mothes, Rob W. M. Van SoestAbstract:This work deals with haplosclerid sponges off the coast of Maranhao State, northeastern coast of Brazilian shelf (southwestern Atlantic). A new species is described, Haliclona (Halichoclona) lernerae. Four species are recorded for the first time for the Brazilian coast: Amphimedon caribica (Pulitzer-Finali, 1986); Niphates lutea Lehnert & Van Soest, 1999; Neopetrosia subtriangularis (Duchassaing, 1850) and Petrosia (Petrosia) weinbergi Van Soest, 1980. Three species were recollected from the southwestern Atlantic and have their geographic distribution extended on the Brazilian coast: Callyspongia (Cladochalina) vaginalis (Duchassaing & Michelotti, 1864), Oceanapia bartschi (De Laubenfels, 1934) and Xestospongia muta (Schmidt, 1870). Two species were collected for the first time from the Maranhao State Coast: Niphates erecta Duchassaing & Michelotti, 1864 and Neopetrosia proxima (Duchassaing & Michelotti, 1864). A taxonomic study of those samples is given, including description, illustrations and geographic distribution.
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amphimelibiosides a f six new ceramide dihexosides isolated from a japanese marine sponge Amphimedon sp
Journal of Organic Chemistry, 2005Co-Authors: Chisato Emura, Tomofumi Miyamoto, Ryuichi Higuchi, Rob W. M. Van SoestAbstract:Six new ceramide dihexosides, amphimelibiosides A−F (1−6), were isolated from a Japanese marine sponge Amphimedon sp. The structure of amphimelibioside C (3), which is a major component of amphimelibiosides, was determined by 2D NMR techniques, chemical degradation, and a semisynthetic method to be 1-O-[β-d-glucopyranosyl-(1→6)-α-d-galactopyranosyl]-(2S,3S,4R,6E)-2-[(2‘R)-2-hydroxydocosanoyl]-2-amino-6-octadecene-1,3,4-triol. The structures of the other constituents were elucidated by a combination of mass spectra, 1H NMR, and GC−MS analysis.
