Auxospore formation inLicmophora (Bacillariophyta)

InLicmophora gracilis var.anglica two auxospores are produced per pair of mother-cells, through the allogamic fusion of migratory and stationary gametes. Both active gametes are produced from the same mother-cell and hence both zygotes are formed in the other mother-cell. Pairing can occur between two stalked cells, or between a stalked cell and a detached cell; in the latter case the migratory gametes derive from the detached cell. The auxospores expand parallel to one another and to the apical axis of the donor mother-cell. Behavioural anisogamy of this kind, which may be termed thecis-type, seems to be characteristic of most araphid pennates and contrasts with thetrans-type exhibited byCymbella, Gomphonema and some other raphid taxa, where each mother-cell produces one migratory and one stationary gamete.     

PERIZONIUM AND INITIAL VALVE FORMATION IN THE DIATOM NAVICULA CUSPIDATA (BACILLARIOPHYCEAE)1

This paper describes the perizonium and initial valve formation in Navicula cuspidata Kütz., based on light microscope (LM) and scanning electron microscope (SEM) observations. The perizonium consists of concentric over-lapping bands, laid down sequentially at the tips of the expanding biconical auxospore during its elongation. The central perizonial band has fimbriate edges and is considerably more rigid than the more distal bands. During auxospore elongation and the band secretion, the chloroplasts continuously oscillate between the two ends of the cell; this oscillation ceases once the elongation is complete. The initial valves, formed within the perizonium, are molded into the basically biconical shape of the perizonium except for a central flattening of each valve face. In contrast to the raphes in gametangial and vegetative valves which are surrounded by a smooth axial area, the raphes in initial valves lie within a raised ridge running along the apical axis of the valve. The regular pattern of apically oriented ridges on the outer surface of vegetative valves is also lacking on initial valves. Comparison of pore–pore spacing within striae of gametangial valves, initial values and post-initial valves (first division and vegetative cells) reveals that the pore–pore distance within striae is conserved at all sexual stages. However, the distance between striae is considerably larger in initial valves than in gametangial and post-initial valves. Vegetative interstriae spacing as well as the planar morphology of the valve face is regained at the first division of the initial cell. This suggests that the spacing between striae is dependent on the sexual stage of the cell during valve formation (i.e. not directly dependent on the cell size) and can be altered independently of the pore–pore spacing.     

Two new species of the diatom genusChaetoceros (Bacillariophyta)

Two new planktonic diatom species of the genusChaetoceros are described herein:Ch. octagonus andCh. rectus, from material of Baja California coasts and the Gulf of California, respectively;Ch. rectus was also found in a sample from Australian waters.Ch. octagonus is a robust species, with a characteristic pyramidal valve shape, heavily silicified valves and setae, and one central rimoportula on every valve in the chain; it is compared with related species, which show a similar morphology, especially by electron microscopy. It belongs into subg.Chaetoceros, sect.Borealia. Ch. rectus is placed in subg.Hyalochaete, and provisionally within sect.Anastomosantia. Its major characters are: chains, cells, and setae are delicate and fragile, cells are joined in the chain by a bridge which connects sibling intercalary setae, valves and setae are weakly silicified, and one rimoportula is present on terminal valves only. Comparison is made with another species of the same section.Ch. rectus shows a different manner of chain-forming within the genus than previously described in the literature.     

Silicification of auxospores in the araphid diatom Tabularia fasciculata (Bacillariophyta)

We report on auxospore wall structure and development in the araphid pennate diatom Tabularia fasciculata. Similar to most other pennates, these auxospores showed a typical bidirectional elongation, but unexpectedly bore no transverse perizonium, and with no detectable silicon during much of their expansion. Energy dispersive X-ray spectroscopy (EDS) analyses segregated auxospores into two types: (1) those containing no detectable silicon and (2) those with measureable amounts. Both types were of similar size. Silica precipitation began throughout the auxospore at or near maximal length, but initially was detectable in isolated regions throughout the structure. Following this initial condition, silicon was consistently detectable throughout auxospores of comparable size and corresponded to deposition of longitudinal perizonium (visible through the thin organic outer layer of the wall in some auxospores), followed by the deposition of the initial valves. Our results raise the question as to how the tubular shape of bidirectionally expanding auxospores up to ∼90 μm long is maintained in the absence of transverse siliceous elements restricting isodiametric expansion of the cell, which are present in all other known pennate auxospores and all but one other diatom. Our study is the first to systematically examine mineral elements of the auxospore wall analytically.     

Cladistic analysis of some freshwater araphid diatoms (Bacillariophyta) with particular reference toDiatoma andMeridion

Interrelationships of the ribbed araphid diatoms have been reinvestigated using a cladistic analysis of a revised morphological dataset. The presence of silicified transapical ribs has previously been used to distinguish between different groups and its ability to serve as a distinguishing feature is reevaluated. The importance of valve symmetry, not fully appreciated in a previous paper, is examined. The evolution of heteropolarity, and its importance in taxonomy appear more complex than is implied by a simple dichotomy dividing groups of species into heteropolar and isopolar forms, involving an inferred evolutionary transformation from one condition to the other. This analysis proposes a close relationship betweenDiatoma, Fragilariforma, Asterionella, Distrionella, Meridion, Tabellaria, andTetracyclus. It also confirms the sub-division ofDiatoma into two closely related taxa. The genusMeridion, however, should not include both heteropolar and isopolar species in a single genus. Although the species are closely related, the isopolar species are more closely related toTabellaria andTetracyclus than they are to the heteropolar species ofMeridion.     

A probabilistic model of cell size reduction in Pseudo-nitzschia delicatissima (Bacillariophyta)

The pennate planktonic diatom Pseudo-nitzschia delicatissima is very common in temperate marine waters and often responsible for blooms. Due to its surrounding rigid silicate frustrule the diatom undergoes successive size reduction as its vegetative reproduction cycle proceeds. Since a long time the life cycle of diatoms has raised scientific interest and some years ago extensive samples of Pseudo-nitzschia have been taken from coastal waters. Mating and cell size reduction experiments were carried out and served us as a data basis for a probabilistic model of cell size reduction.We applied a homogenous non-stationary continuous-time Markov chain to model the development of individual diatoms from an initial size of about until cell death which occurred when the size reached its low at about . In contrast to conventional curve fitting models we are capable of calculating confidence intervals for estimates of the population ages as well as integrate the process of auxospore formation into the model. We thus propose a unique way to describe the stationary size distribution in a diatom population in terms of cell division and auxospore formation probabilities of its individuals.     

Resource allocation to sexual and vegetative reproduction in a forest herbSyneilesis palmata (Compositae)

The balance between sexual and vegetative reproduction inSyneilesis palmata was examined in relation to environmental conditions and the amount of reproductive resources, which is defined here as the total quantity of dry matter invested in both modes of reproduction. The allocation balance was measured for individual plants of two populations, with different densities in an open habitat, for 2 years, and those of two other populations, under different light intensities in a plantation forest (forest floor and edge), for 3 years. Relative allocation to sexual reproduction decreased with increasing reproductive resources in all populations except for the forest edge, which showed a constant allocation balance. The high density population showed lower relative allocation to sexual reproduction than the low density population, irrespective of the amount of reproductive resources. However the between-year comparison of the high density population suggested that under extremely high density, plants with a small amount of reproductive resources enhanced sexual reproduction, while plants with a large amount of reproductive resources reproduced vegetatively. On the forest floor, plants with small amounts of reproductive resources had higher relative allocation to sexual reproduction than the forest edge population, while plants with large amounts of reproductive resources had a somewhat lower one. The adaptive significance of such allocation patterns are discussed, based on qualitative data on the characteristics of both types of offspring.     

The role of sexual and clonal reproduction in maintaining population in Fritillaria camtschatcensis (L.) Ker-Gawl. (Liliaceae)

Fritillaria camtschatcensis can reproduce by means of both sexual reproduction and clonal multiplication. Despite prolific seed production, no seedlings have been found for several years in natural populations on Mt Hakusan. The purposes of this paper are to clarify: (i) whether population size is maintained mainly by clonal multiplication; and (ii) if this is the case, to what extent occasional seedling establishment affects population growth rate and population structure. Two permanent quadrats were placed in subalpine meadows in 1992 on Mt Hakusan. Plant size, location and reproductive states for all ramets in the quadrats were recorded every year. Projection matrices were created based on field census, and computer simulation experiments were performed. Fritillaria camtschatcensis had two types of flower, male flower and cosexual flower, and they were changeable. This is the first report on sex lability in Fritillaria. Clonal growth was more closely correlated with life-history stages, especially with sexual states than with plant size. The population growth rate, , was 1.006 for the Mizuyajiri population and 1.047 for the Nanryu population, respectively. Seedlings were found in 1996 for the first time. These facts indicate that populations of F. camtschatcensis on Mt Hakusan can usually be maintained by clonal multiplication. However, it is not yet certain whether seeds germinate every year or whether a flush of seedling emergence occurs once in every few years in natural populations. Computer simulation revealed that: (i) there was a critical germination rate above which population growth rate suddenly increased; and (ii) occasional seedling establishment could provide almost the same contribution to population growth rate as that of annual seedling establishment. These results suggest that population size can be maintained mainly by clonal multiplication, and the role of sexual reproduction lies beyond maintaining the population size in F. camtschatcensis.     

Evidence for Positive Selection on a Sexual Reproduction Gene in the Diatom Genus Thalassiosira (Bacillariophyta)

Single likelihood ancestor counting (SLAC), fixed effects likelihood (FEL), and several random effects likelihood (REL) methods were utilized to identify positively and negatively selected sites in sexually induced gene 1 (Sig1) of four different Thalassiosira species. The SLAC analysis did not find any sites affected by positive selection but suggested 13 sites influenced by negative selection. The SLAC approach may be too conservative because of low sequence divergence. The FEL and REL analyses revealed over 60 negatively selected sites and two positively selected sites that were unique to each method. The REL method may not be able to reliably identify individual sites under selection when applied to short sequences with low divergence. Instead, we proposed a new alignment-wide test for adaptive evolution based on codon models with variation in synonymous and nonsynonymous substitution rates among sites and found evidence for diversifying evolution without relying on site-by-site testing. The performance of the FEL and REL approaches was evaluated by subjecting the tests to a type I error rate simulation analysis, using the specific characteristics of the Sig1 data set. Simulation results indicated that the FEL test had reasonable Type I errors, while REL might have been too liberal, suggesting that the two positively selected sites identified by FEL (codons 94 and 174) are not likely to be false positives. The evolution of these codon sites, one of which is located in functional domain II, appears to be associated with divergence among the three major Thalassiosira lineages. Electronic Supplementary Material Electronic Supplementary material is available for this article at and accessible for authorised users. [Reviewing Editor: Dr. Martin Kreitman]     

On the biology of female Epiplatys bifasciatus (Cyprinodontidae) from southern Sudan

The biology of female Epiplatys bifasciatus (Steindachner, 1881) from the Sudd region of southern Sudan was investigated. The population was dominated by females whose growth in body dimensions relative to total length was found to be linear. The reproductive system differs from that of oviparous teleosts in having a portion of the ovary (anterior) modified for carrying the developing brood. Gradual development of oocytes towards the nursing portion of the ovary, and low absolute fecundity of the brood, suggests a rather long breeding season (June to September).A large variety of food organisms belonging to three main groups, viz. insects, crustaceans and arachnids, were taken by Epiplatys collected from the littoral zones of two rivers, the Bahr El Ghazal and the White Nile. Little variation occurred in the food of these two populations. Chironomid larvae showed the highest occurrence frequency in the fish food for both localities, and insects in general constituted over 80% of all organisms eaten.     

Clonal and genetic structure of two Mexican oaks: Quercus eduardii and Quercus potosina (Fagaceae)

Quercus eduardii and Q. potosina are dominant oak species in Sierra Fría, Aguascalientes, Mexico. These species have been exploited for multiple purposes since the 16th century. Both species produce clonal offspring through root suckering and acorns through sexual reproduction. To understand clonality for the implementation of the most adequate actions for the conservation of these species, we addressed the following questions: (a) what is the spatial clonal structure of both species? (b) How much clonal and genetic diversity is maintained in these species? Random Amplified Polymorphic DNAs (RAPDs) were used as molecular markers for these analyses. Genets of both species have few ramets and these grow close the parent tree. Autocorrelation analyses at the ramet level showed an aggregated distribution at short distances and a random spatial distribution at larger distances. Also, at the genet level the autocorrelation analyses showed a random distribution. Clonal diversity was high in both species (Q. eduardii: D=0.963, G/N=0.60; Q. potosina: D=0.985, G/N=0.65). Genetic diversity was high within populations (Q. eduardii: H _e =0.33±0.11; Q. potosina: H _e =0.35±0.11). Low levels of genetic differentiation among populations were observed (Q. eduardii ? _st =0.19, P < 0.002; Q. potosina ? _st =0.13, P < 0.002). Both species maintain high levels of clonal and genetic diversity, probably due to successful sexual reproduction, which allows gene flow among populations. Conservation and/or reforestation programs must include seed collections and germplasm banks. Due to the small genet size and the high clonal diversity of these species, seeds can be collected in any place in Sierra Fría, Aguascalientes.     

AUTOGAMOUS AUXOSPORULATION IN PINNULARIA NODOSA (BACILLARIOPHYCEAE)1

Auxosporulation of the freshwater epipelic diatom Pinnularia nodosa (Ehrenb.) W. Sm. was studied in a clonal culture. Interphase cells possessed two chloroplasts with invaginated pyrenoids. The nucleus contained a single small body of heterochromatin at one end, also visible during most of meiotic prophase. During auxosporulation, induced by transfer of stationary‐phase cells to fresh medium and suppressed by high nitrogen (N), an unpaired mother cell produced a single auxospore. Although meiosis II and nuclear fusion were not observed, indirect evidence indicated that auxosporulation was autogamous (rarely reported in pennate diatoms), rather than apomictic; paedogamy was excluded. The protoplast produced after meiosis either (1) matured into a “pseudozygote,” via an asymmetrical contraction after meiosis I to form a single spherical cell at one end of the mother cell (pathway 1); or (2) constricted into two spherical cells (pathway 2). In pathway 2, the “pseudogametes” never fused and only one or none developed into a pseudozygote and then into an auxospore. Pathway 2 could be suppressed by continuous light. During metamorphosis of the spherical pseudozygote into an elongate young auxospore, a complete covering of thin siliceous incunabular strips was formed, separate from the organic wall formed around the pseudozygote when first formed and from the perizonium. Mature auxospores produced via pathway 2 had 60% of the volume as those produced via pathway 1 and had smaller chloroplasts (through loss of fragments during protoplast cleavage), but they achieved exactly the same lengths, suggesting that absolute length is monitored during expansion.     

Spore formation in the Actinoplanaceae (Actinomycetales)

Spore development in four genera, Actinoplanes, Dactylosporangium, Planomonospora, and Streptosporangium, was studied by transmission and scanning electron microscopy. Actinoplanes and Streptosporangium formed spores by fragmentation of a hypha within its expanded outer sheath, as do many other actinomycetes. Dactylosporangium and Planomonospora formed spores endogenously by development of wall material within the parent hypha. In this respect, they resembled the genera Actinobifida and Thermoactinomyces. The term sporangium has therefore been used to describe structures which are not homologous. It was suggested that the term should be confined to structures in which endogenous spore formation occurs.     

Plastid inheritance in the planktonic raphid pennate diatom Pseudo-nitzschia delicatissima (Bacillariophyceae)

Plastid inheritance was followed during sexual reproduction in the raphid pennate diatom Pseudo-nitzschia delicatissima, using rbcL haplotypes as plastid identification tools. Pseudo-nitzschia species are dioecious and show functional anisogamy with 'male' mating type+(PNd(+)) cells and 'female' PNd(-) cells. Vegetative cells possess two plastids. In P. delicatissima, meiosis results in two gametes that both contribute two plastids to the zygote. The latter initially contains four plastids, but during auxospore development two of these four seem to disappear, and the initial cell emerging from the auxospore appears to contain only two. Here we assessed if the plastids are inherited strictly unipaternally, strictly biparentally, or randomly. We traced the source of the plastids in the F(1) generation by using PNd(+) and PNd(-) parental strains with different rbcL genotypes, here denoted AA (homoplastidial, with two plastids of rbcL haplotype A) and BB (homoplastidial; two plastids of haplotype B). Results showed that 16 out of 96 strains raised each from single F(1) cells had retained two paternal (PNd(+)) plastids, 20 had two maternal (PNd(-)) plastids and the remaining 60 had one maternal and one paternal plastid. This pattern is in accordance with the hypothesis that either two of the four plastids are eliminated during auxospore formation, or that all plastids are retained in the auxospore and segregate in pairs joining at random during the first mitotic division of the initial cell. Heteroplastidic F(1)-strains retained the AB genotype throughout the vegetative phase of their life cycle. The finding that 60 out of 96 F(1) strains were heteroplastidial contrasts with an absence of such genotypes in our strains raised from single cells sampled in the Gulf of Naples.     

异极藻属(硅藻门)5种中国新记录

对采自海南省三亚亚龙湾国家森林公园,湖北省神农架大九湖湿地和黑龙江省兴凯湖湿地的硅藻标本进行分类学研究,发现异极藻属植物中国新记录5种,分别是狭状披针异极藻(Gomphonema acidoclinatum LangeBertalotReichardt)、矛形异极藻(Gomphonema hasta D.Metzeltin)、林氏异极藻(Gomphonema kobayashiae MetzeltinLange-Bertalot)、宽缘异极藻(Gomphonema latipes E.Reichardt)和隐形异极藻(Gomphonema occultum ReichardtLange-Bertalot)。对相似种类之间的形态差异进行比较,为中国异极藻属植物的分类学研究提供基本资料。     

Sorus development on median and marginal parts of the sporophyte of Laminaria japonica Areschoug (Phaeophyceae)

Sorus formation on median and marginal parts of Laminaria japoncia Areschoug was investigated by cultivating sporophytes from December 1996 to September 1997 in Southern Hokkaido, Japan. Simultaneously, discs (ca. 3 cm in diameter) that were cut off monthly from median and marginal parts of the cultivated sporophytes were incubated in our laboratory. All the cultivated sporophytes formed sori on the median parts from January to August, while all the median and marginal discs formed sori at 10 under 46 μmol photon m^-2s^-1 (12L:12D) in enriched medium. There was no difference in sorus area between median and marginal discs. These results indicated that the marginal parts are able to produce sori, although the greater part of them are actually washed away before forming sori in the sea. The marginal part is probably in a state of `resource limitation' on the reproduction because the resource outflowed from this part to the basal part for the growth of sporophytes. This revised version was published online in August 2006 with corrections to the Cover Date.     

真卵形藻属(硅藻门)的中国新记录植物

该文报道了采自四川九寨沟自然保护区的2个真卵形藻属(硅藻门)中国新记录种,分别为高山真卵形藻[Eucocconeis alpestris(Brun)Lange-Bertalot]和平滑真卵形藻[Eucocconeis laevis(strup)Lange-Bertalot]。利用光学显微镜和扫描电子显微镜对2个种类进行细致的观察,并对其形态分类学特征和生境进行详细描述。     

芬兰贝氏藻(硅藻门)的超微结构研究

以采集于青海湖的石附生硅藻(芬兰贝氏藻)为样品,用10%的HCl溶液溶解样品中含钙的固体化合物,再加入30% H₂O₂溶液氧化去掉硅藻细胞的有机物质,获得干净的硅藻标本。利用Naphrax^ 封装剂制成永久装片,在光学和电子显微镜下对其形态特征进行了详细研究。结果表明：(1) 在光学显微镜下芬兰贝氏藻具有披针形的壳瓣外形、尖圆的壳面末端和明显伸长的中轴区三个特征。(2) 在扫描电子显微镜下芬兰贝氏藻紧邻中央区的孔纹明显伸长,其长度为2～4个正常孔纹的长度。(3) 通过文献查阅和相似种类的比较发现,同时具有尖圆的末端和明显伸长的紧邻中央区的孔纹是芬兰贝氏藻鉴定的独特性状组合,故芬兰贝氏藻的身份可由此确定。该研究首次报道了芬兰贝氏藻在中国的分布。     

Bud formation in the scyphozoan Cassiopea andromeda: epithelial dynamics and fate map

Cassiopea andromeda scyphistomae reproduce asexually by forming spindle shaped buds which, after detachment, metamorphose into polyps. Parent polyps appear to contribute to the buds' ecto- and endodermal epithelial cells, septal muscle cells, nematocytes and some zooxanthellae. Herein we describe bud morphogenesis, define 5 bud stages, and investigate the recruitment of bud ectoderm. India ink vital marking experiments reveal permanent apicobasal displacement of ectoderm. Labelled polyp cells are displaced towards and incorporated into the emerging bud. Ectoderm is recruited from all angular positions and cells labelled at increasing distances from the bud center are traced at increasingly more proximal positions on the buds. Unlike in Hydra attenuata, the recruitment area appears to be asymmetric since the zone contributing ectoderm from below is smaller than the zones above and lateral to the buds.     

Morphological investigations of the frustule, perizonium and initial valves of the freshwater diatom Achnanthes crenulata Grunow (Bacillariophyceae)

The present study clarifies the fine structure of the vegetative frustules, initial valves and perizonium of Achnanthes crenulata Grunow. The valves of the vegetative cell are distinctly linear‐lanceolate with an undulate margin. The valve face is quite flat and in girdle view is smoothly curved as in species of Gephyria (Bacillariophyceae). However, the valve face of the initial cells is slightly rounded and does not have an undulate margin. Furthermore, the rapheless sternum is centrally positioned along the apical axis of the araphid initial valve. As this taxon develops from auxospore to initial valve, it forms only longitudinal perizonial bands; no transverse bands arise. The perizonium consists of three silicified bands: one large, central longitudinal plate and two bands that underlie this plate; these two bands are either open or closed. This taxon has several conspicuous structures compared to other marine species of Achnanthes, but the structure of the perizonium supports the position of A. crenulata within Achnanthes sensu stricto.