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NMR-Spectroscopic Screening of Spider Venom Reveals Sulfated Nucleosides As Major Components for the Brown Recluse and Related Species

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NMR-Spectroscopic Screening of Spider Venom Reveals Sulfated Nucleosides As Major Components for the Brown Recluse and Related Species NMR-spectroscopic screening of spider venom reveals sulfated nucleosides as major components for the brown recluse and related species Frank C. Schroeder*†‡, Andrew E. Taggi†§, Matthew Gronquist¶, Rabia U. Malik†, Jacqualine B. Grantʈ, Thomas Eisner**, and Jerrold Meinwald†‡ *Boyce Thompson Institute, Cornell University, Tower Road, Ithaca, NY 14853; Departments of †Chemistry and Chemical Biology and **Neurobiology and Behavior, Cornell University, Ithaca, NY 14853; ¶Department of Chemistry, SUNY Fredonia, Fredonia, NY 14063; and ʈSchool of Forest Resources and Environmental Science, Michigan Technological University, Houghton, MI 49931 Contributed by Jerrold Meinwald, July 17, 2008 (sent for review May 12, 2008) Extensive chemical analyses of spider venoms from many species aimed at enrichment or isolation of the compounds of interest. have revealed complex mixtures of biologically active compounds, In the case of the hobo spider, Tegenaria agrestis, direct NMR of which several have provided important leads for drug devel- spectroscopic analysis facilitated detection and characterization opment. We have recently shown that NMR spectroscopy can be of an entire family of natural products, the sulfated nucleosides, used advantageously for a direct structural characterization of the including the sulfated guanosine derivatives 1 and 2 (Fig. 1) (5). small-molecule content of such complex mixtures. Here, we report Except for a single example (6), sulfated nucleosides had com- the application of this strategy to a larger-scale analysis of a pletely escaped detection by conventional isolation and charac- collection of spider venoms representing >70 species, which, in terization protocols, although spider venoms had been subject to combination with mass spectrometric analyses, allowed the iden- intensive chemical scrutiny that had led to identification of tification of a wide range of known, and several previously hundreds of proteins, peptides, acylated polyamines, and various undescribed, small molecules. These include polyamines, common small-molecule neurotransmitters (7, 8). neurotransmitters, and amino acid derivatives as well as two The promising results with T. agrestis prompted us to initiate ECOLOGY additional members of a recently discovered family of natural NMR-spectroscopic screening of venom from a large selection products, the sulfated nucleosides. In the case of the well studied of spider species, both to determine the extent to which sulfated brown recluse spider, Loxosceles reclusa, sulfated guanosine de- nucleosides might occur among the various spider taxa and to rivatives were found to comprise the major small-molecule com- obtain an expanded and unbiased view of spider venom com- ponents of the venom. position in general. chemical prospecting ͉ Loxosceles ͉ metabolomics ͉ Results CHEMISTRY natural products ͉ neurotoxins Spider venom samples representing Ͼ70 species from 27 families (9, 10) were subjected to direct NMR spectroscopic analysis. For iological samples commonly contain complex mixtures of each species, one or more lyophilized samples corresponding to Bsmall molecules with diverse properties and functions. Their 5–10 ␮l of crude venom were dissolved, without purification,†† 1 1 characterization remains a challenge, because many analytical in D2O, and H NMR spectra were recorded. These H spectra approaches are primarily aimed at identifying specific classes of were used to assess the overall composition of nonvolatiles in known compounds. Several recent studies have demonstrated each venom sample. In a number of instances, the 1H NMR the feasibility of NMR-spectroscopic analyses of complex small- spectrum immediately suggested that simple amino acids and molecule mixtures, including the use of diffusion-ordered spec- amino acid derivatives typically found in spider venom repre- troscopy (DOSY) (1) or principal component analysis (PCA) in sented the major components. For example, in the 1H spectrum metabolomics (2) as well as the characterization of crude of Pisaura mirabilis venom, signals representing Ͼ90% of the unfractionated natural products extracts using routine 2D NMR small-molecule content of the venom could be assigned to spectra (3–5). Compared with MS analyses, 2D NMR spectro- glutamic acid, aspartic acid, 4-aminobutyric acid, and spermi- scopic investigations of small-molecule mixtures offer the benefit dine (Fig. 2). For venoms of this type, structural assignments of more detailed structural information, which is of particular were confirmed through NMR-spectroscopic mixing experi- relevance for the detection of unanticipated chemotypes. We ments using authentic samples of the identified compounds. The have recently reported on the utility of direct NMR spectro- scopic analysis of unpurified biological extracts as a powerful method for the discovery of natural products (5). Such analyses Author contributions: F.C.S., A.E.T., M.G., T.E., and J.M. designed research; F.C.S., A.E.T., provide a means for the rapid evaluation of biological samples in M.G., R.U.M., and J.B.G. performed research; F.C.S., A.E.T., M.G., R.U.M., and J.B.G. analyzed an unbiased and nondestructive manner and are particularly data; and F.C.S., M.G., and J.M. wrote the paper. useful for the characterization of glandular secretions with The authors declare no conflict of interest. specific biological functions. A standard set of spectra [1H, ‡To whom correspondence may be addressed. E-mail: [email protected] or fs31@ dqfCOSY, heteronuclear multiple quantum correlation cornell.edu. (HMQC), heteronuclear multiple bond correlation (HMBC), §Present address: DuPont Crop Protection, Stine–Haskell Research Center, Newark, DE and NOESY] collected for a crude biological sample will usually 19711. permit partial or complete structural characterization of its ††In some instances, centrifugation was necessary to remove insoluble material. In addition, major components as well as of many minor components. If these it cannot be excluded that in some cases lyophilization caused loss of volatile venom components; however, for the present study, limited availability of nonlyophilized ‘‘direct’’ NMR-spectroscopic analyses suggest the presence of venom samples precluded additional analyses aimed at volatile components, for exam- compounds of interest, final identification of individual compo- ple, via GC-MS. nents can be accomplished in a subsequent step, for example by This article contains supporting information online at www.pnas.org/cgi/content/full/ using additional MS analyses, through synthesis of proposed 0806840105/DCSupplemental. structures, or via NMR-monitored fractionation of the sample © 2008 by The National Academy of Sciences of the USA www.pnas.org͞cgi͞doi͞10.1073͞pnas.0806840105 PNAS ͉ September 23, 2008 ͉ vol. 105 ͉ no. 38 ͉ 14283–14287 Downloaded by guest on September 23, 2021 O Like many of the venoms investigated, C. pastoralis venom was N O NH found to contain significant amounts of acylated polyamines, a N large family of natural products that has been extensively studied NH N N NH HO SO 2 (7). The presence of acyl polyamines in our unpurified venoms 3 O N N NH HO SO 2 was easily recognized based on the characteristic spin systems of 3 O 2 O OH their aromatic head groups, whose connections to the polyamine 1 chains were evident from the HMBC spectra. Polyamines fre- O OSO H 3 O H3C OH quently occurred as mixtures of several structurally very similar O compounds, and as a result, overlap of the corresponding H3C OH O HO NMR-spectroscopic signals in both 1D and 2D spectra did not O O HO H3C OH always permit complete characterization of individual com- OH HO pounds. Nonetheless, NMR-spectroscopic analyses enabled elu- cidation of the general structural features of the polyamines Fig. 1. Sulfated nucleosides from the hobo spider, Tegenaria agrestis. present in a sample, which, in cases where the spectra suggested the presence of unusual structural features, were investigated further. In the case of C. pastoralis, two major as well as several 1 H NMR spectra of several other spider venom samples, for minor acylated polyamines were detected in the 1H NMR example, Dysdera sp., Phyxioshema sp., and Scytodes fusca, spectrum. Analysis of the 2D NMR spectra yielded partial showed primarily broad, unresolved peaks characteristic of large structures for the two major polyamines (Tables S2 and S3). Both peptides and proteins but were virtually devoid of any significant compounds appeared to possess a 2,5-dihydroxybenzoylamin- signals representing small molecules. A summary of the small opropylamine head group and terminate with a 4-guanidinobu- molecules we identified in our venom library is given in Table 1. tylamine (Agmatine) moiety. Complete characterization of the Venom samples of more interesting composition that could intervening diaminoalkane segments was not possible, however, not be characterized based solely on their 1H NMR spectra were because of multiple signal overlap. Subsequent analysis of the analyzed further by using 2D NMR spectroscopy, including sample by ESI-MS-MS nevertheless allowed the remainder of dqfCOSY, NOESY, HMQC, and HMBC spectra. For the venom both structures, 5 and 6, to be determined (Figs. S2 and S3). For samples analyzed in this way, the resulting sets of spectra these analyses, the molecular [M ϩ 1]ϩ peaks corresponding to provided sufficient signal dispersion and connectivity informa- polyamines 5 and 6 were identified based on the structural tion to assign all major signals to discrete
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