FLAVONOIDS OF THE PRIMITIVE FERNS: STROMATOPTERIS,SCHIZAEA, GLEICHENIA,HYMENOPHYLLUM, AND CARDIOMANES

The flavonoids of the primitive leptosporangiate ferns Stromatopteris moniliformis, Schizaea bifida, Gleichenia cunninghamii, Cardiomanes reniforme, and Hymenophyllum demissum have been identified as 3-O-glycosides of the flavonols kaempferol and quercetin. None of the examined ferns produced flavonoids which are also common to the Psilotoceae. The Psilotaceae have previously been shown to produce O-glycosides of amentoflavone (biflavone) and apigenin (flavone) and traces of C-glycosylflavones (Wallace and Markham, 1978). These data imply that the primitive leptosporangiate ferns are not closely related to the Psilotaceae. The antiquity of rhamnose as a glycosyl moiety is suggested by its occurrence in both the primitive leptosporangiate ferns and in the Psilotaceae.     

Systematic implications of the Psilotaceae

To assess the taxonomic relationships between the Psilotaceae and the Filicales ( = Polypodiales), certain factors including character divergence, gaps in the record, categorical consistency, conservatism of characters, and use of factual vs. hypothetical data have been given consideration. Although it has been proposed that the aerial shoots of Psilotaceae are homologous to fronds, and the appendages to pinnae, and that the Psilotaceae should be classified as Filicales,Psilotum andTmesipteris show a number of profound differences from Filicales. These involve the structure of the aerial organs, vascular patterns, fertile appendages, and sporangia. In view of the taxonomic criteria used, and the nature of the character differences, it is concluded that the Psilotaceae do not warrant inclusion in Filicales. InsteadPsilotum andTmesipteris constitute a separate line of evolution, and should be treated as a class, the Psilopsida, of Tracheophyta, equivalent to Equisetopsida, Lycopodiopsida, and Polypodiopsida.     

Morphologic and taxonomic relationships of the Psilotaceae relative to evolutionary lines in early land vascular plants

Examination of the fossil record shows that there is no direct evidence regarding the origins and early evolution of the Psilotaceae. Past comparisons betweenPsilotum orTmesipteris and certain fossil plants are reviewed. The major types of early land vascular plants are surveyed in order to demonstrate their basic characteristics and evolutionary trends and to point out the absence of any direct links with the Psilotaceae. Fossil plants of problematical affinities which appear to be important in the early diversification of ferns or plants closely related to ferns are discussed. The nature of stem-leaf relationships of some primitive, extant filicalean ferns and possibly the Psilotaceae is considered in terms of possible similar occurrences among fossil plants.     

Taxonomic and morphological relationships of the psilotaceae: Introduction to the symposium

This symposium was organized to publicly discuss the controversy over the relationships of the Psilotaceae, centering around Bierhorst’s thesis thatPsilotum andTmesipteris are filicalean ferns.     

Apigenin di-C-glycosylflavones of Angiopteris (Marattiales)

A survey of the flavonoids of four species of Angiopteris indicates that di-C-glycosylflavones and flavone-O-glycosides may be characteristic of this distinct group of eusporangiate ferns. Derivatives of flavonols, which are typical of leptosporangiate ferns and Ophioglossum, or biflavones, which are characteristic of the Psilotaceae, were not detected in Angiopteris.     

Morphological status of the shoot systems of Psilotaceae

Brittonia - This paper evaluates the morphological bases for Bierhorst’s recent proposals of the systematic alliance of Psilotaceae with leptosporangiate ferns. In particular it examines the...     

Apigenin and amentoflavone glycosides in the psilotaceae and their phylogenetic significance

Documentation of amentoflavone O-glucosides as the predominant flavonoid glycosides in both genera of the Psilotaceae clearly distinguishes this family from all other families of vascular plants. Psilotum and Tmesipteris also possess apigenin C- and O-glycosides as common flavonoid types. Apigenin 7-O-rhamnoglucoside occurs in both genera and the previously undocumented apigenin 7-O-rhamnoglucoside-4′-O-glucoside, although identified only in Tmesipteris, may also be present in Psilotum. The existence of flavone C-glycosides in both genera may provide a phytochemical relationship between the Psilotaceae and some ferns. The phylogenetic significance of these results is discussed.     

ON PHENOTYPIC EXPRESSION,MORPHOGENETIC PATTERN AND SYNANGIUM EVOLUTION IN PSILOTUM

Both the typical form and the appendageless variant of Psilotum nudum produce terminal synangia at the ultimate tips of the aerial axes. One clone in particular of the typical appendaged form produced synangia entirely at the tips of the aerial branches, as in the appendageless variant, and also developed occasional lateral transitional entities on the upper aerial axis displaying appendagelike and axislike morphological qualities. A developmental comparison of synangium development at the ultimate tips of aerial branches and of unusually elongating and normal sized fertile-appendages showed that the morphogenetic pattern of synangium development was similar. Anatomical and morphological evidence showed the synangium to be derived from terminal subdivisions or bifurcations of the apical meristem of each structure studied. This supports the phyletic concept that the synangium of the Psilotaceae is basically terminal to an axis or an axis homologue, and that it probably evolved from terminal bifurcative branching. Occasional multiple sporangium lobes may be formed on a P. nudum synangium which may not be represented by vascular bundles. Two hypothetical phyletic models of synangium evolution are proposed that could be used to explain this phenomenon and which should be tested by further evidence. Typical and appendageless P. nudum were compared in their morphogenetic pattern developed at the upper axis vegetative apical meristem, and a reconciliation was made between the structuring of the apparently disparate forms, which involved the presence or absence of serial ordering in apical derivatives. It is suggested that this could serve as a model for appendage evolution in the family Psilotaceae.     

Refractive spherules in phloem ofMicrosorium scolopendria andPsilotum nudum

Summary Phloem tissues ofMicrosorium scolopendria (Polypodiaceae) andPsilotum nudum (Psilotaceae) were examined with light and electron microscopes. The characteristic refractive spherules in the sieve elements ofM. scolopendria apparently develop from endoplasmic reticulum-derived cytoplasmic vesicles. In both taxa they have not been observed to be spatially related to plastids or mitochondria. Refractive spherules contain protein and often occur in the peripheral cytoplasm of mature sieve elements. InM. scolopendria they also occur in pericycle cells. Significant differences in refractive spherule substructure occur between the two taxa studied.     

Book Reviews

Book reviewed in this article:
Winter Garden Glory. Adrian Bloom
Gardening with Perennials Month by Month. Joseph Hudek
Flora Europaea. Volume 1, second edition, Psilotaceae to Platanaceae. Edited by T. G. Tutin, N. A. Burgess, A. O. Chater, J. R. Edmondson, V. H. Heywood, D. M. Moore, D. H. Valentine, S. M. Walters and D. A. Webb, assisted by J. R. Akeroyd and M. E. Newton
Bark. The Formation, Characteristics and Usefulness of Bark around the World. Ghillean Tolmie Prance and Anne E. Prance     

A PHYTOCHEMICAL APPROACH TO THE GLEICHENIACEAE

The Gleicheniaceae, containing approximately 120 species, is a family of primitive ferns distributed throughout most of the tropics and subtropics. Historically the family was considered to be monogeneric; however, most recent authors using morphological, anatomical and cytological information have subdivided the group into 5 or 6 distinct genera. Some authors have even elevated certain species to family or ordinal levels (e.g., Platyzoma and Stromatopteris). The distribution of flavonol-3-0-glycosides supports the segregation of the Gleicheniaceae, sensu lato, into at least three groupings. Flavonoids similar to those identified within the Psilotaceae (flavones, biflavones, and C-glycosylflavones) were not detected during this study.     

SEM studies on vessels in ferns. 17. Psilotaceae

Perforation plates are reported in aerial and subaerial axes of Psilotum nudum and in aerial axes of Tmesipteris obliqua. In Psilotum, both perforations lacking pit membranes and perforations with pit membrane remnants were observed. Perforation plates in Psilotum may consist wholly of one type or the other. In Tmespteris, perforations have threadlike pit membranes or consist of porose pit membranes. Wide perforations alternating with narrow pits, a conformation observed in various ferns, were observed in Psilotum (subaerial axes). In Psilotum, perforations are more common in metaxylem than in protoxylem; perforations in protoxylem consist of primary wall areas containing small circular porosities or relatively large circular to oval perforations. There are no modifications in the secondary wall framework of protoxylem or metaxylem in Psilotum or Tmesipteris that would permit one to distinguish presence of perforations or perforation plates with light microscopy, and scanning electron microscopy (SEM) is required for demonstration of porose walls or perforations. The tracheary elements of the Psilotaceae studied have no features not also observed in other ferns with SEM.     

Tapetal Organization During Sporogenesis in Psilotum nudum

Stages in the differentiation of the tapetum of Psilotum nudumare described. Two concurrently occurring components of thetapetum can be recognized. A plasmodial tapetum with associatedfunctional nuclei develops within the sporangial loculus duringthe early stages of differentiation, appears to remain viablefor several months, that is during the entire period of sporogenesis,and undergoes reorganization on three occasions. During MeiosisI groups of spore mother cells are enclosed in clear areas withinthe plasmodium: by the end of Meiosis II each tetrad is isolatedin a plasmodial chamber; and, finally, mature spores are enclosedwithin individual tapetal chambers. Typically enlarged cellsare present during the development of a cellular, parietal tapetum.A sporopollenin-containing layer or tapetal membrane characteristicof a secretory tapetum develops on the inner tangential walland lines the surface of the loculus. This tapetal membranepersists even after dehiscence of the sporangium. These observationsare discussed in relation to previously published conflictingdata and may be relevant to the arguments concerning the relationshipof the Psilotaceae to the Filicales. Psilotum nudum, light microscopy, parietal tapetum, plasmodial tapetum, tapetal membrane, tapetal reorganization, sporogenesis, sporopollenin     

Naturally occurring neuronal NO-synthase inactivators found in Nicotiana tabacum (Solanaceae) and other plants

NO-synthase (NOS) is a heme-containing enzyme that catalyzes the oxidation of L-arginine to nitric oxide, an important cellular signaling molecule. Recently, it was found that aqueous extracts of tobacco cigarettes cause the inactivation of the neuronal isoform of NOS (nNOS) and that this may explain some of the toxicological effects of smoking. Although the exact identity of the chemical inactivator(s) is not known, we wondered if extracts prepared from other plants, including those closely related to tobacco, Nicotiana tabacum (Solanaceae), would similarly inactivate nNOS. We examined 33 plants, representing diverse members of the plant kingdom ranging from whisk fern, Psilotum nudum (Psilotaceae) to tobacco and discovered 18 plants that contain a chemical inactivator(s) of nNOS. Of these plants, 16 are members of the core asterids flowering plant group. Of these asterids, 6 are members of the Solanaceae family, of which tobacco is a member. Based on the phylogenetic relationship of the plants, it is possible that the same chemical or related chemical inactivator(s) exist. This, in turn, may help elucidate the structure of the chemical(s), as well as provide a source of a potentially novel inactivator of nNOS. The alkaloid nicotine can be excluded as putative nNOS inhibitor.     

轮藻和陆地植物系统发育及其进化

Charophytic algae and land plants together make up a monophyletic group, streptophytes, which represents one of the main lineages of multicellular eukaryotes and has contributed greatly to the change of the environment on earth in the Phanerozoic Eon. Significant progress has been made to understand phylogenetic relationships among members of this group by phylogenetic studies of morphological and molecular data over the last twenty-five years. Mesostigma viride is now regarded as among the earliest diverging unicellular organisms in streptophytes. Characeae are the sister group to land plants. Liverworts represent the first diverging lineage of land plants. Hornworts and lycophytes are extant representatives of bryophytes and vascular plants, respectively, when early land plants changed from gametophyte to sporophyte as the dominant generation in the life cycle. Equisetum, Psilotaceae, and ferns constitute the monophyletic group of monilophytes, which are sister to seed plants. Gnetales are related to conifers, not to angiosperms as previously thought. Amborella, Nymphaeales, Hydatellaceae, Illiciales, Trimeniaceae, and Austrobaileya represent the earliest diverging lineages of extant angiosperms. These phylogenetic results, together with recent progress on elucidating genetic and developmental aspects of the plant life cycle, multicellularity, and gravitropism, will facilitate evolutionary developmental studies of these key traits, which will help us to gain mechanistic understanding on how plants adapted to environmental challenges when they colonized the land during one of the major transitions in evolution of life.     

Evaluation of atpB nucleotide sequences for phylogenetic studies of ferns and other pteridophytes

Inferring basal relationships among vascular plants poses a major challenge to plant systematists. The divergence events that describe these relationships occurred long ago and considerable homoplasy has since accrued for both molecular and morphological characters. A potential solution is to examine phylogenetic analyses from multiple data sets. Here I present a new source of phylogenetic data for ferns and other pteridophytes. I sequenced the chloroplast gene atpB from 23 pteridophyte taxa and used maximum parsimony to infer relationships. A 588-bp region of the gene appeared to contain a statistically significant amount of phylogenetic signal and the resulting trees were largely congruent with similar analyses of nucleotide sequences from rbcL. However, a combined analysis of atpB plus rbcL produced a better resolved tree than did either data set alone. In the shortest trees, leptosporangiate ferns formed a monophyletic group. Also, I detected a well-supported clade of Psilotaceae (Psilotum and Tmesipteris) plus Ophioglossaceae (Ophioglossum and Botrychium). The demonstrated utility of atpB suggests that sequences from this gene should play a role in phylogenetic analyses that incorporate data from chloroplast genes, nuclear genes, morphology, and fossil data.     

Amplification of noncoding chloroplast DNA for phylogenetic studies in lycophytes and monilophytes with a comparative example of relative phylogenetic utility from Ophioglossaceae

Noncoding DNA sequences from numerous regions of the chloroplast genome have provided a significant source of characters for phylogenetic studies in seed plants. In lycophytes and monilophytes (leptosporangiate ferns, eusporangiate ferns, Psilotaceae, and Equisetaceae), on the other hand, relatively few noncoding chloroplast DNA regions have been explored. We screened 30 lycophyte and monilophyte species to determine the potential utility of PCR amplification primers for 18 noncoding chloroplast DNA regions that have previously been used in seed plant studies. Of these primer sets eight appear to be nearly universally capable of amplifying lycophyte and monilophyte DNAs, and an additional six are useful in at least some groups. To further explore the application of noncoding chloroplast DNA, we analyzed the relative phylogenetic utility of five cpDNA regions for resolving relationships in Botrychium s.l. (Ophioglossaceae). Previous studies have evaluated both the gene rbcL and the trnL(UAA)-trnF(GAA) intergenic spacer in this group. To these published data we added sequences of the trnS(GCU)-trnG(UUC) intergenic spacer + the trnG(UUC) intron region, the trnS(GGA)-rpS4 intergenic spacer+rpS4 gene, and the rpL16 intron. Both the trnS(GCU)-trnG(UUC) and rpL16 regions are highly variable in angiosperms and the trnS(GGA)-rpS4 region has been widely used in monilophyte phylogenetic studies. Phylogenetic resolution was equivalent across regions, but the strength of support for the phylogenies varied among regions. Of the five sampled regions the trnS(GCU)-trnG(UUC) spacer+trnG(UUC) intron region provided the strongest support for the inferred phylogeny.