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Molecular Self-Assembly: Hypothesized for “Hair” of Macroneuropteris Scheuchzeri (Pennsylvanian-Age Seed-Fern)

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Molecular Self-Assembly: Hypothesized for “Hair” of Macroneuropteris Scheuchzeri (Pennsylvanian-Age Seed-Fern) International Journal of Coal Geology 121 (2014) 14–18 Contents lists available at ScienceDirect International Journal of Coal Geology journal homepage: www.elsevier.com/locate/ijcoalgeo Molecular self-assembly: Hypothesized for “hair” of Macroneuropteris scheuchzeri (Pennsylvanian-age seed-fern) Erwin L. Zodrow Palaeobiology Laboratory, Cape Breton University, Postal Box 303, Sydney, B1P 6L2 Nova Scotia, Canada article info abstract Article history: Hoffmann (1827) erected the Carboniferous pteridophyll species Neuropteris Scheuchzeri without mentioning, Received 5 September 2013 nor illustrating, “hair” in the species' diagnosis. However, one to five millimeter-long hair-like structures on Received in revised form 6 November 2013 the abaxial pinnule of the species, called hair or trichome in the literature, have been routinely used since Accepted 7 November 2013 1847 as one of the main taxonomic character states for determining the identity of this species. Results from Available online 13 November 2013 preparatory and microscopic observations, together with infrared spectrochemistry, have clarified that these structures are not the same as trichomes for the following reasons. The hair-like structures of M. scheuchzeri Keywords: Extracuticular deposit (1) are not organically attached to the abaxial surface; (2) differ spectrochemically from the organic material Ex “hair” of the lamina; (3) are composed, in contrast with the trichomes, of relatively long, unbranched aliphatic FTIR (polymythelinic) hydrocarbon chains [CH2)]n, and (4) are acellular and black, unlike true trichomes of the species Carboniferous that are multicellular. Overall, the sum-total of these experimental results supports the postulate for dynamic M. scheuchzeri molecular self-assembly. For this reason the term “extracuticular deposit” is proposed, reflecting the origin and emergent nature of such hair-like structures in the abaxial pinnule. © 2013 Elsevier B.V. All rights reserved. 1. Introduction This paper focuses on the hypothesis that the long, hair-like struc- tures of M. scheuchzeri are really extracuticular deposits resulting from The current concept of Macroneuropteris scheuchzeri (Cleal et al., the process of dynamic molecular self-assembly (summary: Koch and 1990; Hoffmann, 1827) includes four character states on the abaxial Ensikat, 2008). A review of the taxonomy/systematics of M. scheuchzeri pinnule, i.e., extracuticular deposits (this study), hair (trichome in is accordingly recommended. botanical Greek), files (unicellular transparent structures), and papillae which are comparatively small and curved. Of these features, the trichomes very densely populate the abaxial lamina (see Barthel, 2. Materials and methods 1961; Cleal and Zodrow, 1989, and others). In contrast to these features of which extracuticular deposits are clearly observable under a loupe or The Carboniferous seed-fern M. scheuchzeri bears polymorphic even by the naked eye as they are up to 5 mm long, Hoffmann did not pinnate foliage that is seen above and below a basal frond dichotomy; mention the presence of any abaxial features in his diagnosis of the individual pinnule lengths range from 3 mm to 120 mm (summaries: species. However, the presence of extracuticular deposits, using the Cleal and Zodrow, 1989; Laveine, 1997; Laveine and Belhis, 2007; name hair, has been taken as an important taxonomic character state Zodrow, 2003, and many others). since 1847 to distinguish the identity of M. scheuchzeri from among sim- The compression specimens for this study were collected by the ilar larger-leaved Pennsylvanian foliage (literature survey: Bunbury, author from the roof shale of the Lloyd Cove Seam, which is known 1847 to Stull et al., 2012). A notable exception is Leo Lesquereux who in the literature for its rich content in plant fossils that are well- assigned American hairy, long-leaved pinnules to neuropteroid taxa, preserved (Fig. 1). For the experimental work, only compressions other than Neuropteris scheuchzeri (summarized by Darrah, 1969). freed from the rock matrix were used from which extracuticular de- However, Gothan (1916), made the pointed observation that the visible posits were collected in a Petri dish. Trichomes were obtained from a cu- hair of N. scheuchzeri did not survive Schulze's chemical-oxidative treat- ticle of a macerated compression (cf. Cleal and Zodrow, 1989). ment (cf. Cleal and Zodrow, 1989). Barthel (1961) described attached, Spectrochemical analyses of extracuticular deposits and trichomes pointed hairs of N. scheuchzeri, which instead I regard as extracuticular were performed using FTIR (Fourier transform infrared spectrometry) deposits. and the KBr-pellet technique. Interpretive details, particularly of IR (infrared) spectra of M. scheuchzeri, supported by carbon 13 nuclear mag- netic resonance studies, are found in Lyons et al. (1995),orD'Angelo et al. E-mail address: [email protected]. (2010, 2013). 0166-5162/$ – see front matter © 2013 Elsevier B.V. All rights reserved. http://dx.doi.org/10.1016/j.coal.2013.11.002 E.L. Zodrow / International Journal of Coal Geology 121 (2014) 14–18 15 Fig. 1. Location map. (A) Canada. (B) Sydney Sub-Basin as integral geological structure of the Maritimes Basin. (C) Coal lithostratigraphy and sample seam (S). CANT lower Cantabrian strata. 3. Results the midvein (Fig. 2B). The distribution pattern, as correctly drawn by Bunbury in 1847, follows a trend, i.e., it is biogenetically non-random. 3.1. Physical aspects of extracuticular deposits Extracuticular deposits are straight in shape, doubly-pointed, black, solid, and fracture easily (Fig. 2C to E). Microscopic examination of the The confusion in the literature between hair [trichomes] and abaxial surfaces of compressions at ×250 magnification, and critical ob- extracuticular deposits stems mainly from lack of observing the abaxial servation and photography of the compressions during maceration, surface of M. scheuchzeri compressions after being freed from show no evidence for organic attachment of the extracuticular deposits. the entombing rock matrix, though previous exceptions are noted Most importantly, they are acellular (Fig. 3). (Barthel, 1961; Cleal and Zodrow, 1989; Gothan, 1916). To untangle Fig. 4A, B shows IR spectra of individual extracuticular deposit and the confusion necessitates (1) examining freshly unearthed compres- trichome, respectively. In particular, the former is relatively aliphatic- sions immediately after collecting, (2) examining the HF solution used rich, as indicated by the larger CH2/CH3 ratio of 3.0, which at the same for freeing the compressions from the rock matrix, (3) real-time study- time implies comparatively longer and straight hydrocarbon chains ing of the compressions during Schulze's (1855) maceration process, with relatively few side branchings. This ratio is computed after and (4) examining the ammonium hydroxide solution that produces deconvolution in the 3000–2800 cm−1 aliphatic stretching region (see the cuticle. Summarizing my experimental results, extracuticular Zodrow and Mastalerz, 2001, Figs. 6 or 7). The Al/Ox ratio of aliphatics deposits (1) tend to drop-off in storage because of dehydration of the to oxygen-containing compounds [(3000–2800/1800–1500) cm−1 exposed compression (not of the extracuticular deposits), (2) were band] is comparatively very small at 0.32, which suggests a significant found loose in the plastic dishes, (3) had slowly solubilized on the com- joint contribution of oxygen-containing groups and aromatic carbon. pression in ca 3–5 h, and (4) intact trichomes were found in Petri The peak at ~3400 cm−1 is due to hydroxyl absorbance, at 1727 cm−1 dishes, correlating with structural holes found subsequently in the (C_O) ester, at 1634 cm−1(C_O) ketones, at 1385 cm−1 (symmetric corresponding abaxial cuticles. COO\), and at 1029 cm−1 (C\O\C) ether, or Si\O stretch in silicates Microscopic observations include that trichomes (ca. 300 μmlong) (see Chen et al., 2012). The inescapable conclusion is that extracuticular are not ordinarily visible on compressions, freed or still attached to deposits are the last physiological event in the development of the the rock matrix, but molds of extracuticular deposits are marked M. scheuchzeri pinnule, representing an excreted biochemical deposit. (Fig. 2A), and extracuticular deposits flatly overlie the abaxial venation In contrast, the trichomatous IR spectrum is relatively poor in terms at an acute angle in a more or less consistent parallel arrangement with of functional groups, confirmed by a second spectrum. In particular, 16 E.L. Zodrow / International Journal of Coal Geology 121 (2014) 14–18 Fig. 2. Macroneuropteris scheuchzeri. (A) Impressions (molds) of extracuticular deposits. (B) Extracuticular deposits overlying lateral veins of the abaxial pinnule. (A) and (B) represent the same compression. (C) Extracuticular deposit, round, opaque, solid. (D) Diagenetically altered extracuticular deposits in situ showing pointed terminals. No chemical treatment was ap- plied. (E) Extracuticular deposit, detail of a pointed terminus. (C) and (E) Nomarski phase-contrast micrsocopy. absent are the aliphatic stretching bands (C\H) that are necessary for calculating the two ratios mentioned (D'Angelo et al., 2010; D'Angelo et al., 2013). 4. Discussion 4.1. Comparison of IR-spectra: extracuticular deposit vs compression The combination of long and straight aliphatic chains with increas- ing contents of oxygenated/aromatic carbon groups does not fitthe usual
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