This thread ended up meandering far enough that I think it needs a table of contents :lol:. Initially I just posted some notes about Aaron Jenks' Ph.D. dissertation that deals with the evolutionary history of S. divinorum. Then the thread got stickied so I added some notes on the pre- and early- recorded history of the plant. Then one of our members was kind enough to provide me with a wide variety of theses and dissertations dealing with S. divinorum, and this thread seems like as good a place as any to share my notes on those sources. I've added a post for each paper as I finished reading it. There are a few other things peppered in here and there.
Table of Contents
In terms of the evolutionary history of Salvia divinorum, there are a couple of particularly contentious points that have been discussed or speculated on over the years.
One is the question of why the plant seems to have such low reproductive viability. While some cultivators have been able to induce the plant to set seed, this has never been observed in the wild in its native region; there the plant seems to exclusively propagate vegetatively (whether though humans taking cuttings or tall plants breaking off at the stem and setting new roots). Some have speculated that this is because the plant is a hybrid with low reproductive viability, while others suggest that it's a result of inbreeding depression, a consequence of having been subject to human cultivation for a long period of time (particularly if the plants in cultivation had limited genetic diversity to begin with). The other issue is whether S. divinorum is monoclonal (that is, if all the plants are genetically identical). This speculation arose from the fact that the plant propagates vegetatively, and there are no truly wild populations of the species known. All populations are either under human cultivation, or appear to be feral populations that were once under human cultivation, so it would not be inconceivable that they all represent a single genotype.
In 2007, Daniel Siebert mentioned that he had a colleague at UC Riverside doing research on the plant for a PhD dissertation, including genetic analysis of S. divinorum to resolve the issue of whether the plant is monoclonal. Then just last month, the Journal of Plant Research published a paper by Aaron Jenks discussing the evolution and origins of S. divinorum. The paper did resolve the question of hybridity: long story short, S. divinorum is not a hybrid species. If you're really into botany, the paper also established that S. divinorum does not belong in section Dusenostachys of subgenus Calosphace, where Epling had originally placed it, but it does clearly belong within Calosphace. Aside from those two points, the paper left me with more unanswered questions than I'd have liked.
However, in reading the paper I noticed that Jenks was affiliated with UC Riverside, and that the paper was originally part of a PhD dissertation. So I hopped over to UC Riverside's archive of dissertations, and sure enough he'd submitted the dissertation in 2009, and the full text of it was freely available. And luckily, the dissertation contained a lot more juicy bits of information than the recently-published paper. I guess this information has been out there for a while, I just have never seen any mention of it (in the published literature or elsewhere). I've summarized the more interesting bits:
Further Reading
Table of Contents
- Post #1: Notes on Jenks' "Systematics and ethnobotany of Salvia subgenus Calosphace and origins of the hallucinogenic sage, Salvia divinorum"
- Post #8: Notes on the prehistory and early recorded history of S. divinorum
- Post #9: Notes on Willard's "Forensic analysis of Salvia divinorum and related Salvia species using chemometric procedures"
- Post #14: Notes on Addy's "That deep internal voice: controlled administration of Salvia divinorum"
- Post #15: Notes on Gruber's "Quantification of salvinorin A from tissues of Salvia divinorum (Epling & Játiva-M.)"
- Post #16: 1994 Palenque Norte talk by Rob Montgomery with Jonathan Ott and Terence McKenna
- Post #18: Guide to the literature on models for salvinorin A binding to opioid receptors
- Post #19: Notes on Kane's "Molecular recognition of G-protein coupled receptor ligands: insights into salvinorin A and xanomeline"
- Post #20: Notes on Yan's "Molecular mechanisms by which salvinorin A binds to and activates the kappa-opioid receptor"
- Post #24: Notes on Kutrzeba's "Biosynthesis of salvinorin A, a potent hallucinogen from Salvia divinorum Epling & Jativa"
- Post #27: Notes on Lawrence's "Total synthesis of salvinorin A"
- Post #28: Notes on Lovell's "Synthesis and pharmacological evaluation of salvinorin A analogues as opioid receptor probes"
- Post #30: Notes on Lozama's "Preparation of neoclerodane diterpenes as probes for the opioid receptor system”
- Post #31: Notes on McGovern's "Salvinorin A: fragment synthesis and modeling studies"
- Post #32: Notes on Nygård's "Listening to the sage: the experience of learning from the Salvia divinorum altered state"
- Post #33: Notes on Pandit's "3D-QSAR studies of salvinorin A analogs as kappa opioid agonists"
- Post #34: Notes on Phipps' "The neuropharmacological assesment [sic] of Salvia divinorum Epling and Jativa-M"
- Post #35: Notes on Stevens' "Extraction and photodegradation of the plant species Salvia divinorum"
- Post #36: Notes on Stewart's "Phytochemical investigation of Salvia divinorum"
- Post #37: Notes on Tidgewell's "Development of novel analgesics from the neoclerodane diterpene natural product salvinorin A"
- Post #38: Notes on Valdés' "The pharmacognosy of Salvia divinorum (Epling & Jativa-M)"
- Post #39: Notes on Rasakham's "Kappa opioid receptor regulation of ERK1/2 MAP kinase signaling cascade: molecular mechanisms modulating cocaine reward"
- Post #40: Relevant patents and patent applications
In terms of the evolutionary history of Salvia divinorum, there are a couple of particularly contentious points that have been discussed or speculated on over the years.
One is the question of why the plant seems to have such low reproductive viability. While some cultivators have been able to induce the plant to set seed, this has never been observed in the wild in its native region; there the plant seems to exclusively propagate vegetatively (whether though humans taking cuttings or tall plants breaking off at the stem and setting new roots). Some have speculated that this is because the plant is a hybrid with low reproductive viability, while others suggest that it's a result of inbreeding depression, a consequence of having been subject to human cultivation for a long period of time (particularly if the plants in cultivation had limited genetic diversity to begin with). The other issue is whether S. divinorum is monoclonal (that is, if all the plants are genetically identical). This speculation arose from the fact that the plant propagates vegetatively, and there are no truly wild populations of the species known. All populations are either under human cultivation, or appear to be feral populations that were once under human cultivation, so it would not be inconceivable that they all represent a single genotype.
In 2007, Daniel Siebert mentioned that he had a colleague at UC Riverside doing research on the plant for a PhD dissertation, including genetic analysis of S. divinorum to resolve the issue of whether the plant is monoclonal. Then just last month, the Journal of Plant Research published a paper by Aaron Jenks discussing the evolution and origins of S. divinorum. The paper did resolve the question of hybridity: long story short, S. divinorum is not a hybrid species. If you're really into botany, the paper also established that S. divinorum does not belong in section Dusenostachys of subgenus Calosphace, where Epling had originally placed it, but it does clearly belong within Calosphace. Aside from those two points, the paper left me with more unanswered questions than I'd have liked.
However, in reading the paper I noticed that Jenks was affiliated with UC Riverside, and that the paper was originally part of a PhD dissertation. So I hopped over to UC Riverside's archive of dissertations, and sure enough he'd submitted the dissertation in 2009, and the full text of it was freely available. And luckily, the dissertation contained a lot more juicy bits of information than the recently-published paper. I guess this information has been out there for a while, I just have never seen any mention of it (in the published literature or elsewhere). I've summarized the more interesting bits:
- As previously mentioned, S. divinorum does not appear to be a hybrid based on the low incidence of polymorphisms in genetic sequences that are inherited biparentally.
- In the sequences analyzed to assess hybridity, two of the six plants analyzed each displayed a single polymorphism (both at the same locus). This means that there is at least some genetic variability in the species, so it can safely be said that it's not monoclonal. It's still possible that genetic variations are extremely limited; more thorough examination of a larger number of plants collected from wild populations would be necessary to assess the species' genetic diversity.
- S. divinorum is closely related to a Columbian species, S. venulosa. The data indicates that Central America was the primary region of diversification of the subgenus Calosphace, with dispersal events carrying some species to South America. The simplest explanation would be that the common ancestor of these two species arose in Mexico, and a dispersal event carried the ancestor of S. venulosa to the Northern Andes. But it can't be ruled out that the common ancestor of both species arose in South America (after being carried there by a dispersal event), and the ancestor of modern S. divinorum was returned to Mexico in a back-dispersal event. To resolve this question, phylogenetic studies would have to be conducted on a variety of related South American species.
- It's not certain whether S. divinorum once occurred over a wider geographic range, but if this were the case it would help explain the plant's apparently limited reproductive viability in its current natural habitat. Nahua trade routes pssed though the eastern Mazatec region, possible providing an explanation of S. divinorum's presence there (assuming it previously occurred over a wider geographic range). “Regardless, of whether or not S. divinorum is native to the Sierra Mazateca, the conditions there do not seem fully adequate for sexual reproduction, suggesting that the origins of this plant are elsewhere.”
- Jenks frequently mentions the possibility pipiltzintzintli being S. divinorum as possible evidence for a more broad distribution in the past... I find the evidence for that identification problematic, as detailed in my Notes on Pipiltzintzintli. But the fact that a deity's headdress appears to be decorated with the flower of S. divinorum in the Dresden Codex of the pre-Columbian Maya would certainly tend to support the notion that S. divinorum occurred outside the Mazatec region some centuries ago.
- The close association of the plant with humans does not, in itself, mean that the species is a cultigen (requiring human intervention to prosper). While S. divinorum is not reproductively robust (about 50% of pollen grains aborted), it does seem to be reproductively viable. Nutlet maturation in self-pollinated flowers has been reported to be fairly low (11 nutlets from 108 flowers), seed set in some species of Calosphace appears to be naturally low. The reported 30% germination rate is not abnormally low for species in this subgenus.
Further Reading
- Jenks, A.A. 2009. "Systematics and ethnobotany of Salvia subgenus Calosphace and origins of the hallucinogenic sage, Salvia divinorum." PhD dissertation, Plant Biology: University of California Riverside. (fulltext pdf).
- Jenks, A.A., J.B. Walker, and S.-C. Kim. 2011. "Evolution and origins of the Mazatec hallucinogenic sage, Salvia divinorum (Lamiaceae): a molecular phylogenetic approach." Journal of Plant Research 124(5): 592-600.
- Reisfield, A.S. "The botany of Salvia divinorum (Labiatae)." SIDA 15(3): 349-366. (fulltext pdf)