Auxospore Development Research Articles

Sexual reproduction and auxospore development in the diatom Biddulphia biddulphiana.

Phylogenetic relationships among mediophycean diatoms with elliptical valve outline and elevated apices have long been a subject of interest and debate, particularly with respect to their relationship to pennates. However, results remain inconclusive, whether based on vegetative valve morphology, reproduction, or molecular phylogenetic data. Searching for phylogenetically informative features, we re-examined sexual reproduction, auxospore structure and development in the diatom Biddulphia biddulphiana. Several unique or unusual features and processes characterized its sexual reproduction. A unique spermatogenesis occurs with premeiotic separation of an anucleate protoplast containing all chloroplasts and likely other organelles. Additionally, their auxospore walls are some of the most complex documented, retaining earlier deposited layers that obscure layers formed during later stages of development. The oldest layer consists of thick, mostly organic incunabulum, underlain by outer and inner epizonia and finally transverse (TP) and longitudinal (LP) perizonia. The complexity of the fine structure of these layers is unprecedented. The orientation of some TP bands is also unique in mediophytes, with some perpendicular to the auxospore apical axis, parallel to each other, and open with aligned ends, as typically seen in pennates. The TP also contains rings slanting toward the apices, as in some other mediophytes, e.g., eupodiscaceans. However, both eupodiscaceans and biddulphiaceans show perizonial band structure derived from anastomosing radial scales, thus termed “scaly bands”. Pinnate TP bands, common among pennate auxospores, were not found. Thus B. biddulphiana auxospore wall structure contains a mixture of characters specific to this clone but also known from mediophytes and araphid pennates. However, these features do not provide unequivocal evidence that this or the other Biddulphia species examined to date are the closest extant relatives of basal araphid pennates.

Enlarge or die! An auxospore perspective on diatom diversification

Sexual reproductive cells and processes are highly conserved in a wide range of organisms. A specialized, expandable zygote called the auxospore is a cell known only in diatoms. Auxospores perform two critical functions in the life cycle of most diatoms: (1) as in other sexual organisms, they confer the benefits of genetic recombination to diatom progeny, and (2) restore larger cell size and allow a new cycle of vegetative cell size diminution to commence. Because of this dual importance in the diatom life cycle, the auxospore is strongly conserved, in contrast to the astounding diversity of forms among diatom vegetative cells. The auxospore is a complex cell which develops in a specific sequence of stages with a variety of characteristic structures recognizable across all diatoms. With only two major blueprints for auxospore growth and development (isodiametric and anisodiametric) and a number of highly conserved associated structures, studying this aspect of the diatom life cycle may reveal much about their deep evolutionary history. In the evolutionarily more recent arrivals (i.e., pennates), additional reproductive characters may be informative, e.g., mode of gamete motility, anisogamy vs. isogamy. We argue that combined with the widely varying but taxon-specific aspects of diatom gametogenetic processes, auxospores are excellent tools for cross group comparisons that provide insight into their evolutionary relationships. Expanding our dataset from 21 genera presented in 2004 to 65 polar centrics, nonpolar centrics, and araphid pennates, we repostulate that the evolutionarily informative capacity of auxospore development stands on its own.

Auxosporulation in Biddulphia tridens (Ehrenberg) Ehrenberg (Mediophyceae, Bacillariophyta)

ABSTRACT We documented male and female gametogenesis, auxospore structure and development of the biddulphioid diatom species Biddulphia tridens (Ehrenberg) Ehrenberg, a close relative of the generitype. Our mating experiments showed that the B. tridens clones were homothallic, self-fertile and produced two eggs per oogonium. During spermatogenesis, spermatogonia developed by depauperating mitosis from spermatogonangium cells and four hologenous uniflagellate male gametes were produced per spermatocyte. In the early stage of auxospore development, cells were globular to sub-globular in shape and covered with a siliceous envelope. The incunabular scales on these auxospores showed a great diversity in size and shape. In the later stages of development, auxospores grew anisodiametrically, facilitated by transverse perizonial bands. Initial epivalve formation was preceded by the retraction of the protoplast from the auxospore wall. The structural features of sex cells such as incunabular scales found in B. tridens are distinct from other broadly defined biddulphioid diatoms thus far examined. This is consistent with the distant relationship between true Biddulphia species and the Biddulphia-like diatoms indicated by molecular phylogenies. Morphological and molecular examination of B. tridens clones were performed here only to confirm species identity. This complete reproductive dataset from a true Biddulphia species will provide valuable insights for future studies of the relationships between Biddulphia-like diatoms and true Biddulphia species.

Auxosporulation in Chaetoceros acadianus sp. nov. (Bacillariophyceae), a new member of the Section Compressa

ABSTRACTWe document the fine structure of auxospores in a Chaetoceros species isolated from the Acadian coast of New Brunswick, Canada. Auxospore development in this species occurs in a terminal rather than lateral position, a characteristic never before observed in this genus. Our observations suggest that auxosporulation was uniparental, probably an extreme form of autogamy with sister nuclei fusing following meiosis II. Mature auxospores were adze-shaped to sub-globular and contained both scales and transverse perizonia in their walls. The transverse perizonial band structure was similar to longitudinal perizonial bands found in other species of Chaetoceros and differed from the pinnate bands of pennate transverse perizonia, which consisted of a central rib and bilateral fimbria. Instead, the band structure in C. acadianus was more similar to unilateral fimbriate bands in cymatosiroids. We also propose that our diatom represents a species new to science and is a member of the Chaetoceros Section Compressa. We provide its morphological, molecular and reproductive characterization.

Sexual reproduction in plagiogrammacean diatoms: First insights into the early pennates.

The genera Plagiogramma and Dimeregramma are members of a small, but evolutionarily important group of diatoms, the "basal" araphids. They are sister to all other pennates, both araphid and raphid taxa. Thus, their phylogenetic position carries the potential for providing insights into the earliest pennates. We documented sexual reproduction, mating system and sex cell development in the first members of the "basal" araphid clade ever investigated. The mating system in all these species involved heterothally. It was, however, more complex in P. tsawwassen, where in addition to heterothallic clones, intraclonal and polysexual clones also exist. Auxospore development and wall structure was similar in all three species and demonstrated several characters also reported from "core" araphids. Of these, vigorous, pseudopodial motility of male secondary spermatocytes and gametes was most notable because it indicates that this character was likely present in the last common ancestor of all the pennates. Pseudopodial motility of the male sex cells might have afforded sufficient compensation and/or benefits to the emerging pennates for replacing flagellated sperm, present in centrics. The characters thus far uniquely present among our plagiogrammaceans but not reported from other pennates were: the "gametic" fusion between sex-compatible secondary spermatocytes, in some cases before completion of Meiosis II in males, transverse perizonial bands produced all together or in quick succession rather than being added to the auxospore apex one at a time, and expanding auxospores with 3–4 nuclei. An initial epivalve, similar in morphology to what in some diatoms had been interpreted as a “longitudinal” perizonium, may be more widespread among pennates than thus far appreciated. In addition, we discovered two species new to science (D. acutumontgo, P. tsawwassen), and refined delineation of P. staurophorum by including metric data from the original material.

Gametogenesis and Auxospore Development in the Bipolar Centric Diatom Brockmanniella brockmannii (Family Cymatosiraceae)

We examined gametogenesis and auxospore development in the cymatosiroid diatom Brockmanniella brockmannii. Our mating experiments demonstrated that the clones were homothallic, self-compatible, produced two eggs per oogonium and four hologenous uniflagellate male gametes per spermatogonium. Auxospores grew free in the culture medium and the most external layer of the auxospore wall was made up of spinescent scaly elements in their early globular stage of development. The anisodiametric expansion of the globular auxospore was rather limited, at best proceeding only to a sub-globular stage. Our data suggest that the initial epivalves, nearly pennate-like in outline, developed within such auxospores curled up against the ventral side of the auxospore wall, not across its equatorial plane, and apparently without the help of rings, hoops or bands characteristic of pennate-type transverse perizonia. Thus, B. brockmannii showed a mode of initial epivalve development thus far unknown among diatoms. We propose that progressive silicification of the initial epivalve facilitated the breakdown of the sub-globular auxospore dorsal wall to accommodate for its straightening. The structural features of the spinescent scales on the auxospore wall suggest that Cymatosiraceae could be related to the mediophycean family Eupodiscaceae, although most current molecular phylogenies do not support a direct relationship.

Auxosporulation in Paralia guyana MacGillivary (Bacillariophyta) and Possible New Insights into the Habit of the Earliest Diatoms.

BackgroundDiatoms are one of the most ecologically important aquatic micro-eukaryotes. As a group unambiguously recognized as diatoms, they seem to have appeared relatively recently with a limited record of putative remains from oldest sediments. In contrast, molecular clock estimates for the earliest possible emergence of diatoms suggest a considerably older date. Depending on the analysis, Paralia and Leptocylindrus have been recovered within the basal molecular divergences of diatoms. Thus these genera may be in the position to inform on characters that the earliest diatoms possessed.FindingsHere we present auxospore development and structure of initial and post-auxospore cells in a representative of the ancient non-polar centric genus Paralia. Their initial frustules showed unusual, but not unprecedented, spore-like morphology. Similarly, initial frustules of Leptocylindrus have been long considered resting spores and a unique peculiarity of this genus. However, even though spore-like in appearance, initial cells of Paralia readily resumed mitotic divisions. In addition, Paralia post-auxospore cells underwent several rounds of mitoses in a multi-step process of building a typical, “perfect” vegetative valve. This degree of heteromorphy immediately post-auxosporulation is thus far unknown among the diatoms.ImplicationsA spore-related origin of diatoms has already been considered, most recently in the form of the “multiplate diploid cyst” hypothesis. Our discovery that the initial cells in some of the most ancient diatom lineages are structurally spore-like is consistent with that hypothesis because the earliest diatoms may be expected to look somewhat similar to their ancestors. We speculate that because the earliest diatoms may have appeared less diatom-like and more spore-like, they could have gone unrecognized as such in the Triassic/Jurassic sediments. If correct, diatoms may indeed be much older than the fossil record indicates, and possibly more in line with some molecular clock predictions.

Spermatogenesis and auxospore structure in the multipolar centric diatom Hydrosera.

Spermatogenesis and auxospore development were studied in the freshwater centric diatom Hydrosera triquetra. Spermatogenesis was unusual, lacking depauperating cell divisions within the spermatogonangium. Instead, a series of mitoses occurred within an undivided cell to produce a multinucleate plasmodium with peripheral nuclei, which then underwent meiosis. 32 or 64 sperm budded off from the plasmodium leaving a large residual cell containing all the chloroplasts. Similar development apparently occurs in Pleurosira, Aulacodiscus, and Guinardia, these being so distantly related that independent evolution of plasmodial spermatogenesis seems likely. After presumed fertilization, the Hydrosera egg cell expanded distally to form a triangular end part. However, unlike in other triangular diatoms (Lithodesmium, Triceratium), the development of triradiate symmetry was not controlled by the "canonical" method of a perizonium that constrains expansion to small terminal areas of the auxospore wall. Instead, the auxospore wall lacked a perizonium and possessed only scales and a dense mat of thin, apparently entangled strips of imperforate silica. No such structures have been reported from any other centric diatoms, the closest analogs being instead the incunabular strips of some raphid diatoms (Nitzschia and Pinnularia). Whether these silica structures are formed by the normal method (intracellular deposition within a silica deposition vesicle) is unknown. As well as being more rounded than vegetative cells, the initial cell is aberrant in its structure, since it has a less polarized distribution of the "triptych" pores characteristic of the species.

Sexual reproduction and auxospore structure in Diploneis papula (Bacillariophyta)

Idei M., Sato S., Watanabe T., Nagumo T. and Mann D.G. 2013. Sexual reproduction and auxospore structure in Diploneis papula (Bacillariophyta). Phycologia 52: 295–308. DOI: 10.2216/12–021.1We give a detailed account of sexual reproduction and auxospore development in the diatom genus Diploneis, principally from clonal cultures of the marine benthic Diploneis papula. Sexual reproduction of D. papula was apparently homothallic. After pairing side to side, cells entered meiosis, and each gametangium produced two gametes. Fertilization was physiologically anisogamous, and both gametes of one gametangium were active and those of the other gametangium passive so that the two zygotes were formed within or close to the ‘passive’ gametangium. Each gamete contained a single chloroplast. The zygote became surrounded by delicate incunabula that contained circular or elliptical scales; this confirmed recent observations that scales (or apparently homologous strips or plates) are quite commonly formed by the zygotes of raphid diatoms. Subsequently, a robust transverse perizonium was built up as the auxospore expanded, and the perizonium was comprised of a closed primary band at the centre and open secondary bands towards each pole. The ends of the secondary bands, which met along one side of the auxospore to form a suture as in other pennate diatoms, were curved inwards towards the centre of the auxospore. Novel features were the extension of the transverse perizonium to cover the tips of the auxospores (in other diatoms it stopped short of the poles, leaving an apical dome covered only by the incunabula) and a differentiation of the ends of the transverse bands into truncated and prolonged variants, which alternated along the suture. The longitudinal perizonium possessed the same highly conserved configuration as in other pennate diatoms, with a wide bifacial central band flanked by two other bands, which here differed in shape and structure. All the perizonial bands were fimbriate. The possible homology of the longitudinal perizonium to the thecae of vegetative cells was discussed. During formation of the initial cell, the protoplast contracted away from the perizonium, but the latter nevertheless moulded the outline shape of the initial valves, which were more strongly constricted than preauxospore cells and gametangia. Other Diploneis species produced either one or two gametes per gametangium.

Gametogenesis and Auxospore Development in Actinocyclus (Bacillariophyta)

cGametogenesis and auxospore development have been studied in detail in surprisingly few centric diatoms. We studied the development of sperm, eggs and auxospores in Actinocyclus sp., a radially symmetrical freshwater diatom collected from Japan, using LM and electron microscopy of living cultures and thin sections. Actinocyclus represents a deep branch of the ‘radial centric’ diatoms and should therefore contribute useful insights into the evolution of sexual reproduction in diatoms. Spermatogenesis was examined by LM and SEM and involved the formation of two spermatogonia (sperm mother-cells) in each spermatogonangium through an equal mitotic division. The spermatogonia produced a reduced ‘lid’ valve, resembling a large flat scale with irregular radial thickenings. Sperm formation was merogenous, producing four sperm per spermatogonium, which were released by dehiscence of the ‘lid’ valve. The sperm were spindle-shaped with numerous surface globules and, as usual for diatoms, the single anterior flagellum bore mastigonemes. One egg cell was produced per oogonium. Immature eggs produced a thin layer of circular silica scales before fertilization, while the eggs were still contained within the oogonium. Sperm were attracted in large numbers to each egg and were apparently able to contact the egg surface via a gap formed between the long hypotheca and shorter epitheca of the oogonium and a small underlying hole in the scale-case. Auxospores expanded isodiametrically and many new scales were added to its envelope during expansion. Finally, new slightly-domed initial valves were produced at right angles to the oogonium axis, after a strong contraction of the cell away from the auxospore wall. At different stages, Golgi bodies were associated with chloroplasts or mitochondria, contrasting with the constancy of Golgi–ER–mitochondrion (G-ER-M) units in some other centric diatoms, which has been suggested to have phylogenetic significance. Electron-dense bodies in the vacuole of Actinocyclus are probably acidocalcisomes containing polyphosphate.

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.

Sexual reproduction, mating system, chloroplast dynamics and abrupt cell size reduction in Pseudo-nitzschia pungens from the North Sea (Bacillariophyta)

Clonal cultures of Pseudo-nitzschia pungens were isolated at various times from seven sites in the North Sea. During the mitotic cell cycle, the two plate-shaped chloroplasts were girdle-appressed during interphase and mitosis. After cytokinesis, the chloroplasts moved onto the parental valve and remained there during the formation of the new hypovalve and until separation and re-arrangement of the sibling cells within the cell chain had been completed. Clones were almost always heterothallic and cultures of opposite mating type isolated from different localities were compatible. Meiosis I was cytokinetic and accompanied by chloroplast division. Meiosis II involved karyokinesis but not cytokinesis and preceded the rearrangement and contraction of the two gametes. Sexual reproduction involved physiological anisogamy. With one exception, gamete behaviour was clone-specific, gametes being active in clones of one mating type but passive in clones of the other mating type. Auxospore development was accompanied by deposition of a transverse and then a longitudinal perizonium. Infrequently, triploid auxospores and presumably haploid auxospores were produced. The four chloroplasts of diploid auxospores did not divide, and behaved synchronously during the two acytokinetic mitotic cycles accompanying the deposition of the initial thecae. Just before the first division of the initial cell, the chloroplasts shifted onto the valves (two per valve). The division of the initial cell was not accompanied by chloroplast division and so the two daughter cells received two chloroplasts each. Two modes of abrupt cell size reduction were detected. One occurred during initial cell formation when part of the expanded auxospore aborted. The other pattern was more gradual and was observed in growing cultures; during successive cell divisions a frustule constriction appeared and intensified, one chloroplast split into two, and part of the protoplast aborted. A simple naming system is proposed for mating types in pennate diatoms.

Fine-structure of the vegetative frustule, perizonium and initial valve of Achnanthes yaquinensis (Bacillariophyta)

This study clarifies the fine structure of the vegetative and initial valves of Achnanthes yaquinensis and briefly compares them to other Achnanthes species. It also elucidates the structure of the perizonium, based on auxospore development in short-term cultures. The araphid valve has marginal ridges and terminal spines that allow connecting valves to form a chain. The terminal spines develop from the rapheless sternum. The complete cingulum consists of 3–5 split bands with two rows of areolae. These features can be used to discriminate species within the genus. Sexual reproduction is isogamous with two mother cells producing two auxospores, which are enclosed in mucilage. The perizonium develops on one side of an auxospore only, comprising one large central longitudinal band and four closed bands. There are no transverse perizonial bands. The raphid valve of the initial cell forms first, underneath the longitudinal perizonium, followed by the araphid valve, which is not covered by any perizonial bands. The araphid valve of the initial cells lacks a marginal spine, and the rapheless sternum lies more centrally than in the vegetative cell. The relationship of the genus Achnanthes to other monoraphid diatoms is discussed briefly.

Sexual behaviour, auxosporulation and chloroplast dynamics in a marine Amphora (Bacillariophyceae) studied in culture

Chloroplast, mating behaviour, and auxosporulation are described for Amphora cf. proteus Gregory. The interphase chloroplast is butterfly-shaped and lies with its central part, containing a flat discoid pyrenoid, pressed against the ventral side of the girdle. Chloroplast division is longitudinal and begins and ends before mitosis and cytokinesis. Over the cell cycle, no rotations or lateral translocations of the chloroplast were noticed. A. cf. proteus is the first species of this genus in which sexual reproduction and breeding behaviour have been studied in clonal cultures. The cells isolated exhibited strictly dioecious behaviour, reproducing only in mixed cultures of sexually compatible clones. Sexual reproduction in A. cf. proteus is morphologically and behaviourally isogamous. Each gametangium forms two functional gametes, which do not rearrange within the gametangium, and two small aborted cells. The aborted cells are produced after an unequal cytokinesis accompanying meiosis II; the second cytokinesis during gametogenesis represents a phenomenon hitherto unknown in pennate diatoms. Auxospore development takes place by bipolar expansion. The expansion is unequal on the two long sides of the auxospore, so that the auxospore becomes curved and dorsiventral. The auxospore forms a transverse and longitudinal perizonium.

TRANSITION TO SEXUAL REPRODUCTION AND CONTROL OF INITIAL CELL SIZE IN NITZSCHIA LANCEOLATA

Induction of gametogenesis in some diatoms occurs after a change in illumination. The amount of light energy supplied and the pattern of supply not only condition the physiological status of the cells, but can also switch the mitotic cycle into meiosis. Previous studies have dealt with monoecious species. Here, the transition to sexual reproduction was studied in the dioeciously reproducing species Nitzschia lanceolata, where a compatible partner must be present for vigorous auxosporulation. Successful completion of auxospore development is ultimately dependent on light energy captured through photosynthesis, although sufficient storage material can be accumulated before sexual interaction or the shift to meiosis to allow initial cells to be formed without any further supply of energy. However, if the gametes lack sufficient stored energy then auxospore development ceases. The fact that the cells can exit from the mitotic cycle in the beginning of the Gl phase indicates that possession of a particular material stock is not essential for sexual induction. Factors inducing gametogenesis do not influence the growth of the auxospores or initial cell formation. In N. lanceolata, the lengths of the initial cells are correlated with the lengths of the gametangia. However, for a given parent pair, initial cell size is ± constant and appears to conform to the principle “all or nothing”.

Sexual reproduction and systematics of Placoneis (Bacillariophyta)

Sexual reproduction and protoplast structure are used to check the classification of the naviculoid genus Placoneis Mereschkowsky (Bacillariophyta). The gametangia are closely associated within a robust mucilage capsule throughout auxosporulation, and pairing is ± random with respect to asymmetries of the frustule and protoplast. Two gametes are produced per gametangium, which are morphologically identical but differentiated into active and passive (physiological anisogamy). The supernumerary nuclei from meiosis II begin to degenerate soon after telophase II, but can still be detected in the initial cells. The gametes become rearranged within the gametangia after meiosis II and plasmogamy takes place via a single, simple aperture. The auxospores expand parallel to the apical axes of the gametangia.The classification of Placoneis in the Cymbellales, as the sister group to the cymbelloid and gomphonemoid diatoms, is supported by valve and protoplast structure, the method ofchloroplast division, and by sexual reproduction and auxospore development. Placoneis seems to occupy an intermediate position in the Cymbellales between the cymbelloid-gomphonemoid cluster and more ancient lineages that include Anomoeoneis Pfitzer, Dickieia Berkeley ex Kiitzing, Rhoicosphenia Grunow and Campylopyxis Medlin. No features can be found that are common to Placoneis and Navicula Bory (from which Placoneis was separated) except for the bilateral symmetry of the valve and the central raphe system, which would be synapomorphies only at a much higher taxonomic level.

SEXUAL REPRODUCTION IN A MARINE MEMBER OF THE BACILLARIACEAE

Sexual reproduction and auxospore development are described for a marine species of Bacillariaceae related to Nitzschia reversa. Oogamy, recently claimed to occur in N. pungens f. multiseries, was not found. No copulation envelope is produced and reproduction is isogamous. The orientations of the auxospores and gametangia bear no fixed relation to each other. A similar mode of auxosporulation occurs in some other species of Bacillariaceae, such as N.fonticola, N. angustata and apparently also N. subcurvata. Certain features are also shared by N. longissima, which shares a similar cell shape, with long rostrate apices. Two of the four plastids present in the zygotes and auxospores are often contracted and apparently degenerate. Oogamy is unlikely to occur in any members of the Bacillariaceae, though some slight doubt lingers; an old report (by Pascher), of the formation of four protoplasts (gametes?) within frustules of Nitzschia cf. hantzschiana, requires reinvestigation. The nature of the caps that cover the ends of the auxospores in many pennate diatoms is discussed.

VALVE DEVELOPMENT INSTEPHANODISCUS HANTZSCHIIGRUNOW (BACILLARIOPHYCEAE) AND ITS IMPLICATIONS ON SPECIES IDENTIFICATION

Clonal isolates of Stephanodiscus hantzschii Grunow were grown in batch culture and growth rates were measured during the exponential growth phase. Clones differing in size grew at significantly different rates at 20°C but not at 12°C. The reduction in cell size (diameter) over a period of time was variable within and among clones at 12°C over the course of one year. Mean concentration of Si/cell (pg Si/cell) was related to mean cell size of the respective clones. Valve morphology in older clones at 12°C and two grown at 20°C showed visual signs of decreased silicification, typical of the morphotype referred to as S. hantzschii fo. tenuis (Hust.) Håkansson & Stoermer. Occluded valve types diagnostic of Pseudo-stephanodiscus perforatus Sieminska (Genkal & Kusmin) appeared at the onset of the exponential growth phase. The rimoportula was the final process to develop during frustule formation. Auxospore development showed conclusively that forms hantzschii, tenuis and P. perforatus are just different phases in the growth cycle of the species S. hantzschii Grunow. An emended description of the species should include all the morphotypes in the life cycle.

Ultrastructure of the centric diatom, Cyclotella meneghiniana: vegetative cell and auxospore development

Abstract In Cyelotella meneghiniana the location of the nucleus in the vegetative cell changes as the cell diameter is decreased by successive cell divisions. Other features of the vegetative cell are ultrastructurally similar to other diatoms: chloroplasts are enveloped by chloroplast endoplasmic reticulum; pyrenoids are traversed by a single thylakoid; mitochondria contain tubular cristae; and dictyosomes are located near the nucleus. The auxospore has one or more large vacuoles that restrict the organelles to a narrow, peripheral band of cytoplasm. The organelles of the auxospore are morphologically similar to those found in the vegetative cells. Peripheral endoplasmic reticulum is present in the auxospore but not in the vegetative cell. The wall of the immature auxospore is composed of an organic component that is supplemented with siliceous scales as the auxospore matures. A metagamic mitosis precedes formation of each of the valves of the initial cell. Both valves of the initial cell are formed insi...