A NOVEL ASSOCIATION BETWEEN AN ENDEMIC STICKLEBACK AND A PARASITIC DINOFLAGELLATE. 2. MORPHOLOGY AND LIFE CYCLE1

An unusual dinoflagellate has been discovered in association with an endemic population of stickleback, Gasterosteus (L.), from the Queen Charlotte Islands, Canada. The dinoflagellate spends most of its life cycle as a coccoid vegetative cyst, not as a parasitic trophont. The vegetative cyst is unique in containing a rigid fenestrated matrix, which is penetrated by cytoplasmic process that emanate from a central area containing the dinokaryotic nucleus and associated chloroplasts. Some pores in the matrix are filled by oil droplets or starch granules. Intracellular bacteria are found throughout the cyst, sometimes in association with the nucleus. The cytoplasm contains accumulation bodes, microbodies, polyhedral crystals, chloroplasts and polyvesicular bodes. The encysted dinoflagellate has several potential strategies. It can 1) shed its wall and become amoeboid; 2) undergo sporogenesis and give rise to both regular and resistant spores; 3) divide mitotically, with a gradual reduction in the size of daughter cells down to 20 μm; and 4) apparently form a resting cyst, during which it secretes a thick outer wall composed of five layers. Taxonomically, this unusual dinoflagellate appears to be a new member of the Blastodiniales, although its position will become clearer when details of the motile stage are known.     

CYST FORMATION IN THE FRESHWATER DINOFLAGELLATE CERATIUM HIRUNDINELLA (DINOPHYCEAE)1

Cyst formation in Ceratium hirundinella (O. F. Müll.) Bergh was studied by light and electron microscopy, using material from several lakes and reservoirs and also laboratory cultures. Cells preparing to encyst build up large quantities of starch and lipid and at the same time reduce their other cell components. The cyst is released from the theca as a naked cell bounded by a double membrane. The most commonly found cyst deposits a layer of electron-dense granules containing silicon on the outer membrane and lays down a cellulose-like material between the two membranes. Cysts without the electron-dense granules are commonly formed in cultures but rarely found in lakes. These cysts appear less resistant to decay and do not show the reorganization of cell contents for dormancy. It is suggested that C. hirundinella has both a resting cyst, forming part of the life cycle, and a temporary cyst stage.     

Life cycle of the ichthyotoxic dinoflagellate Cochlodinium polykrikoides in Korean coastal waters

Since 1995, blooms of the harmful dinoflagellate, Cochlodinium polykrikoides, have caused considerable mortality of aquatic organisms and economic loss in Korea. However, little is known about the life cycle of the species, except for the planktonic vegetative stage; therefore, the aim of this paper was to elucidate the life cycle of C. polykrikoides. Its life cycle has two morphologically different stages: an armored and an unarmored vegetative stage. Armored vegetative cells were found in seawater samples collected in late-November and developed into four-cell chained, unarmored vegetative cells under laboratory culture. In samples collected in late-May, both the armored and unarmored types (vegetative swimming stage) occurred; the former easily developed into an unarmored vegetative cell type, suggesting that the armoured–unarmored transition occurs as early as May. A presumptive resting cyst, round but folded at one side, was produced from armored type cells in laboratory conditions. It was also collected from natural bottom sediments, which suggests it is the dormant resting cyst of C. polykrikoides.     

Recruitment of Planktonic Dinoflagellates: Importance of Benthic Resting Stages and Resuspension Events

Many factors have ken put forward to account for the development of nuisance phytoplankton blooms in coastal zones. Usually hydrological factors as temperature or salinity stratification and adequate nutrient and trace metal availability are held responsible for the phenomenon. The most frequent causative organisms for nuisance blooms are dinoflagellates, many of which have a dormant stage (resting cyst) in their life cycle. The role of the complex life-strategies of these forms in initiating bloom formation is the focus of this study. Special attention is given to 25 different dinoflagellate resting cyst types isolated from recent German North Sea and Baltic Sea sediments, and their germination frequency under different environmental conditions. Also, the role of cyst resuspension in relationship to the timing, persistence and recurrence of dinoflagellate blooms is extensive discussed.     

TAXONOMIC STUDY OF BIGELOWIELLA LONGIFILA SP. NOV. (CHLORARACHNIOPHYTA) AND A TIME‐LAPSE VIDEO OBSERVATION OF THE UNIQUE MIGRATION OF AMOEBOID CELLS1

A new species of a chlorarachniophyte alga, Bigelowiella longifila sp. nov., is described. It is classified as a member of Bigelowiella as flagellate cells constitute the main stage of the life cycle. However, this alga is different from the only described species of the genus, B. natans Moestrup, in having a unique amoeboid stage in the life cycle. We observed an interesting behavior of amoeboid daughter cells after cell division: One of the two daughter cells inherits the long filopodium of the parental cell, and it subsequently transports its cell contents through the filopodium to develop at its opposite end. The other daughter cell forms a new filopodium. This unequal behavior of daughter cells may have evolved before the chlorarachniophytes and some colorless cercozoans diverged.     

Lotharella amoeboformis sp. nov.: A new species of chlorarachniophytes from Japan

The ultrastructure, morphology and life cycle of a new chlorarachniophyte alga collected from Okinawa in Japan have been studied. The life cycle of this alga consists of amoeboid, wall‐less round, coccoid and flagellated cells in culture condition; however, the coccoid and flagellate cells are very rare. The pyrenoid ultra‐structure of this alga is the same as that of a previously described species, Lotharella globosa. Since pyrenoid ultrastructure has been adopted as the main criterion for the generic classification of the chlorarachniophytes, the present alga is placed in Lotharella. However, the present alga has a dominant amoeboid cell stage and a reduced walled‐cell stage in the life cycle, while in L. globosa, the walled‐cell stage is dominant and there is no amoeboid cell stage. Therefore the present alga is described as a new species of Lotharella: Lotharella amoeboformis Ishida et Y. Hara sp. nov.     

The life cycle of the amoeboid alga Synchroma grande (Synchromophyceae, Heterokontophyta)--highly adapted yet equally equipped for rapid diversification in benthic habitats

Synchroma grande (Synchromophyceae, Heterokontophyta) is a marine amoeboid alga, which was isolated from a benthic habitat. This species has sessile cell stages (amoeboid cells with lorica and cysts) and non‐sessile cell stages (migrating and floating amoebae) during its life cycle. The different cell types and their transitions within the life cycle are described, as are their putative functions. Cell proliferation was observed only in cells attached to the substrate but not in free‐floating or migrating cells. We also characterised the phagotrophy of the meroplasmodium in comparison to other amoeboid algae and the formation of the lorica. The functional adaptations of S. grande during its life cycle were compared to the cell stages of other amoeboid algae of the red and green chloroplast lineages. S. grande was found to be highly adapted to the benthic habitat. One sexual and two asexual reproductive strategies (haplo‐diploid life cycle) support the ability of this species to achieve rapid diversification and high adaptivity in its natural habitat.     

细弱绒泡菌的生活史

利用燕麦-琼脂培养、基物培养及扫描电镜技术研究了细弱绒泡菌的个体发育过程,在燕麦琼脂培养基上完成了从孢子到孢子的生活史。结果表明,细弱绒泡菌生活史包括单核的黏变形体或游动胞、多核的营养体原质团以及孢子形成阶段。孢子球形,表面具细小疣点。孢子萌发为裂式,释放1黏变形体。黏变形体行变形运动,在有水的条件下,可转变为游动胞。成熟原质团橘黄色。原质团类型为显型,具有扇形网络状菌脉。成熟原质团可形成多个孢囊。琼脂培养基上获得的细弱绒泡菌孢子与野生型相似,并具有可育性。     

黄柄钙皮菌的生活史

利用基物培养、燕麦-琼脂培养技术及扫描电镜技术研究了黄柄钙皮菌的个体发育过程,在燕麦琼脂培养基上完成了从孢子到孢子的生活史。结果表明,生活史包括单核的黏变形体或游动胞、多核的营养体原质团以及孢子形成阶段。孢子球形,表面具疣突。孢子萌发为裂式,释放1黏变形体。黏变形体行变形运动,在有水的条件下,可转变为游动胞并游动。可观察到1长具极性的鞭毛。合子形成原质团。成熟原质团棕色。原质团类型为显型,具有扇形网络状菌脉。琼脂培养基上获得的黄柄钙皮菌孢子与野生型相似,并具有可育性。     

The Fine Structure of the Phoront of the Apostomatous Ciliate,Hyalophysa chattoni*

The phoront of the apostomatous ciliate, Hyalophysa chattoni, is an encysted stage that is carried on the exoskeleton of its crustacean host until the ecdysis of the host. At molting the phoront rapidly metamorphoses to the feeding stage, excysts, and immediately begins to feed on exuvial fluid trapped in the cast-off exoskeleton. The fine structure of the resting phoront resembles that of the preceding migratory stage, the tomite. A prominent ventral tuft of cilia, the ogival field, has vanished, and the trichocysts that paralleled the kinetics have disappeared. The dense inclusion bodies that were concentrated around the mouth and falciform fields have dispersed and greatly decreased in number. The cytoplasm and its membranous organelles do not appear visibly condensed or altered from the preceding stage in the life cycle. The phoront is merely quiescent instead of dormant. Unlike the few ciliate cysts previously examined by electron microscopy, the phoront's cyst is not divisible into separable layers. It resembles the loricae of certain suctoria in being formed principally of a fibrous substance, the outer surface of which has a paracrystalline pattern. The peduncle attaching the cyst to the crab's gill is a continuation of the cyst wall although its structure is somewhat modified. The most conspicuous innovation in the phoront's fine structure is the massive tracts of microtubules that run longitudinally through the macronucleus. The microtubules are in intimate contact with Feulgen-positive chromatin masses which are crowded toward the periphery of the macronucleus.     

OBSERVATIONS ON “VAMPYRELLA PENULA-STYLODINIUM SPHAERA” AND THE ULTRASTRUCTURE OF THE REPRODUCTIVE CYST

“Vampyrella-Stylodinium,” an artificial name for a predaceous organism of uncertain taxonomic position, has at least three distinct phases in its life history: the amoeboid phase, both free-floating and attached; the feeding cyst or immobile phase; and flagellated gymnodinoid swarmers. The orange free-floating amoeba has unbranched, filose pseudopodia and several contractile vacuoles. When feeding on the filamentous green alga Oedogonium, the pseudopodia shorten and rearrange. After dissolution of part of the Oedogonium cell wall, the amoeba ingests the host protoplast. Then a stalked reproductive cyst may form. This cyst changes color from green to light orange as it matures. At the time of excystment, the cyst has a smooth outer wall, a spinose inner wall, and a well-delineated phagocytic vacuole. As this vacuole moves from its central position to the cyst's periphery, the walls rupture and 2-4 amoebulae emerge. With TEM observations, the reproductive cyst is shown to be multinucleate. Each nucleus is eukaryotic in organization and possesses one nucleolus. Mitochondria have tubular cristae and no structures unique to the division Pyrrhophyta are observed. Although this stage of the life history does not have a dinokaryotic nucleus, the gymnodinoid swarmers that can emerge from the reproductive cyst, do. Like other parasites which have been assigned to the division Pyrrhophyta, “Vampyrella-Stylodinium” does not conform well to the generalized concept of a dinoflagellate.     

海洋浮游纤毛虫包囊的研究进展*

作为微型浮游动物的重要组成部分,海洋浮游纤毛虫是连接微食物环和经典食物链的重要中介。有些浮游纤毛虫在生活周期中会形成包囊,条件适宜时包囊会萌发,这对纤毛虫种群动态有重要的意义。目前国际上对于浮游纤毛虫生态学的研究主要集中在其营养期细胞,浮游纤毛虫包囊的研究尚少,中国还没有这方面的研究。本文对浮游纤毛虫包囊研究进展进行概述,包括包囊的形态、沉积物中包囊的丰度、包囊形成的环境因素、包囊萌发过程及环境因素对包囊萌发的影响等方面,希望对国内开展浮游纤毛虫包囊的研究有所裨益。     

Generalist Life Cycle Aids Persistence of Alexandrium ostenfeldii (Dinophyceae) in Seasonal Coastal Habitats of the Baltic Sea

In seasonal environments, strong gradients of environmental parameters can shape life cycles of phytoplankton. Depending on the rate of environmental fluctuation, specialist or generalist strategies may be favored, potentially affecting life cycle transitions. The present study examined life cycle transitions of the toxin producing Baltic dinoflagellate Alexandrium ostenfeldii and their regulation by environmental factors (temperature and nutrients). This investigation aimed to determine whether genetic recombination of different strains is required for resting cyst formation and whether newly formed cysts are dormant. Field data (temperature and salinity) and sediment surface samples were collected from a site with recurrent blooms and germination and encystment experiments were conducted under controlled laboratory conditions. Results indicate a lack of seasonal germination pattern, set by an endogenous rhythm, as commonly found with other dinoflagellates from the Baltic Sea. Germination of quiescent cysts was triggered by temperatures exceeding 10°C and combined nutrient limitation of nitrogen and phosphorus or a drop in temperature from 16 to 10°C triggered encystment most efficiently. Genetic recombination was not mandatory for the formation of resting cysts, but supported higher numbers of resistant cysts and enhanced germination capacity after a resting period. Findings from this study confirm that A. ostenfeldii follows a generalist germination and cyst formation strategy, driven by strong seasonality, which may support its persistence and possibly expansion in marginal environments in the future, if higher temperatures facilitate a longer growth season.     

针箍菌的生活史

利用基物和燕麦-琼脂技术研究了针箍菌的个体发育过程,在燕麦琼脂培养基上完成了从孢子到孢子的生活史。结果表明,生活史包括单核的黏变形体或游动胞、多核的营养体原质团以及孢子形成阶段。琼脂培养基获得的针箍菌孢子与野生型相似,并具有可育性。     

The toxic dinoflagellate Cochlodinium polykrikoides (Dinophyceae) produces resting cysts

While harmful algal blooms (HABs) caused by the toxic dinoflagellate Cochlodinium polykrikoides have been known to science for more than a century, the past two decades have witnessed an extraordinary expansion of these events across Asia, North America, and even Europe. Although the production of resting cysts and subsequent transport via ships’ ballast water or/and the transfer of shellfish stocks could facilitate this expansion, confirmative evidence for cyst production by C. polykrikoides is not available. Here, we provide visual confirmation of the production of resting cysts by C. polykrikoides in laboratory cultures isolated from North America. Evidence includes sexually mating cell pairs, planozygotes with two longitudinal flagella, formation of both pellicular (temporary) cysts and resting cysts, and a time series of the cyst germination process. Resting cyst germination occurred up to 1 month after cyst formation and 2–40% of resting cysts were successfully germinated in cultures maintained at 18–21 °C. Pellicular cysts with hyaline membranes were generally larger than resting cysts, displayed discernable cingulum and/or sulcus, and reverted to vegetative cells within 24 h to ∼1 week of formation. A putative armored stage of C. polykrikoides was not observed during any life cycle stage in this study. This definitive evidence of resting cyst production by C. polykrikoides provides a mechanism to account for the recurrence of annual blooms in given locales as well as the global expansion of C. polykrikoides blooms during the past two decades.     

圈绒泡菌的生活史

利用基物培养、燕麦-琼脂培养技术及扫描电镜技术研究了圈绒泡菌的个体发育过程, 在燕麦琼脂培养基上完成了从孢子到孢子的生活史。结果表明,生活史包括单核的黏变形体或游动胞、多核的营养体原质团以及孢子形成阶段。琼脂培养基上获得的圈绒泡菌孢子与野生型相似,并具有可育性。     

淡黄绒泡菌和全白绒泡菌原生质流向的观察

显型原质团是绒泡菌目黏菌的营养生长阶段,其最明显的现象是往返原生质流,但一直并不清楚原生质流反向流动的原因。观察研究了淡黄绒泡菌和全白绒泡菌原质团中的原生质流,结果表明：由于菌脉中堵塞或是在原质团前缘尚未分化通道引起反向原生质流,从而引起原质团多方向生长使原质团前缘呈现扇面状。原生质流总的方向是扇面端,并完成原质团运动。     

扁绒泡菌的生活史

黏菌的生活史对于研究其营养方式的多样性以及系统发育等具有重要价值,目前国内外有关黏菌生活史的报导很少。利用燕麦-琼脂培养、基物培养及扫描电镜技术研究了扁绒泡菌的个体发育过程,在燕麦琼脂培养基上完成了从孢子到孢子的生活史。结果表明,扁绒泡菌生活史包括单核的黏变形体或游动胞、多核的营养体原质团以及孢子形成阶段。孢子球形,表面具细小疣点。孢子萌发为裂式,释放1黏变形体。原质团类型为显型。成熟原质团乳白色,可形成多个孢囊。琼脂培养基上获得的扁绒泡菌孢子与野生型相似,并具有可育性。