Molecular evolution of pteridophytes and their relationship to seed plants: Evidence from complete 18S rRNA gene sequences

Complete 18S ribosomal RNA sequence data from representatives of all extant pteridophyte lineages together with RNA sequences from different seed plants were used to infer a molecular phylogeny of vascular plants that included all major land plant lineages. The molecular data indicate that lycopsids are monophyletic and are the earliest diverging group within the vascular land plants, whereasPsilotum nudum is more closely related to the seed plants than to other pteridophyte lineages. The phylogenetic trees based on maximum likelihood, parsimony and distance analyses show substantial agreement with the evolutionary relationships of land plants as interpreted from the fossil record.     

Ultrastructure and development during meiosis and the tetrad period of sporogenesis in the leptosporangiate fern Alsophila setosa (Cyatheaceae) compared with corresponding stages in Psilotum nudum (Psilotaceae)

The pre-meiotic, meiotic and tetrad stages of development in microsporangia of Alsophila setosa were studied with particular emphasis on the early establishment of patterning in the microspore wall and the subsequent development of the sporoderm. The data obtained were compared with corresponding ontogenetic stages of Psilotum nudum. Tapetal behaviour was also examined. During the tetrad period, only one layer, a thin undulating sheet, appeared alongside the plasma membrane of the tetraspores, and this was evidently formed on a pre-patterned structure – a fibrillar layer, corresponding to a kind of primexine matrix. The early free microspores had a wavy plasma membrane with a parallel, sinusoidal, thin initial sporoderm layer. The proximal apertural fold was observed to be an extended outgrowth of this initial spore envelope. Sporoderm ontogeny during the tetrad period in Alsophila and Psilotum show some common points, but also fundamental differences, mainly in the relative timing of events: in Alsophila the end of the tetrad period is the starting point for exospore development, whereas in Psilotum the exospore is already complete at this stage. Considerable differences were also observed in the tapetum of the two species.     

Cell division patterns in the apices of subterranean axis and aerial shoot of Psilotum nudum (Psilotaceae): morphological and phylogenetic implications for the subterranean axis

The cell division pattern in the apical meristem of Psilotum nudum was examined using epi-illumination microscopy and a paraffin method. In the subterranean axis, about half of the derivative cells of the apical cell produce tetrahedral daughter apical cells by the first three or more oblique divisions. Roughly half of these apical cells give rise to the apical meristems of axes, whereas the other half do not. Various relative activities of the mother and daughter apical cells give rise to disordered branching patterns. In the ill-organized apical meristem as well as the leafless and capless structure, the Psilotum subterranean axis differs from the basic organs of vascular plants such as stem and root and seems to be an independent organ. The cell division pattern characteristic of the subterranean axis persists in the young unbranched aerial shoots, although fewer daughter apical cells are produced. Dichotomous branching of the aerial shoots, as in a variety of organs of pteridophytes, involves loss of the mother apical cell followed by appearance of two daughter apical cells.     

Non‐angiosperm phytochromes and the evolution of vascular plants

The phytochromes, a class of plant light‐sensing pigments, are a gene family with a long, complex evolutionary history. Angiosperms each have five or more phytochromes (designated A to E in Arabidopsis ) with distinct functions as light receptors and only moderate sequence identities for different types within a species. The long‐term challenge taken up here is to trace the origin and function of the various motifs within the angiosperm phytochromes through gymnosperm phytochromes (types N, O and P) and lower plant phytochromes, sometimes reaching even to bacterial progenitor molecules. Particularly intriguing are the findings of homology of a C‐terminal region of phytochromes with bacterial transmitter modules and of a large N‐terminal region with a protein encoded by a gene from the cyanobacterum Synechocystis . Phylogenetic analysis helps to answer general questions such as the times of divergence of mono‐ and dicotyledons, of groups of gymnosperms or of ferns. Phytochrome sequences suggest (1) that mono‐ and dicotyledons became separated 150‐200 million years earlier than indicated by the fossil record and (2) that Ginkgo and Cycas have been separated unexpectedly late from the lineage giving rise to the Pinidae. (3) The status of Psilotum as a close relative of the primeval vascular plants is not supported. Phytochrome gene sequences additionally reveal that (4) moss and fern phytochromes have erratically acquired C‐termini which, though kinase‐like, are different from the common ones and that (5) introns have been lost, gained or shifted in position from algae to angiosperms. Phytochromes promise to be a rich source of phylogenetic information into the future as more sequences and functional data emerge, not least from studies of lower plants.     

Deterministic cellular descendance and its relationship to the branching of plant organ axes

Summary. A double-wall map L-system, designated as S_5-5, was developed to simulate the cellular pattern found at the summit of shoot apices of Psilotum nudum. Commencing from a 3-sided autoreproductive founder cell, fives steps of simulation established a basic set of ten different cell types. Continuing the simulation beyond the fifth step revealed that, in addition to the regular production of new 3-sided cells, a group of autoreproductive 5-sided cells came into being. A close correspondence exists between the cells of the two-dimensional simulation and the two-dimensional cellular patterns found on the epidermis of the apices of Psilotum species. The 3-sided cells produced during the simulation correspond to the potentially organogenetic 3-sided cells that can be seen upon the apical surfaces. Successive generations of these 3-sided apical cells (which are actually 4-sided tetrahedral cells when viewed in three dimensions) and their immediate descendants are thought to be selected to organise the successive pairs of apices that bring about the repeated bifurcation of the Psilotum shoots. The 5-sided cells contribute to the cellular pavements which separate these pairs of organogenetic centres, each with their 3-sided apical cells. The cellular patterns simulated by the S_5-5 system may also correspond to the cellular patterns found on the surfaces of some other pteridophyte apices, including that of the rhizophores of Selaginella species. Tritiated-thymidine labelling of rhizophore apices revealed a group of nonproliferating cells that was associated with rhizophore bifurcation and which may correspond to a group of pavement cells. Nonproliferating cells, by regulating the siting of new organogenetic centres, may have evolved as an accompaniment to branching events such as the bifurcation of root and organ axes.Correspondence and reprints: School of Biological Sciences, University of Bristol, Woodland Road, Bristol BS8 1UG, United Kingdom.     

The Distribution of a Phytochrome-Like Protein in the Fern Psilotum nudum.; An Immunoblotting Analysis of an Early Ancestor of all Vascular Plants

The distribution of a 125 kg . mol^?1 protein recognized by a monoclonal antibody raised against phytochrome of maize was analyzed in the sporophyte of the fern Psilotum nudum. Highest amounts (up to 5 μg per fresh weight) of this protein were found in the tips of expanding shoots. Green sporangia as well as the pale tips of the rhizome contained this 125 kg . mol^?1 protein, too. In the brown parts of the rhizome it was more rarely contained. Unlike phytochrome from etiolated higher plants, the Psilotum protein appeared to be scarcely degraded by the illuminated plants. In this respect the protein of Psilotum seems to resemble the small fraction of phytochrome contained in green and illuminated higher plants. Moreover, after illuminating the Psilotum rhizome for 3 d, higher amounts of this protein were detected therein as before.     

Phylogenetic affinity of arbuscular mycorrhizal symbionts in <Emphasis Type="Italic">Psilotum nudum</Emphasis>

Many lineages of land plants (from lycopsids to angiosperms) have non-photosynthetic life cycle phases that involve obligate mycoheterotrophic arbuscular mycorrhizal (AM) associations where the plant host gains organic carbon through glomalean symbionts. Our goal was to isolate and phylogenetically identify the AM fungi associated with both the autotrophic and underground mycoheterotrophic life cycle phases of Psilotum nudum. Phylogenetic analyses recovered 11 fungal phylotypes in four diverse clades of Glomus A that form AM associations with P. nudum mycoheterotrophic gametophytes and autotrophic sporophytes, and angiosperm roots found in the same greenhouse pots. The correspondence of identities of AM symbionts in P. nudum sporophytes, gametophytes and neighboring angiosperms provides compelling evidence that photosynthetic heterospecific and conspecific plants can serve as the ultimate sources of fixed carbon for mycoheterotrophic gametophytes of P. nudum, and that the transfer of carbon occurs via shared fungal networks. Moreover, broader phylogenetic analyses suggest greenhouse Psilotum populations, like field-surveyed populations of mycoheterotrophic plants, form AM associations with restricted clades of Glomus A. The phylogenetic affinities and distribution of Glomus A symbionts indicate that P. nudum greenhouse populations have the potential to be exploited as an experimental system to further study the physiology, ecology and evolution of mycoheterotrophic AM associations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

安徽大别山区蕨类植物新记录种——松叶蕨

报道了安徽大别山区蕨类植物新记录种--松叶蕨Psilotum nudum (L.) Beauv.,并描述其形态特征和国内地理分布。     

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.     

The influence of air temperature on the starting date of Quercus pollination in the South of Europe

In temperate zones the air temperature influences many aspects of the plant growth and also the time of flowering is often correlated with this environmental parameter. It is a generally accepted idea that higher temperatures in the period preceding ripening of the flowers determine earlier pollination. To verify if a correlation between the air temperature and the date of onset of the pollination period of Quercus spp. exists, a comparative study was carried out over 7 years (1995–2001) in two South-European towns: Vigo (Spain) and Perugia (Italy). Quercus pollen is released in the atmosphere of Perugia on average in the last two weeks of April while in the Spanish region the pollination occurs on average one month before.

In order to overcome the dormancy period Perugia requires 1110 Chilling Hours (CH)-884 Growth Degree-Days (GDD°C) and Vigo 709 CH-861 GDD°C. With the Ashcroft method Perugia needs 1075 CH-1000 GDD°C and Vigo 625 CH-1512 GDD°C. Heat accumulation from the end of winter dormancy to the onset of pollination, showed the highest significance when mean temperature in Perugia and maximum temperature in Vigo were used. Every year we have found that the colder station needed a lower heat accumulation: Perugia required a higher quantity of chilling and heat than Vigo. However, the correlation detected between temperature and flowering was, on average, less significant that those found in the same regions for other arboreal taxa that present winter pollination (e.g. Corylus, Alnus). This preliminary study suggests that there is an effect of air temperature on Quercus pollination, but other environmental factors, such as photoperiod, hours of light, rainfall, relative humidity, may be of great influence in determining the onset of pollination in plants with a spring flowering.