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1.
Am J Bot ; 105(3): 291-301, 2018 Mar.
Artigo em Inglês | MEDLINE | ID: mdl-29603143

RESUMO

PREMISE OF THE STUDY: For the past one billion years, green plants (Viridiplantae) have dominated global ecosystems, yet many key branches in their evolutionary history remain poorly resolved. Using the largest analysis of Viridiplantae based on plastid genome sequences to date, we examined the phylogeny and implications for morphological evolution at key nodes. METHODS: We analyzed amino acid sequences from protein-coding genes from complete (or nearly complete) plastomes for 1879 taxa, including representatives across all major clades of Viridiplantae. Much of the data used was derived from transcriptomes from the One Thousand Plants Project (1KP); other data were taken from GenBank. KEY RESULTS: Our results largely agree with previous plastid-based analyses. Noteworthy results include (1) the position of Zygnematophyceae as sister to land plants (Embryophyta), (2) a bryophyte clade (hornworts, mosses + liverworts), (3) Equisetum + Psilotaceae as sister to Marattiales + leptosporangiate ferns, (4) cycads + Ginkgo as sister to the remaining extant gymnosperms, within which Gnetophyta are placed within conifers as sister to non-Pinaceae (Gne-Cup hypothesis), and (5) Amborella, followed by water lilies (Nymphaeales), as successive sisters to all other extant angiosperms. Within angiosperms, there is support for Mesangiospermae, a clade that comprises magnoliids, Chloranthales, monocots, Ceratophyllum, and eudicots. The placements of Ceratophyllum and Dilleniaceae remain problematic. Within Pentapetalae, two major clades (superasterids and superrosids) are recovered. CONCLUSIONS: This plastid data set provides an important resource for elucidating morphological evolution, dating divergence times in Viridiplantae, comparisons with emerging nuclear phylogenies, and analyses of molecular evolutionary patterns and dynamics of the plastid genome.


Assuntos
Sequência de Aminoácidos , Evolução Biológica , Genes de Plantas , Genomas de Plastídeos , Filogenia , Viridiplantae/genética , Aminoácidos , Briófitas/genética , Classificação , Cycadopsida/genética , DNA de Plantas/análise , Conjuntos de Dados como Assunto , Evolução Molecular , Gleiquênias/genética , Genoma de Planta , Genômica/métodos , Ginkgo biloba/genética , Gnetophyta/genética , Magnoliopsida/genética , Proteínas de Plantas/genética , Plastídeos/genética
2.
J Plant Res ; 129(2): 175-87, 2016 Mar.
Artigo em Inglês | MEDLINE | ID: mdl-26858202

RESUMO

The blue light (BL) receptor phototropin (phot) is specifically found in green plants; it regulates various BL-induced responses such as phototropism, chloroplast movement, stomatal opening, and leaf flattening. In Arabidopsis thaliana, two phototropins--phot1 and phot2--respond to blue light in overlapping but distinct ways. These BL-receptor-mediated responses enhance the photosynthetic activity of plants under weak light and minimize photodamage under strong light conditions. Welwitschia mirabilis Hook.f. found in the Namib Desert, and it has adapted to severe environmental stresses such as limiting water and strong sunlight. Although the plant has physiologically and ecologically unique features, it is unknown whether phototropin is functional in this plant. In this study, we assessed the functioning of phot-mediated BL responses in W. mirabilis. BL-dependent phototropism and stomatal opening was observed but light-dependent chloroplast movement was not detected. We performed a functional analysis of the PHOT1 gene of W. mirabilis, WmPHOT1, in Arabidopsis thaliana. We generated transgenic A. thaliana lines expressing WmPHOT1 in a phot1 phot2 double mutant background. Several Wmphot1 transgenic plants showed normal growth, although phot1 phot2 double mutant plants showed stunted growth. Furthermore, Wmphot1 transgenic plants showed normal phot1-mediated responses including phototropism, chloroplast accumulation, stomatal opening, and leaf flattening, but lacked the chloroplast avoidance response that is specifically mediated by phot2. Thus, our findings indicate that W. mirabilis possesses typical phot-mediated BL responses that were at least partially mediated by functional phototropin 1, an ortholog of Atphot1.


Assuntos
Proteínas de Arabidopsis/metabolismo , Arabidopsis/fisiologia , Gnetophyta/fisiologia , Transdução de Sinal Luminoso , Fosfoproteínas/metabolismo , Fototropinas/metabolismo , Arabidopsis/genética , Arabidopsis/efeitos da radiação , Proteínas de Arabidopsis/genética , Cloroplastos/metabolismo , Gnetophyta/genética , Gnetophyta/efeitos da radiação , Luz , Mutação , Fosfoproteínas/genética , Fotossíntese/efeitos da radiação , Fototropinas/genética , Fototropismo/efeitos da radiação , Folhas de Planta/genética , Folhas de Planta/fisiologia , Folhas de Planta/efeitos da radiação , Plantas Geneticamente Modificadas , Proteínas Serina-Treonina Quinases , Plântula/genética , Plântula/fisiologia , Plântula/efeitos da radiação
3.
Mol Biol Evol ; 33(6): 1448-60, 2016 06.
Artigo em Inglês | MEDLINE | ID: mdl-26831941

RESUMO

Mitochondrial genomes (mitogenomes) of flowering plants are well known for their extreme diversity in size, structure, gene content, and rates of sequence evolution and recombination. In contrast, little is known about mitogenomic diversity and evolution within gymnosperms. Only a single complete genome sequence is available, from the cycad Cycas taitungensis, while limited information is available for the one draft sequence, from Norway spruce (Picea abies). To examine mitogenomic evolution in gymnosperms, we generated complete genome sequences for the ginkgo tree (Ginkgo biloba) and a gnetophyte (Welwitschia mirabilis). There is great disparity in size, sequence conservation, levels of shared DNA, and functional content among gymnosperm mitogenomes. The Cycas and Ginkgo mitogenomes are relatively small, have low substitution rates, and possess numerous genes, introns, and edit sites; we infer that these properties were present in the ancestral seed plant. By contrast, the Welwitschia mitogenome has an expanded size coupled with accelerated substitution rates and extensive loss of these functional features. The Picea genome has expanded further, to more than 4 Mb. With regard to structural evolution, the Cycas and Ginkgo mitogenomes share a remarkable amount of intergenic DNA, which may be related to the limited recombinational activity detected at repeats in Ginkgo Conversely, the Welwitschia mitogenome shares almost no intergenic DNA with any other seed plant. By conducting the first measurements of rates of DNA turnover in seed plant mitogenomes, we discovered that turnover rates vary by orders of magnitude among species.


Assuntos
Evolução Biológica , Genoma Mitocondrial , Ginkgo biloba/genética , Gnetophyta/genética , Mitocôndrias/genética , Sequência de Bases , Mapeamento Cromossômico , Evolução Molecular , Genes de Plantas , Genoma de Planta , Filogenia , Edição de RNA
5.
Am J Bot ; 102(2): 312-24, 2015 Feb.
Artigo em Inglês | MEDLINE | ID: mdl-25667083

RESUMO

PREMISE OF THE STUDY: The female gametophyte of Welwitschia has long been viewed as highly divergent from other members of the Gnetales and, indeed, all other seed plants. However, the formation of female gametes and the process of fertilization have never been observed. METHODS: Standard histological techniques were applied to study gametophyte development and the fertilization process in Welwitschia. KEY RESULTS: In Welwitschia, fertilization events occur when pollen tubes with binucleate sperm cells grow down through the nucellus and encounter prothallial tubes, free nuclear tubular extensions of the micropylar end of the female gametophyte that grow up through the nucellus. Entry of a binucleate sperm cell into a vacuolate prothallial tube appears to stimulate the rapid coagulation of cytoplasm around a single female nucleus, which differentiates into an egg cell. One sperm nucleus enters the female gamete, while the second sperm nucleus remains outside and ultimately degenerates. Only a single fertilization event occurs per mating pair of pollen tube and prothallial tube. CONCLUSIONS: Welwitschia lacks the gnetalean pattern of regular double fertilization, as found in Ephedra and Gnetum, involving sperm from a single pollen tube to yield two zygotes. Moreover, an analysis of character evolution indicates that the female gametophyte of Welwitschia is highly apomorphic both among seed plants, and specifically within Gnetales, but also shares several key synapomorphies with its sister taxon Gnetum. Finally, the biological role of prothallial tubes in Welwitschia is examined from the perspectives of gamete competition and kin conflict.


Assuntos
Evolução Biológica , Células Germinativas Vegetais/crescimento & desenvolvimento , Gnetophyta/genética , Filogenia , Polinização , Fertilização , Desenvolvimento Vegetal , Tubo Polínico
6.
Mol Phylogenet Evol ; 75: 24-40, 2014 Jun.
Artigo em Inglês | MEDLINE | ID: mdl-24565948

RESUMO

Living gymnosperms comprise only a little more than 1000 species, but represent four of the five main lineages of seed plants, including cycads, ginkgos, gnetophytes and conifers. This group has huge ecological and economic value, and has drawn great interest from the scientific community. Here we review recent advances in our understanding of gymnosperm evolution and biogeography, including phylogenetic relationships at different taxonomic levels, patterns of species diversification, roles of vicariance and dispersal in development of intercontinental disjunctions, modes of molecular evolution in different genomes and lineages, and mechanisms underlying the formation of large nuclear genomes. It is particularly interesting that increasing evidence supports a sister relationship between Gnetales and Pinaceae (the Gnepine hypothesis) and the contribution of recent radiations to present species diversity, and that expansion of retrotransposons is responsible for the large and complex nuclear genome of gymnosperms. In addition, multiple coniferous genera such as Picea very likely originated in North America and migrated into the Old World, further indicating that the center of diversity is not necessarily the place of origin. The Bering Land Bridge acted as an important pathway for dispersal of gymnosperms in the Northern Hemisphere. Moreover, the genome sequences of conifers provide an unprecedented opportunity and an important platform for the evolutionary studies of gymnosperms, and will also shed new light on evolution of many important gene families and biological pathways in seed plants.


Assuntos
Evolução Biológica , Cycadopsida/classificação , Genoma de Planta , Filogenia , Mapeamento Cromossômico , Cycadopsida/genética , Gnetophyta/classificação , Gnetophyta/genética , Filogeografia , Retroelementos , Análise de Sequência de DNA
7.
BMC Evol Biol ; 13: 72, 2013 Mar 27.
Artigo em Inglês | MEDLINE | ID: mdl-23530702

RESUMO

BACKGROUND: The extant Gnetales include three monotypic families, namely, Ephedraceae (Ephedra), Gnetaceae (Gnetum), and Welwitschiaceae (Welwitschia), all of which possess compound female cones that comprise a main axis and 1 to multiple pairs/whorls of bracts subtending a female reproductive unit or having lower pairs/whorls of bracts sterile. However, the evolutionary origin of such a reproductive architecture in Gnetales is controversial in the light of the competing anthophyte versus gnetifer hypotheses of seed plant relationships. Hence, macrofossils demonstrating the structure of compound female cones of the Gnetales should be important to decipher the early evolution of the order. RESULTS: A new ephedroid plant Chengia laxispicata gen. et sp. nov. is described from the Early Cretaceous Yixian Formation of western Liaoning, Northeast China. The fossil represents a part of a leafy shooting system with reproductive organs attached. The main shoot bears internodes and swollen nodes, from which lateral branches arise oppositely. Reproductive organs consist of female spikes terminal to twigs or axillary to linear leaves. Spikes are loosely arranged, having prominent nodes and internodes. Bracts of the spikes are decussately opposite and comprise 4-8 pairs of bracts. Each bract subtends an ellipsoid seed. Seeds are sessile, with a thin outer envelope and a distal micropylar tube. CONCLUSIONS: Chengia laxispicata gen. et sp. nov. provides a missing link between archetypal fertile organs in the crown lineage of the Gnetales and compound female cones of the extant Ephedraceae. Combined with a wealth of Ephedra and ephedroid macrofossils from the Early Cretaceous, we propose a reduction and sterilization hypothesis that the female cone of the extant Ephedraceae may have stemmed from archetypal fertile organs in the crown lineage of the Gnetales. These have undergone sequentially intermediate links similar to female cones of Cretaceous Siphonospermum, Chengia, and Liaoxia by reduction and sterilization of the lower fertile bracts, shortenings of internodes and peduncles as well as loss of reproductive units in all inferior bracts. The basal family Ephedraceae including Ephedra of the extant Gnetales was demonstrated to have considerable diversity by the Early Cretaceous, so an emended familial diagnosis is given here. The Jehol Biota in Northeast China and adjacent areas contains a plethora of well-preserved macrofossils of Ephedra and ephedroids that show different evolutionary stages including primitive and derived characters of Ephedraceae, so Northeast China and adjacent areas may represent either the centre of origination or one of the centres for early diversification of the family.


Assuntos
Evolução Biológica , Fósseis , Gnetophyta/classificação , Gnetophyta/genética , China , Gnetophyta/anatomia & histologia , Gnetophyta/fisiologia , Filogeografia
8.
Genome Biol Evol ; 3: 1340-8, 2011.
Artigo em Inglês | MEDLINE | ID: mdl-22016337

RESUMO

Resolving the closest relatives of Gnetales has been an enigmatic problem in seed plant phylogeny. The problem is known to be difficult because of the extent of divergence between this diverse group of gymnosperms and their closest phylogenetic relatives. Here, we investigate the evolutionary properties of conifer chloroplast DNA sequences. To improve taxon sampling of Cupressophyta (non-Pinaceae conifers), we report sequences from three new chloroplast (cp) genomes of Southern Hemisphere conifers. We have applied a site pattern sorting criterion to study compositional heterogeneity, heterotachy, and the fit of conifer chloroplast genome sequences to a general time reversible + G substitution model. We show that non-time reversible properties of aligned sequence positions in the chloroplast genomes of Gnetales mislead phylogenetic reconstruction of these seed plants. When 2,250 of the most varied sites in our concatenated alignment are excluded, phylogenetic analyses favor a close evolutionary relationship between the Gnetales and Pinaceae-the Gnepine hypothesis. Our analytical protocol provides a useful approach for evaluating the robustness of phylogenomic inferences. Our findings highlight the importance of goodness of fit between substitution model and data for understanding seed plant phylogeny.


Assuntos
Genoma de Cloroplastos , Gnetophyta/classificação , Filogenia , Sementes/genética , Traqueófitas/classificação , DNA de Cloroplastos/genética , Gnetophyta/genética , Modelos Genéticos , Traqueófitas/genética
9.
Mol Biol Evol ; 27(12): 2855-63, 2010 Dec.
Artigo em Inglês | MEDLINE | ID: mdl-20601411

RESUMO

The phylogenetic position of Gnetales is one of the most contentious issues in the seed plant systematics. To elucidate the Gnetales position, an improved amino acid substitution matrix was estimated based on 64 chloroplast (cp) genomes and was applied to cp genome data including all three lineages of Gnetales in maximum likelihood analyses of proteins. Although the initial analysis strongly supported the sister relation of Gnetales with Cryptomeria (Cupressophyta or non-Pinaceae conifers) (the "Gnecup" hypothesis), the support seems to be caused by a long-branch attraction (LBA) artifact. Indeed, by removing fastest evolving proteins that are most likely associated with the LBA, the support drastically declined. Furthermore, another analysis of partial genome data with dense taxon sampling of conifers showed that, in psbC, rpl2, and rps7 proteins, there are many parallel amino acid substitutions between the lineages leading to Gnetales and to Cryptomeria, and by further excluding these three genes, the sister relation of Gnetales with Pinaceae (the "Gnepine" hypothesis) became supported. Overall, our analyses indicate that the LBA and parallel substitutions cause a seriously biased inference of phylogenetic position of Gnetales with the cp genome data.


Assuntos
Cloroplastos/genética , Genoma de Cloroplastos , Gnetophyta/genética , Filogenia , Substituição de Aminoácidos , Evolução Molecular , Genes de Plantas , Funções Verossimilhança , Dados de Sequência Molecular , Pinaceae/genética , Análise de Sequência de DNA
10.
BMC Evol Biol ; 10: 183, 2010 Jun 17.
Artigo em Inglês | MEDLINE | ID: mdl-20565755

RESUMO

BACKGROUND: Knowledge on fossil and evolutionary history of the Gnetales has expanded rapidly; Ephedra and ephedroids as well as the Gnetum-Welwitschia clade are now well documented in the Early Cretaceous. However, hypotheses on evolutionary relationships among living and fossil species are hampered by restricted knowledge of morphological variation in living groups and recent studies indicate that gnetalean diversity and character evolution may be more complex than previously assumed and involve additional extinct groups (Bennettitales, Erdtmanithecales and unassigned fossil taxa). RESULTS: Here we describe a new fossil related to Gnetales, Siphonospermum simplex from the Early Cretaceous Yixian Formation, an impression/compression of a reproductive shoot. The slender main axis bears one pair of opposite and linear leaves with primary parallel venation. The reproductive units are ovoid, without supporting bracts and borne on one median and two lateral branches. The most conspicuous feature of the fossil is the long, thread-like micropylar tube formed by the integument. Each ovule is surrounded by two different layers representing one or two seed envelopes; an inner sclerenchymatous layer and an outer probably parenchymatous layer. CONCLUSIONS: The vegetative and reproductive features of Siphonospermum simplex exclude a relationship to any other group than the Gnetales. A combination of opposite phyllotaxis, linear leaves and ovules surrounded by seed envelope(s) and with a long exposed micropylar tube are known only for extant and extinct Gnetales. Siphonospermum simplex constitutes a new lineage within the Gnetales. Its morphology cannot be directly linked to any previously known plant, but the organization of the reproductive units indicates that it belongs to the Gnetum-Welwitschia clade. Based on the absence of cone bracts and the inferred histology of the seed envelope(s) it could be related to Gnetum, however, there are also affinities with the ephedran lineage, some of which are likely plesiomorphic features, others perhaps not. Phylogeny and character evolution in the Bennettitales, Erdtmanithecales and Gnetales are currently only partly understood and under debate; the exact systematic position of Siphonospermum simplex, i.e., its position within the Gnetales, cannot be resolved with certainty.


Assuntos
Fósseis , Gnetophyta/classificação , Evolução Biológica , China , Filogenia
11.
Curr Genet ; 55(3): 323-37, 2009 Jun.
Artigo em Inglês | MEDLINE | ID: mdl-19449185

RESUMO

The exact phylogenetic position of Gnetales, a small, highly modified group of gymnosperms with an accelerated rate of molecular evolution, is one of the most challenging issues for seed plant systematics. Recent results from entire plastid genome (ptDNA) sequencing revealed the absence of the entire suite of plastid ndh genes in several species of Gnetales and the pine family (Pinaceae) potentially highlighting a major structural feature linking these two groups-concerted loss of all plastid genes for the NADH dehydrogenase complex. However, the precise extent of ndh gene loss in gymnosperms has not been surveyed. Using a slot-blot hybridization method, we probed all 11 ndh genes in 162 species from 70 of 85 gymnosperm genera. We find that all ndh genes are absent across Gnetales and Pinaceae, but not in any other group of gymnosperms. This feature represents either a major synapomorphy for a clade consisting of these two lineages or, less likely, a convergent loss. Our survey substantially extends previous inferences based on more limited sampling and, if the former evolutionary interpretation is correct, it provides additional support for the contentious "gnepine" hypothesis, which places Gnetales as sister to Pinaceae.


Assuntos
Evolução Molecular , Gnetophyta/genética , NADPH Desidrogenase/genética , Filogenia , Proteínas de Plantas/genética , DNA de Plantas/química , DNA de Plantas/genética , Variação Genética , Gnetophyta/classificação , Gnetophyta/enzimologia , Mutação , Hibridização de Ácido Nucleico/métodos , Plastídeos/enzimologia , Plastídeos/genética , Especificidade da Espécie
12.
Mol Phylogenet Evol ; 52(1): 115-24, 2009 Jul.
Artigo em Inglês | MEDLINE | ID: mdl-19166950

RESUMO

The cpDNA of Welwitschia mirabilis (the only species of Welwitschiales) was recently reported to be the most reduced and compact among photosynthetic land plants. However, cpDNAs of the other two gnetophyte lineages (viz. Ephedrales and Gnetales) have not yet been studied. It remains unclear what underlining mechanisms have downsized the cpDNA. To pin down major factors for cpDNA reduction and compaction in gnetophytes, we have determined 4 complete cpDNAs, including one from each of the 3 gnetophyte orders, Ephedra equisetina, Gnetum parvifolium, and W. mirabilis, and one from the non-Pinus Pinaceae, Keteleeria davidiana. We report that the cpDNAs of E. equisetina (109,518bp) and G.parvifolium (114,914bp) are not only smaller but more compact than that of W. mirabilis (118,919bp). The gnetophyte cpDNAs have commonly lost at least 18 genes that are retained in other seed plants. Furthermore, they have significantly biased usages of AT-rich codons and shorter introns and intergenic spaces, which are largely due to more deletions at inter-operon than intra-operon spaces and removal of segment sequences rather than single-nucleotides. We show that the reduced gnetophyte cpDNAs clearly resulted from selection for economy by deletions of genes and non-coding sequences, which then led to the compactness and the accelerated substitution rates. The smallest C-values in gnetophyte nuclear DNAs and the competitive or resource-poor situations encountered by gnetophytes further suggest a critical need for an economic strategy.


Assuntos
Evolução Molecular , Genoma de Cloroplastos , Gnetophyta/genética , Filogenia , Seleção Genética , DNA de Cloroplastos/genética , DNA de Plantas/genética , Íntrons , Óperon , Edição de RNA , Alinhamento de Sequência , Análise de Sequência de DNA , Deleção de Sequência
13.
BMC Evol Biol ; 8: 130, 2008 May 01.
Artigo em Inglês | MEDLINE | ID: mdl-18452621

RESUMO

BACKGROUND: Welwitschia mirabilis is the only extant member of the family Welwitschiaceae, one of three lineages of gnetophytes, an enigmatic group of gymnosperms variously allied with flowering plants or conifers. Limited sequence data and rapid divergence rates have precluded consensus on the evolutionary placement of gnetophytes based on molecular characters. Here we report on the first complete gnetophyte chloroplast genome sequence, from Welwitschia mirabilis, as well as analyses on divergence rates of protein-coding genes, comparisons of gene content and order, and phylogenetic implications. RESULTS: The chloroplast genome of Welwitschia mirabilis [GenBank: EU342371] is comprised of 119,726 base pairs and exhibits large and small single copy regions and two copies of the large inverted repeat (IR). Only 101 unique gene species are encoded. The Welwitschia plastome is the most compact photosynthetic land plant plastome sequenced to date; 66% of the sequence codes for product. The genome also exhibits a slightly expanded IR, a minimum of 9 inversions that modify gene order, and 19 genes that are lost or present as pseudogenes. Phylogenetic analyses, including one representative of each extant seed plant lineage and based on 57 concatenated protein-coding sequences, place Welwitschia at the base of all seed plants (distance, maximum parsimony) or as the sister to Pinus (the only conifer representative) in a monophyletic gymnosperm clade (maximum likelihood, bayesian). Relative rate tests on these gene sequences show the Welwitschia sequences to be evolving at faster rates than other seed plants. For these genes individually, a comparison of average pairwise distances indicates that relative divergence in Welwitschia ranges from amounts about equal to other seed plants to amounts almost three times greater than the average for non-gnetophyte seed plants. CONCLUSION: Although the basic organization of the Welwitschia plastome is typical, its compactness, gene content and high nucleotide divergence rates are atypical. The current lack of additional conifer plastome sequences precludes any discrimination between the gnetifer and gnepine hypotheses of seed plant relationships. However, both phylogenetic analyses and shared genome features identified here are consistent with either of the hypotheses that link gnetophytes with conifers, but are inconsistent with the anthophyte hypothesis.


Assuntos
Variação Genética , Genoma de Cloroplastos , Genoma de Planta , Gnetophyta/genética , Sequência de Bases , Mapeamento Cromossômico , Evolução Molecular , Ordem dos Genes , Especiação Genética , Dados de Sequência Molecular , Filogenia , Pseudogenes
14.
Ann Bot ; 98(1): 123-40, 2006 Jul.
Artigo em Inglês | MEDLINE | ID: mdl-16675607

RESUMO

BACKGROUND AND AIMS: The extant species of the seed plant group Gnetales (Ephedra, Gnetum and Welwitschia) have been considered a remnant of a much greater, now extinct, diversity due to the pronounced differences in form and ecology among the genera. Until recently, this hypothesis has not been supported by evidence from the fossil record. This paper adds to the expanding information on Gnetales from the Early Cretaceous and describes coalified seeds from Barremian-Albian localities in Portugal and USA. METHODS: The fossils were extracted from sediment samples by sieving in water. Adhering mineral matrix was removed by chemical treatment. Seeds were investigated using light and scanning electron microscopy. Morphology and anatomy of the seeds were documented and compared with those of extant species. KEY RESULTS: The fossils share characters with extant Ephedra, for example papillae on the inner surface of the seed envelope and in situ polyplicate pollen grains that shed the exine during germination. They differ from extant Ephedra seeds in morphological and anatomical details as well as in their smaller size. Two new species of Ephedra are described together with one species assigned to a new genus of Gnetales. Other Ephedra-like seeds, for which pollen and critical morphological details are currently unknown, are also present in the samples. CONCLUSIONS: These Cretaceous seeds document that key reproductive characters and pollen germination processes have remained unchanged within Ephedra for about 120 million years or more. There is sufficient variety in details of morphology to suggest that a diversity of Ephedra and Ephedra-like species were present in the Early Cretaceous flora. Their presence in Portugal and eastern North America indicates that they were widespread on the Laurasian continent. The fossil seeds are similar to seeds of Erdtmanithecales and this supports the previously suggested relationship between Erdtmanithecales and Gnetales.


Assuntos
Ephedra/ultraestrutura , Fósseis , Sementes/ultraestrutura , Evolução Biológica , Ephedra/classificação , Ephedra/embriologia , Gnetophyta/classificação , Gnetophyta/embriologia , Gnetophyta/ultraestrutura , Microscopia Eletrônica de Varredura , Filogenia , Pólen/anatomia & histologia , Portugal , Sementes/anatomia & histologia , Sementes/citologia , Virginia
15.
J Hered ; 97(3): 270-8, 2006.
Artigo em Inglês | MEDLINE | ID: mdl-16614133

RESUMO

Aspergillus niger is an asexual, haploid fungus which infects the seeds of Namibia's national plant, Welwitschia mirabilis, severely affecting plant viability. We used 31 randomly amplified polymorphic DNA markers to assess genetic variation among 89 A. niger isolates collected from three W. mirabilis populations in the Namib Desert. While all isolates belonged to the same vegetative compatibility group, 84% were unique genotypes, and estimates of genotypic evenness and Simpson's index of diversity approached 1.0 in the three populations. Analysis of molecular variance revealed that 78% of the total variation sampled was among isolates from individual W. mirabilis plants. Lower, but significant, amounts of variation detected among isolates from different plants (12%) and different sites (10%) also indicated some site- and plant-level genetic differentiation. Total gene diversity (H(T) = 0.264) was mostly attributable to diversity within populations (H(S) = 0.217); the relatively low level of genetic differentiation among the sites (G(ST) = 0.141) suggests that gene flow is occurring among the three distant sites. Although sexual reproduction has never been observed in this fungus, parasexuality is a well-known phenomenon in laboratory strains. We thus attribute the high levels of genetic variation to parasexuality and/or wind-facilitated gene flow from an as of yet undocumented broader host range of the fungus on other desert vegetation. Given the apparent ease of transmission, high levels of genetic diversity, and potentially broad host range, A. niger infections of W. mirabilis may be extremely difficult to control or prevent.


Assuntos
Aspergillus niger/genética , Variação Genética , Gnetophyta/microbiologia , Sequência de Bases , Primers do DNA
16.
J Mol Evol ; 61(4): 425-36, 2005 Oct.
Artigo em Inglês | MEDLINE | ID: mdl-16155750

RESUMO

The trnT-trnF region is located in the large single-copy region of the chloroplast genome. It consists of the trnL intron, a group I intron, and the trnT-trnL and trnL-trnF intergenic spacers. We analyzed the evolution of the region in the three genera of the gymnosperm lineage Gnetales (Gnetum, Welwitschia, and Ephedra), with especially dense sampling in Gnetum for which we sequenced 41 accessions, representing most of the 25-35 species. The trnL intron has a conserved secondary structure and contains elements that are homologous across land plants, while the spacers are so variable in length and composition that homology cannot be found even among the three genera. Palindromic sequences that form hairpin structures were detected in the trnL-trnF spacer, but neither spacer contained promoter elements for the tRNA genes. The absence of promoters, presence of hairpin structures in the trnL-trnF spacer, and high sequence variation in both spacers together suggest that trnT and trnF are independently transcribed. Our model for the expression and processing of the genes tRNA(Thr)(UGU), tRNA(Leu)(UAA), and tRNA(Phe) (GAA) therefore attributes the seemingly neutral evolution of the two spacers to their escape from functional constraints.


Assuntos
DNA de Cloroplastos/genética , Evolução Molecular , Genes de Plantas/genética , Gnetophyta/genética , Composição de Bases , Sequência de Bases , DNA Intergênico/genética , Gnetophyta/química , Íntrons/genética , Dados de Sequência Molecular , Filogenia , Processamento Pós-Transcricional do RNA/genética , RNA de Transferência/genética , Alinhamento de Sequência
17.
Chem Biodivers ; 2(6): 773-9, 2005 Jun.
Artigo em Inglês | MEDLINE | ID: mdl-17192020

RESUMO

Four new trimeric stilbene glucosides, mirabilosides C-F (1-4) were isolated from MeOH extract of stem and root of Welwitschia mirabilis (Welwitschiaceae) along with three known stilbenoids, resveratrol (5), gnemonoside B (6), and gnetin G (7). The structures of these compounds were elucidated by spectroscopic methods.


Assuntos
Glucosídeos/química , Gnetophyta/química , Estilbenos/química , Estrutura Molecular , Raízes de Plantas/química , Caules de Planta/química
18.
J Hered ; 94(3): 212-7, 2003.
Artigo em Inglês | MEDLINE | ID: mdl-12816961

RESUMO

Welwitschia mirabilis is a monotypic member of the family Welwitchiaceae which, along with Ephedra and Gnetum species, comprises the gymnospermous order Gnetales. While the monophyly of this order is now widely accepted, the relationship of the Gnetales to other seed plants is still contentious. Despite the unique phylogenetic position of W. mirabilis and its extraordinary physiological and anatomical adaptations, little is known about the plant's phylogeny or its current distribution in isolated locations throughout the Namib Desert. As a preliminary step in the design of an more extensive phylogeographic study, we analyzed 37 random amplified polymorphic DNA (RAPD) loci from 59 plants distributed among five sites separated by distances of 6-440 km. Cluster analysis and analysis of molecular variance (AMOVA) revealed significant levels of variation within and between populations and little evidence of inbreeding. Genetic differences between populations reflect the geographic distances separating them. Three of the populations formed discernable genetic clusters, suggesting that little gene flow occurs between populations separated by > or = 18 km. In contrast, gene flow is occurring between two populations separated by only 6 km, supporting previous observations that pollen dispersal is primarily local and that seeds are not readily wind-born over the large distances separating most W. mirabilis populations. As a working hypothesis, we propose that W. mirabilis had a continuous distribution across its current range as much as 105 million years ago, and that as a consequence of subsequent drying trends and physical disturbance, populations became progressively isolated, accounting for their current distribution.


Assuntos
Variação Genética , Gnetophyta/genética , Evolução Biológica , Namíbia , Técnica de Amplificação ao Acaso de DNA Polimórfico
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