Each day, some planozygotes fall to the sediments as cysts, but new planozygotes appear following gamete fusion. The concept of a discrete seedbed may not be appropriate in some locations, however, due to the widespread, dispersed distribution of some cysts and the likelihood that germination will occur over a large area. Similar findings have been obtained with A. tamiyavanichi and A. cohorticula. The appearance of conjugating gametes and formation of larger and biflagellate planozygotes is generally obtained by transferring vegetative cells into diluted N- or P-deprived culture medium (e.g., Anderson and Lindquist, 1985). For example, cyst mapping within the Nauset Marsh System on Cape Cod revealed three highly localized seedbeds at the extreme ends of the complex network of channels and salt ponds that comprise that system (Crespo et al., in press.). Flynn K, Jones KJ, Flynn KJ. compressum. Stken A, Orr RJS, Kellmann R, Murray SA, Neilan BA, Jakobsen KS. Examples of cyst seedbeds in deeper coastal waters are less common, perhaps due to the expense and difficulty of large-scale mapping, but some large studies have been conducted, revealing accumulations stretching hundreds of km along the shore, and 50 km or more offshore, such as those for A. fundyense in the Gulf of Maine (e.g., Anderson et al., 2005c). Monroe EA, Van Dolah FM. Nevertheless, in situ studies provide the necessary validation of the experimental approach and are in turn source of new questions for experimental work. Iron is a likely candidate for a stimulatory micronutrient, as Wells et al. Oshima Y, Hirota M, Yasumoto T, Hallegraeff GM, Blackburn SI, and Steffensen DA (1989) Production of paralytic shellfish toxins by the dinoflagellate Alexandrium minutum Halim from Australia. Light and nutrient limitation in, MacKenzie L, de Salas M, Adamson J, Beuzenberg V. The dinoflagellate genus, MacKenzie L, White D, Oshima Y, Kapa J. Half-saturation constants range from 0.01 to 2.6 M, and were related to growth rate in A. catenella (Jauzein et al., 2010). However, allelopathic activity is not ubiquitous among Alexandrium populations or universally effective against all potential targets in natural plankton assemblages; even in extremely dense Alexandrium blooms grazing by tintinnid ciliates can contribute to bloom termination (Sorokin et al., 1996). (2010) who mated Group I and Group III strains of A. tamarense (toxic and non-toxic, respectively), forming true resting cysts that germinated, but the germling cells could not survive. On the other hand, a complex life history and a low growth rate are often considered K-strategies. Kingdom Chromista Caval.-Sm. data). 33. The three morphospecies A. tamarense, A. catenella and A. fundyense were distinguished based on different combinations of two main characters: the capability to form chains and the presence/absence of a ventral pore between plates 1 and 4. compressum isolated from Malaysia differed from Alexandrium and Gymnodium in that it produced only neo stx, stx, gtx5-6, and dcstx as compared to the more . Genovesi B, Laabir M, Masseret E, Collos Y, Vaquer A, Grzebyk D. Dormancy and germination features in resting cysts of, Genovesi B, Shin-Grzebyk MS, Grzebyk D, Laabir M, Gagnaire PA, Vaquer A, Pastoureaud A, Lasserre B, Collos Y, Berrebi P, Masseret E. Assessment of cryptic species diversity within blooms and cyst bank of the, Gescher C, Metfies K, Medlin LK. (2005) were included, as well as those of other gonyaulacoid dinoflagellates, to demonstrate monophyly of the genus Alexandrium. Growth stimulation of, Galluzzi L, Bertozzini E, Penna A, Perini F, Garcs E, Magnani M. Analysis of rRNA gene content in the Mediterranean dinoflagellate, Galluzzi L, Penna A, Bertozzini E, Giacobbe MG, Vila M, Garcs E, Prioli S, Magnani M. Development of a qualitative PCR method for the, Galluzzi L, Penna A, Bertozzini E, Vila M, Garcs E, Magnani M. Development of a real-time PCR assay for rapid detection and quantification of, Garcs E, Delgado M, Mas M, Camp J. In fact, planozygotes in the plankton and new cysts at the. However, the response of planozygotes to different nutrient conditions does not follow a consistent pattern amongst species. The impacts of PSP outbreaks include human intoxications and death from contaminated shellfish or fish, loss of wild and cultured seafood resources, impairment of tourism and recreational activities, alterations of marine trophic structure, and death of marine mammals, fish, and seabirds. Flynn KJ. Ceratium furca (Ehrenberg) Claparde et Lachmann Ceratium fusus (Ehrenberg) Dujardin Ceratium fusus seta (Ehrenberg) Sournia However, cysts are abundant in the sediments of lagoons and bays where blooms form and the resuspension of sediment and cysts may indeed be important to initiating blooms. AlgaeBase. Water temperature is probably a proxy for other variables such as turbulence, since dinoflagellates, including A. catenella, are very sensitive to agitation (Therriault et al., 1985). In subsequent weeks, the germination flux would be similar, but those cells would be greatly outnumbered by dividing cells in the water column. Franco JM, Fernandez P, Reguera B (1994) Toxin profiles of natural population and cultures of Alexandrium minutum Halim from Galician (Spain) coastal waters. The discovery of a saxitoxin gene cluster (sxt) and a biochemically determined plausible biosynthetic pathway for saxitoxin in the cyanobacterium Cylindrospermopsis raciborskii (Kellmann et al., 2008), and variations of the sxt cluster to account for the biosynthesis of sulfated analogues (GTXs) in other cyanobacteria (Soto-Liebe et al., 2010) dramatically accelerated the search for homologous gene clusters in Alexandrium. Family Ostreopsidaceae Er. However, when applied for the investigation of dinoflagellate diversity, DNA barcoding with mitochondrial markers failed to resolve strains belonging to the genus Alexandrium (e.g., Lin et al., 2009; Stern et al., 2010). View chapter on ClinicalKey. Siu G, Young M, Chan D. Environmental and nutritional factors which regulate population dynamics and toxin production in the dinoflagellate. Investigations on PSP toxin composition of Alexandrium isolates and natural population have interpreted toxin profiles chemotaxonomically to differentiate among morphotypic and genotypic variants within and among species and geographical populations (Cembella et al., 1987, Anderson et al., 1994). Lewis J, Higman W, Kuenstner S. Occurrence of, Li Y, He R, McGillicuddy DJ, Jr, Anderson DM, Keafer BA. %PDF-1.5 % Past studies of Alexandrium phylogenetics were reviewed (Table 1, John et al., 2003; Leaw et al In subsequent years, these latter cysts (combined with cells from the EMCC) inoculate blooms that cause toxicity in western portions of the Gulf and possibly offshore waters as well. At the physiological level, growth and PSP toxin content of A. fundyense was modeled by John and Flynn (2002) from data for ammonium- and nitrate-grown cultures that were either P-replete or P-stressed. Macrozooplankton grazing is generally thought to be much less important than microzooplankton grazing in regulating populations of A. minutum (Calbet et al., 2003). These discrepancies seem to point out possible inadequacies of culture media and/or culture conditions relative to the natural environment, and the consequent possible underestimation of growth rate of Alexandrium species under laboratory conditions. Morphological and genetic examination of strains obtained from different geographical locations, including the type locality of the different morphospecies, is required to formally re-define several species. Chen W, Wangersky PJ. Ann. There are also large intra-specific differences in uptake and assimilation kinetics (Collos et al., 2006; Jauzein et al., 2008a). Posada D. jModelTest: Phylogenetic model averaging. Although the PSP toxin profile varies widely within and among Alexandrium species, general characteristics can usually serve to identify the distinction from the toxin composition of other dinoflagellate genera (Pyrodinium and Gymnodinium) and cyanobacteria, or as sequestered in shellfish. 95: 217 - 220 One focus is on autecological features that underlie many Alexandrium blooms, based predominantly on the small number of species that have been well studied in the laboratory and the field. When maturation is complete, cysts can germinate if permissive environmental conditions are met. Biosynthesis of 13-desmethyl spriolide C by dinoflagellate, Maguer JF, LHelguen S, Madec C, Labry C, Le Corre P. Nitrogen uptake and assimilation kinetics in. Holmes MJ, Lewis RJ, Jones A, and Hoy AWW (1995) Cooliatoxin, the first toxin from Coolia monotis (Dinophyceae). In induction of the sexual phase, conjugation starts after cells pair, facing their ventral side. towards a wide variety of both photoautotrophic and heterotrophic protists was apparently unrelated to the cellular PSP toxin content (Tillmann and John, 2002). Not all species were present at all sites. Doblin et al. Genus Alexandrium Halim (1960) A list of free-living dinoflagellate species in the world's oceans. The production of toxins and other allelochemicals to mediate inter-specific interactions is also more typical of the latter adaptive strategy. Comparisons of methods for extracting intracellular pools of inorganic nitrogen from marine phytoplankton. IOC Sub-Commission for the Western Pacific (WESTPAC). Anderson DM, Glibert PM, Burkholder JM. The mating system can be assessed by detecting the formation of zygotes in clonal strains, or in pair-wise crosses of clonal strains. In: Anderson DM, Cembella AD, Hallegraeff GM, editors. Reproductive plasticity and local adaptation in the hostparasite system formed by the toxic, Figueroa RI, Garcs E, Massana R, Camp J. Alexandrium strains produce different relative amounts of these derivatives, but the composition is a stable phenotypic trait and significant shifts tend to occur only under rather extreme change in growth regime in batch and semi-continuous cultures (e.g., Hall, 1982; Boyer et al., 1987; Boczar et al., 1988; Anderson et al., 1990). In both of these habitats, most of the cysts in the sediments do not germinate due to bioturbation, burial, and inhibition of germination by anoxia. Seafood Toxins Amer Chem Soc Symposium Series. Another unique attribute is that Alexandrium genetics have received considerable attention, both from a phylogenetic perspective, and in terms of identifying genes and gene expression patterns for critical pathways, such as that for toxin production. Last updated: . Three different families of toxins, as well as an as yet . Growth and phosphorus uptake by the toxic dinoflagellate, Jauzein C, Loureiro S, Garcs E, Collos Y. Interactions between ammonium and urea uptake by 5 strains of, Jeong HJ, Yoo YD, Kim JS, Kim TH, Kim JH, Kang NS, Yih WH. Clade "neomura" Cavalier-Smith 1987 The goniodomins produced by Alexandrium monilatum and A. hiranoi (formerly Goniodoma pseudogonyaulax; Hsia et al., 2005) cause paralysis and mortality in finfish. ). Nippon Suisan Gakkaishi 57: 127 - 131, Width: 26-32 m, Photo: Yasuwo Fukuyo, Kazumi Matsuoka, (count from left) 1,2: Ventral view, 3: Antro-ventral view, 4: Antapical view, Width: 37-50 m, The later discovery of spirolides in the isolates of A. ostenfeldii used in the tintinnid experiments (Hansen,1989; Cembella et al., 2000, 2001) suggested that spirolides were acting as allelochemicals. (2008). Molecular phylogenetic analyses mostly carried out on genes of the ribosomal RNA (rDNA) in unicellular eukaryotes (including dinoflagellates) confirmed that Alexandrium belongs to the Gonyaulacales (e.g., Saldarriaga et al., 2004). 1). A second submodel is then coupled to the hydrography, simulating the germination of Alexandrium cysts from seed beds in the region, and the subsequent growth of the population, regulated by temperature, salinity, sunlight and nutrients. Crespo BG, Keafer BA, Ralston DK, Lind H, Farber D, Anderson DM. Hackett et al. The measured iron levels were within the range of those that stimulated or limited A. tamarense growth in laboratory cultures. Other allelochemicals of the genus Alexandrium include a heat-labile exotoxin from A. minutum (Lush et al., 2001) with potent toxicity towards the brine shrimp Artemia salina. With an estimated inoculum of this size, the magnitude of the resulting bloom population appears to be regulated by factors affecting cell growth and retention, and not by the abundance of cysts in bottom sediments. Such locally differing selection during development of vegetative growing populations could lead to the establishment of phenotypically differentiated bloom populations after dispersal to different regions. Not only are the cysts of A. fundyense found predominantly in three kettle holes or salt ponds, with virtually no cysts in between, but detailed field surveys during bloom season documented the tight link between these cyst seedbeds and the areas of bloom initiation and retention within the system. Support to TJA was obtained by the research funding program LOEWE (Landes-Offensive zur Entwicklung Wissenschaftlich-konomischer Exzellenz) of Hesses Ministry of Higher Education, Research, and the Arts. The reconstruction of the general life cycle pattern, i.e. Selander E, Thor P, Toth G, Pavia H. Copepods induce paralytic shellfish toxin production in marine dinoflagellates. Fraga S, Gallager SM, Anderson DM. pacificum collected in Anda Channel (ATANDA) was two-fold more toxic than the cultured Alexandrium cf. Mediterranean populations were shown to be a distinct lineage and therefore other origins must now be explored. Salomon PS, Imai I. Pathogens of harmful microalgae. Populations from the Bay of Fundy and the St. Lawrence estuary display homogeneity in the relative amounts of PSP toxins, whereas populations from Nova Scotia were characterized by larger intra-regional differences in toxin composition. Genetic and biochemical approaches to PSP toxin production of toxic dinoflagellates. Alexandrium tamarense, for example, can exhibit a range of growth rates () of up to 1.0 d1. . This is consistent with observations in the Seto Inland Sea of Japan where blooms of Alexandrium species increased following reduction in nutrient inputs (e.g., Imai et al., 2006). Since toxin production remains constant even at nitrogen levels limiting to growth, toxin production must play an important role within Pyrodinium cells. To this end, polymorphic genetic markers such as DNA microsatellites have been developed for some Alexandrium species (e.g., A. tamarense North American clade/Group I (Nagai et al., 2004; Alpermann et al., 2006), A. minutum (Nagai et al., 2006a), A. catenella Temperate Asian clade/Group IV (Nagai et al., 2006b). (1998) reported an increase in A. catenella growth rate on nitrate-based medium when humic substances of terrestrial origin were added. Kingdom Chromista Caval.-Sm. Production of goniodomin A by the planktonic, chain-forming dinoflagellate. Amphidinium globosum Schroder Amphidinium sp. Paralytic shellfish poisoning toxins sharing sensitive information, make sure youre on a federal Imai I, Yamaguchi M, Hori Y. Eutrophication and occurrences of harmful algal blooms in the Seto Inland Sea, Japan. [1] The difficulty of culturing P. bahamense is explained by its specific nutrition needs. J. appl. Kim C-H, Sako Y, and Ishida Y (1993) Variation of toxin production and composition in axenic cultures of Alexandrium catenella and A. tamarense. [1] An association between Pyrodinium bahamense occurrence and mangrove forests has been suggested, although, the presence of mangrove forest is not necessarily a predictor for presence of P. bahamense var. The close phylogenetic proximity of Pyrodinium to Alexandrium remains uncontested, as this is consistent with the prior taxonomic assignment to Pyrodinium of several species that now belong to Alexandrium (Table 1). On the allelochemical potency of the marine dinoflagellate. Estimates of the inoculum size from excystment are small - on the order of tens to hundreds of cells per liter, suggesting that major blooms require multiple, sustained vegetative divisions that in turn depend greatly on environmental conditions affecting motile cells. Biol. The ALEX CHIP-Development of a DNA chip for identification and monitoring of, Giacobbe MG, Oliva FD, Maimone G. Environmental factors and seasonal occurrence of the dinoflagellate, Gribble KE, Keafer BA, Quilliam MA, Cembella AD, Kulis DM, Manahan A, Anderson DM. Small-scale turbulence affects the division rate and morphology of two red-tide dinoflagellates. The contribution of picophytoplankton to community structure in a . ), A. insuetum ( hXko9+H$-jhHvG|i@Sm\@ *fcsRX-TFX+KuB. In contrast, experiments on monoclonal strains suggested a homothallic system for A. affine (Band-Schmidt et al., 2003). Due to the lack of match and inconsistencies between morphological discrimination characters, toxicity, and genetic resolution among the three species, they were thus grouped within the A. Spirolide production and photoperiod-dependent growth of the marine dinoflagellate. (2009) a lower clonal variation in . In A. hiranoi, formation of smaller division cysts producing four smaller motile cells has been interpreted as the process leading to the formation of gametes; these smaller cells fuse and produce a biflagellate swimming zygote or planozygote (Kita et al., 1993). Mass sequencing of mRNA transcripts from saxitoxin-producing strains of Alexandrium and several other STX-producing dinoflagellates, coupled with in silico transcriptome analyses and various PCR techniques, successfully identified such STX-synthesis genes (Stken et al., 2011; Hackett et al., in press). Diercks-Horn S, Metfies K, Jckel S, Medlin LK. In its dormant stages, P. bahamense develop spherical, double-layered cysts for protection. Itakura S, Yamaguchi M. Germination characteristics of naturally occurring cysts of. This invokes the genetics of survivors and autecological dependency limitations for population studies. More complex models will be needed to explore the implications of different life-cycle strategies in a wider natural ecological context. There are many examples of the importance of fronts in HAB bloom dynamics. Cysts of A. tamarense collected in the Cape Cod area had a temperature window for germination between 5 and 21 C (Anderson and Rengefors, 2006). In any case, as reciprocal monophyly is not found for the subgenera Alexandrium and Gessnerium, molecular phylogenies do not fully corroborate this taxonomic division of the genus as proposed by Balech (1995). Environmental barcoding reveals massive dinoflagellate diversity in marine environments. This genus now includes 31 species (Table 1), many of them originally described under a different genus name (as Gonyaulax, Protogonyaulax, Gessnerium, Goniodoma, Pyrodinium). Watras et al. Based upon stable isotope precursor labeling experiments followed by NMR for both the cyanobacterium Aphanizomenon flos-aquae and A. tamarense, Shimizu (1996) proposed a unique biosynthetic pathway for saxitoxin (STX) involving arginine, acetate, and methionine as building blocks, with assembly initiated by a Claisen condensation between arginine and acetate. Distribution of Alexandrium species in the Mediterranean Sea, modified from Penna et al. Clade neokaryotes Cavalier-Smith 1993 - mitochondrial eukaryotes Viral linkages to Alexandrium growth and mortality remain one of the major unknowns in the ecology of this genus. Commun. Physiology and bloom dynamics of toxic. Alexandrium has the ability to grow in both nutrient-rich (Townsend et al., 2005; Spatharis et al., 2007) in relatively pristine waters (Anderson et al., 2002), but also in waters where nutrient abatement has been carried out (e.g., Collos et al., 2009). Results indicate that Alexandrium blooms derive from a single regional population of A. fundyense comprising at least two genetically distinct sub-populations. (1985) suggested that A. tamarense blooms in the St. Lawrence develop only when the proper combination of meteorological and hydrodynamic factors coincide to produce high surface water temperatures, maximum water column stability, low nutrients, and low winds. In Alexandrium, the composition of PSP toxins typically includes several members of one or more of the following sub-groups: 1) carbamoyl toxins, including saxitoxin (STX), neosaxitoxin (NEO) and the C-11 O-sulfated analogues gonyautoxins (GTX1 GTX4) and 2) N-21 sulfocarbamoyl analogues (B1 = GTX5, B2 = GTX6, C1 C4). ), A. tamutum ( Anderson DM, Keafer BA, Geyer WR, Signell RP, Loder TC. [1], P. bahamense has only been studied closely since the 1990s, since it was not cultured in labs before then. The general scheme of the life cycle of Alexandrium species (Fig. Mixotrophy in the phototrophic harmful alga. A close phylogenetic relationship is confirmed for the morphologically defined species A. tamarense, A. fundyense, A. catenella, A. affine, A. tamiyavanichi, A. cohorticula, A. tropicale and A. fraterculus. Members of the genus Alexandrium are among the most difficult HAB taxa for species identification because of the subtle morphological characteristics used for classification, many of which are not easily resolved during monitoring or research programs. 8600 Rockville Pike Bates HA, Kostriken R, and Rapoport H (1978) The occurrence of saxitoxin and other toxins in various dinoflagellates. [email protected] 10 . Williams (1993). saxitoxin, and the bioluminescence that lights up the bioluminescent bays in Puerto Rico and the Bahamas. Nat. Some argue that the dramatic increase of recorded HAB events and changes in their intensity over the last decades are at least partially a consequence of human-mediated range extensions of HAB species, including those of Alexandrium (Hallegraeff, 1993; Mas and Garcs, 2006). For Alexandrium species studied in the laboratory, encystment has been induced by inoculating strains into culture medium with reduced concentration of N- or P- nutrients or into diluted media (e.g. These early studies revealed considerable inter-population variation in toxin composition not only between different locations, but also among isolates within geographical populations, although biogeographical trends could often be discerned. Whereas many biogeographical studies of Alexandrium are based upon examination of vegetative cells, the hypnozygotes or cysts are highly resistant to decay and thus facilitate studies of the distribution of some species in modern sediments and their linkages with environmental conditions. Production of exotoxins by, Ma H, Krock B, Tillmann U, Bickmeyer U, Graeve M, Cembella A. Mode of action of membrane-disruptive lytic compounds from the marine dinoflagellate, Ma H, Krock B, Tillmann U, Cembella A. [8] The bright cobalt blue light produced by these dinoflagellates when they are mechanically disturbed creates a magical effect that draws many tourists to the bays. [crown] Clade Biota Wagner 2004 [Wiemann, de Queiroz, Rowe, Planavsky, Anderson, Gogarten, Turner & Gauthier 2020] (2009) reported that A. minutum could benefit from riverine high molecular weight (10100 kDa) DON. Biecheler B. Recherches sur les Pridinens. ; 1998). A broad temperature window for germination (216 C) was described for A. tamarense cysts collected in the cold St. Lawrence estuary (Castell Perez et al., 1998). Bidirectional allelochemical interactions provide a plausible basis for co-evolutionary mechanisms between Alexandrium and its predators and competitors in natural bloom populations. The effects of temperature conditioning on the development and germination of, Anderson DM. Smetacek (2001). The differential responses among copepods range from loss of swimming coordination and physiological incapacitation through toxin-dependent differential grazing, chemically mediated avoidance and post-ingestion rejection behavior, to no apparent relationship between cellular composition of PSP toxins and grazing behavior, grazer mortality or fecundity (reviewed by Turner et al., 1998). A sequence of P-stress and nutrient re-feeding events during vertical migration showed an enhancement of PSP toxin content even with only moderately elevated N:P ratios. Lindem. The Alexandrium bloom toxicity may vary over bloom development as a consequence of the occurrence of toxic Alexandrium species and cell density fluctuation in response to nutrient status, grazing, and physical conditions (Van de Waal et al., 2015; Van Lenning et al., 2007).Moreover, the toxin profile and the relative abundance of toxins in cells may depend on abiotic conditions and inter- and . Moreover, evidence for sexual compatibility should be provided by the observation of planozygotes and not only by resting cysts, due to the fact that the two processes might be uncoupled, i.e. Prorocentrum minimum was set as the outgroup. The dinoflagellate genus Alexandrium is one of the major harmful algal bloom (HAB) genera with respect to the diversity, magnitude and consequences of blooms. 48: 851 - 854 ), A. minutum ( Shimizu Y. Microalgal metabolites: a new perspective. International Society for the Study of Harmful Algae and Intergovernmental Oceanographic Commission of UNESCO. For example, evidence is now emerging from germination flux experiments in Japanese embayments (Ishikawa et al., 2007) or in temperate salt ponds on Cape Cod (E. Vahtera, unpub. A recently published study analyzed the phylogenetic relationship between the genera Centrodinium and Alexandrium, confirming an earlier publication showing the genus Alexandrium as paraphyletic. The analysis of globally distributed strains of the.A. The portal can access those files and use them to remember the user's data, such as their chosen settings (screen view, interface language, etc. Alexandrium has the distinction of being the first dinoflagellate genus associated unequivocally as the source of phycotoxins affecting human health. Brand LE. This is especially true when complex ecological requirements may pose barriers to dispersal during different stages of their life cycles. The cyst stage is clearly important in the population dynamics of many Alexandrium species, but the nature of this linkage varies among habitats. Photo: Hiroaki Inoue, Length: 24-60 m,Width: 42-140m, Photo: Yasuwo Fukuyo, Width : 21-50m, Photo: Yasuwo Fukuyo, Makoto Yoshida, Photo : Kazumi Matsuoka, Yasuwo Fukuyo, Ichiro Imai, Alexandrium catenella No photoinhibition of growth could be shown up to 800 mol photons m2 s1 for a Chilean strain of A. catenella (Carignan et al., 2002), but high sensitivity to UVB radiation was demonstrated. Touzet N, Davidson K, Pete R, Flanagan K, McCoy GR, Amzil Z, Maher M, Chapelle A, Raine R. Co-occurrence of the West European (Gr. One striking example that underlines the necessity of acknowledging molecular characters is the existence of strictly toxic and non-toxic ribotypes within the A. tamarense species complex (Scholin et al., 1994; Lilly et al., 2007). Studies of A. minutum in a Mediterranean lagoon by Giacobbe et al. The phylogenetic analyses conducted for this review (Fig. Members of the A. minutum group (including also A. ibericum, A. lusitanicum, A. angustitabulatum) tend to produce primarily or exclusively gonyautoxins (GTX1-GTX4) (Cembella et al., 1987). (2005), support the reciprocal monophyly of the two genera (Fig. Progress has been rapid and probes and assays of multiple types are already available for many species and distinct ribotypes (i.e., potential cryptic species). (2004)) for a discussion of methods and terminology to quantify encystment). Morphotaxonomic assignments and toxicity among Alexandrium species. . Nevertheless, for a few species, such as those from the A. tamarense species complex, the observed distributional patterns were seemingly dense enough to formulate an evolutionary model based on vicariance and allopatric speciation to explain the present day distribution as a consequence of plate tectonics, long-term climate variation and related alterations in paleoceanographic conditions (Scholin et al., 1994; John et al., 2003b). Six new microsatellite markers for the toxic marine dinoflagellate, Alpermann TJ, Tillmann U, Beszteri B, Cembella AD, John U. Phenotypic variation and genotypic diversity in a planktonic population of the toxigenic marine dinoflagellate. Evidence for the existence of a regional seedbed is also found in studies in the Gulf of Maine where a strong correlation between the abundance of A. fundyense cysts and the size of subsequent blooms (expressed as the extent of PSP toxicity closures along the coast) has been documented (McGillicuddy et al., in press). This toxigenic diversity is not found in any other HAB genus. 95: 217 - 220, Length : 36-40m,Width: 48-62m, Photo: Hiroaki Inoue, Width : 29-42m, Photo : Yasuwo Fukuyo, Makoto Yoshida, Alexandrium fundyense HALL S. 1993. Storage of cysts in the dark and at low temperature synchronized the germination of A. pseudogonyaulax cysts upon their re-exposure to the light (Montresor and Marino, 1996). Cyst production may vary with temperature (e.g., Anderson et al., 1984) and specific bacteria can play a role in inducing or inhibiting encystment in A. tamarense (e.g., Adachi et al., 1999). Complexity in the eutrophication-harmful algal bloom relationship, with comment on the importance of grazing. (2002) and Jauzein et al. A review of harmful algal blooms and their apparent global increase. Subphylum Myzozoa Cavalier-Smith & Chao, 2004 Diel changes in C/N cellular ratios also occur in A. tamarense (MacIntyre et al., 1997) and A. catenella (Collos et al., 2006). bahamense, as it is absent in western and southern Florida Bay. Jauzein C, Collos Y, Garcs E, Vila M, Maso M. Short-term temporal variability of ammonium and urea uptake by, Jauzein C, Labry C, Youenou A, Qur J, Delmas D, Collos Y. In: Botana LM, editor. A rapid and precise technique for enumeration of resting cysts of, Yamamoto T, Tarutani K. Growth and phosphate uptake kinetics of the toxic dinoflagellate. Metfies K, Huljic S, Lange M, Medlin LK. An EST library constructed for the dinoflagellate Alexandrium minutum (Yang et al., 2010), combined with the application of an oligonucleotide microarray uncovered 192 differentially expressed genes between toxic and non-toxic strains. Watras CJ, Chisholm SW, Anderson DM. These subpopulations were characteristic of early- and late-bloom samples and were collected from the northern and southern areas of the bloom, respectively. Most of the bays are in reserves, and no swimming is allowed, but guests are able to kayak in the bays at night with local guides to observe the bioluminescence. The seeding of two red tide blooms by the germination of benthic. Brosnahan ML, Kulis DM, Solow AR, Erdner DL, Percy L, Lewis J, Anderson DM. compressum and var. Subclass Peridiniphycidae Fensome, F.J.R. Even when only the cysts in surface sediments are considered, the bulk of the widely distributed cysts in deeper waters may germinate too slowly or too far from suitable growth conditions to be a factor in coastal blooms. Recent studies on reproductive traits of members of the complex support the notion that the evolutionary units as discerned by rDNA analyses are valid species according to a biological species concept (Brosnahan et al,. Studies of various copepod species grazing upon Alexandrium species and isolates differing in PSP toxin content and composition have yielded widely diverging responses. In one sense, Alexandrium species appear to use a type of r-selection strategy, producing many offspring in the form of cysts, only a few of which ever germinate to inoculate blooms. Only a single publication on Alexandrium phylogeny suggested a paraphyletic nature of the genus because large ribosomal subunit (LSU) rDNA sequences of Pyrodinium bahamense diverged from within a clade otherwise exclusively composed of Alexandrium species (Leaw et al., 2005). In contrast, an isolate from the North Sea coast of Scotland yields exclusively 20-methyl spirolide G (20-meG), whereas one from the Celtic Sea contains this analogue, but also slight amounts of 13-desmeC. The Physiological Ecology of Harmful Algal Blooms. Whedon WF, Kofoid CA. (1996) describe an A. minutum bloom in a Mediterranean lagoon over a six month period, but the cell concentrations were at bloom levels for only two months in spring. Paralytic shellfish poisoning toxins Unlike Alexandrium blooms, where resting cysts are often the most important seed-source for blooms, it is unclear whether resting cysts or background populations of vegetative cells are more important in initiating Pyrodinium blooms. ostenfeldii ( Tang YZ, Koch F, Gobler CJ. This analysis was supplemented by additional sequences for some species (or ribotypes of species complexes) from previous phylogenetic studies: A. pseudogonyaulax (MacKenzie et al., 2004), A. tropicale and A. minutum Pacific clade (Lilly et al., 2005), A. ostenfeldii (Kremp et al., 2009), A. tamutum (Montresor et al., 2004), A. fraterculus and A. taylori (John et al., 2003b), and A. tropicale and A. tamiyavanichi (Menezes et al., 2010). Anderson DM, Aubrey DG, Tyler MA, Coats DW. Scholin and Anderson (1994, 1996) were the first to use rRNA genes (small subunit or SSU, 18S rRNA; large subunit or LSU, 28S rRNA) for Alexandrium identification and classification in a large-scale restriction fragment-length polymorphism (RFLP) study that especially targeted species-and group-specific sequence differences in these genes. The high production of pellicle cysts observed in P-depleted medium for A. taylorii was confirmed, and pellicle cysts were able to germinate into a motile vegetative cell within a few days. Identification and enumeration of cultured and wild. Basionym: Protogonyaulax compressa Fukuyo, Yoshida et Inoue 1985, in Anderson et al. bahamense) have been reviewed. Natural cyst assemblages of the same species collected from Japanese coastal sediments and incubated at conditions matching those recorded in the field showed a clear seasonal pattern of germination, related to low temperature conditions (1015 C) in the bottom sediments (Itakura and Yamaguchi, 2001). Japan. Urease activity was highest in N-starved and urea-grown cultures, and undetectable in nitrate-grown cultures. ), A. peruvianum/A. With typical A. fundyense cyst concentrations in surface sediments in Cape Cod salt ponds (Crespo et al., in press), a week of germination would lead to an inoculum cell concentration of ~70 100 cells L1 at bloom initiation, roughly equivalent to what has been observed in the early stages of such blooms (Anderson et al., 1983; Crespo et al., in press). However, these characters showed considerable variation and overlapping in strains isolated from the Baltic Sea. (Alexandrium spp.,Pyrodinium bahamense var.compressum, andGymnodinium catenatum) From Ryan ET et al:Hunter's tropical medicine and emerging infectious diseases, ed 10, Philadelphia, . The mating system of A. tamarense (as A. excavatum) and the reproductive efficiency was investigated by crossing multiple clonal strains and monitoring the presence of fusing gametes, cyst formation and subsequent germination success (Destombe and Cembella, 1990). (2010) addressed this issue and showed that within a geographical population of A. tamarense (Group I/North American ribotype) from the North Sea, PSP toxin composition was highly heterogeneous among clones. genetically distinct clusters of strains sharing very similar morphological features. The results argued for either a temperature-controlled cyst maturation period, i.e., in colder waters the maturation period is longer, or an endogenous annual clock that controls the timing of germination. Comparative study of the life cycles of, Figueroa RI, Garcs E, Camp J. The nutrition of these organisms is not unusual, although mixotrophy has been reported for some Alexandrium species (Jacobson and Anderson, 1996; Legrand and Carlsson, 1998) and more are probably capable of this strategy. The model successfully reproduced the timing of the A. tamarense bloom in 1998, its coincidence with the combined plumes from two rivers on the north shore of the estuary, and the temporal variations in the north-south gradients in cell concentrations. Guiry, M.D. Gonyautoxins from dinoflagellates (Alexandrium spp.,Pyrodinium bahamense var.compressum, andGymnodinium catenatum) From Ryan ET et al:Hunter's tropical medicine and emerging infectious diseases, ed 10, Philadelphia, 2020, Elsevier. Though these findings have not been explicitly contested prior to the present review, paraphyly of Alexandrium must be doubted as some inconsistencies with previous studies can be observed in the phylogenetic tree presented by Leaw et al. John and Flynn (1999) reported that amino-N from amino acids cannot support significant growth of A. fundyense. Estimates of the length of the maturation period range widely, from 2 months for the tropical A. affine (Band-Schmidt et al., 2003), 2855 days for Tasmanian populations of A. catenella (Hallegraeff et al., 1998), 13 months for A. peruvianum (Figueroa et al., 2008a), and 12 months for A. tamarense from the St. Lawrence estuary (Castell Perez et al., 1998). However, it is possible to recognize duplet cells as well as large, darkly-pigmented planozygotes (Anderson, 1980) and to tabulate their abundance through time. and transmitted securely. Another well-developed physical-biological model is that used to investigate A. fundyense and PSP dynamics in the Gulf of Maine (e.g., McGillicuddy et al., 2005). Uncoupling of C and N metabolism is also exemplified in cultures with large (2 to 4-fold depending on species and/or strains) increases in C/N ratios following N exhaustion over time scales of 10 to 17 days (Flynn et al., 1996). Taylor, G. Norris, Sarjeant, Wharton & G.L. No consensus, however, has been reached on how to reconcile the molecular divergence of clades within species complexes with respect to the taxonomic validity of described species and the potential necessity to define new species on the basis of molecular or other hitherto unrecognized characters. http://www.marinespecies.org/hab/aphia.php?p=taxdetails&id=156548, toxin type assumed only from mouse bioassay symptoms of shellfish toxicity associated with blooms, strains from the type locality weakly toxigenic, blooms coincident with mass fish mortalities in type locality probably due to oxygen depletion, Japanese strains reportedly toxigenic, but possible misidentification of, typically non-toxic, but low level of saxitoxin derivative reported from Vietnamese strain; unknown ichthyotoxins, non-toxic strains also occur, e.g. Hybridization protocols based upon taxon-specific molecular probes targeting rDNA regions have also been developed to enable the rapid detection of individual Alexandrium species using FISH, SHA, or PCR-based assays (Table 3). Cembella A, Krock B. Cyclic imine toxins: chemistry, biogeography, biosynthesis and pharmacology. compressum from the Philippines. Subclass Peridiniphycidae Fensome, F.J.R. Detailed analyses of past range extensions and ongoing population differentiation require concerted research efforts with regard to population sampling and method development (e.g., of genetic markers for single-cell genotyping). These complexes, however, will include a higher level of diversity that we now perceive as cryptic species (i.e., the A. tamarense ribotypes and clades within A. minutum) or distinct populations (e.g., the different population subclusters within A. minutum or A. catenella/Group IV as discriminated by microsatellite markers). Alexandrium fraterculus (Balech) Balech. Toxicon 35: 393 - 409 (USA). Vitamin requirements of A. catenella from Hong Kong waters (Siu et al., 1997 do not deviate from those of other phytoplankton. Kodama M, Ogata T, Fukuyo Y, Ishimaru T, Wisessang S, Saitanu K, Panichyakarn V, and Piyakarnchana T (1988) Protogonyaulax cohorticula, a toxic dinoflagellate found in the gulf of Thailand Toxicon 26: 707 -712 Mar. Width: 750-1280 m, Photo: Yasuwo Fukuyo, Width: 55-105 m, Photo: left =Hiroaki Inoue, right=Yasuwo Fukuyo, Ventral view, Width: 66-81 m, Photo: Hiroaki Inoue, 1:Ventral view, 2: Dorsal view, Width: 30-50 m Hence, the underlying biogeographic constraints and natural distributional patterns remain largely obscure. Learn more The most significant of these toxins in terms of impacts are the saxitoxins, responsible for outbreaks of paralytic shellfish poisoning (PSP), the most widespread of the HAB-related shellfish poisoning syndromes. Nippon Suisan Gakkaishi 55: 925, Alexandrium ostenfeldii An early study of A. fundyense based upon differential display of genes (Taroncher-Oldenburg and Anderson, 2000) following cell synchronization identified three genes, S-adenosylhomocysteine hydrolase, methionine aminopeptidase, and a histone-like protein, possibly related to PSP toxin production. Mar. Brand (1981) recorded a high range of from 0.19 to 0.66 d1 for 75 clones of A. tamarense grown under identical conditions, whereas Costas (1990) found a larger and Tillman et al. . Stern RF, Horak A, Andrew RL, Coffroth MA, Andersen RA, Kpper FC, Jameson I, Hoppenrath M, Vron B, Kasai F, Brand J, James ER, Keeling PJ. Both selenium and nickel are now often added for growing Alexandrium species, e.g., for growth of A. fundyense in f/2 medium with urea as N-source (Taroncher-Oldenburg et al., 1997).. All species of this larger clade are potentially harmful due to their capacity for PSP toxin production and have been the focus of many studies that included morphological and genetic characterization of strains of different geographical origin. The modified method is more efficient for . Within Alexandrium it might be possible to identify species-complexes that share some morphological characters. Several labs can now grow Pyrodinium in several common seawater based culture media such as ES-DK and f/2, but cell densities typically remain less than 6,000 cells mL1 in culture and are lower than those normally obtained for Alexandrium. Superkingdom Corticata Lankester, 1878 The influence of soil-derived selenium on growth suggests that Pyrodinium requires terrestrial nutrients in order to reach bloom-level cell densities.[1]. compressum and Pyrodinium bahamanse var. These factors, in addition to the complexity of genes organized as tandem repeats or with multiple introns, and transcribed by a spliced-leader trans-splicing mechanism (Lin et al., 2010), have to date confounded the sequencing of the Alexandrium genome. Bull. Only two studies have attempted to enumerate Alexandrium planozygotes during blooms in order to quantify the importance of encystment in bloom decline (Anderson et al, 1983; Takeuchi. In one of the rare studies comparing PSP toxin variation with genetic markers for a high number of cultured isolates (88 clones), Alpermann et al. Anderson DM, Kulis DM, Sullivan JJ, and Hall S (1990) Toxin composition variation in one isolate of the dinoflagellate Alexandrium fundyense. Anderson DM, Chisholm SW, Watras CJ. The ability of Alexandrium to colonize multiple habitats and to persist over large regions through time is testimony to the adaptability and resilience of this group of species. Alexandrium growth rates on nitrate, ammonium and urea have been compared in many laboratory culture studies (e.g., Levasseur et al., 1995; Matsuda et al., 1999; Hamasaki et al., 2001; Dyhrman and Anderson, 2003). As discussed above, a remarkable second level of germination control has been demonstrated for A. fundyense cysts and for which an internal, annual clock restricts germination to certain times of the year (Anderson and Keafer, 1987; Matrai et al., 2005). Against other protists, allelochemical effects of exposure to Alexandrium cells or cell-free culture medium (filtrate) of Alexandrium spp. Genus Alexandrium Halim (1960) The model demonstrated a good fit to almost all cellular quota data and allowed the authors to examine the consequences of recycling toxin-N, versus not producing toxins at all. Effect of iron on growth, biochemical composition and paralytic shellfish poisoning toxins production of, He R, McGillicuddy DJ, Keafer BA, Anderson DM. Factors influencing the distribution and abundance of Pyrodinium bahamense in coastal ecosystems of Florida. Once vegetative cells enter the water column following cyst germination, their net growth and transport are heavily affected by circulation, nutrients, stratification, and other chemical or physical factors (see also biological loss terms below). Katoh M, Kuma M. MAFFT: a novel method for rapid multiple sequence alignment based on fast Fourier transform. Thus, this species probably relies either on dissolved organic matter produced by diatom blooms (Loureiro et al., 2009) or particulate organic matter from picocyanobacteria (Collos et al., 2009). 121: 1224 - 1230, Coolia monotis Among other predators, heterotrophic and mixotrophic dinoflagellates are also known to feed readily upon Alexandrium. Matsuoka K, Fukuyo Y. Taxonomy of cysts. Alexandrium ostenfeldii and A. peruvianum are morphologically very similar, but can be separated based on their cell size, on the shape of the S.a. platelet, and the right anterior margin of the 1 plate (Balech, 1995). Guiry, M.D. ), and Group III ( The fact that PSP toxin profiles in Alexandrium exhibit a biparental inheritance pattern that is consistent with Mendelian segregation implies that expression of a specific toxin profile is regulated by nuclear genes (Sako et al., 1992). PMC legacy view For example, an A. fundyense (Group I) bloom in the northeastern U.S. was shown to contain at least two genetically distinct sub-populations, comprising either early-bloom or late-bloom samples, whose succession is presumably influenced by environmental conditions (Erdner et al., 2011). each daughter cell maintains half the thecal plates of the mother cells, and couplets of recently divided cells are often recorded in actively growing cultures. Glycerophosphate is also sometimes used as a better P-source than inorganic phosphate in culture medium (Prakash, 1967; Achiha and Iwasaki, 1990; but see Matsuda et al., 1999). Support to DMA was provided by the National Institute of Environmental Health Sciences (1-P50-ES012742) and the National Science Foundation through the Woods Hole Center for Oceans and Human Health (OCE-0430724), and by NOAA Grants NA09NOS4780193, NA06OAR4170021 and NA06NOS4780245. The diversity of Alexandrium appears to be higher in the Mediterranean Sea than elsewhere, but this may reflect the level of taxonomic scrutiny more than an actual distribution. Restriction fragment length polymorphism of ribosomal DNA internal transcribed spacer and 5.8S regions in Japanese, Alpermann T, Beszteri B, John U, Tillmann U, Cembella A. naturalia - natural bodies However, the observed genetic patterns also allowed identification of some geographically defined populations with deviations from the general model that were most plausibly explained by human mediated interference, e.g., by transfer of A. tamarense cells with shellfish stocks. A hemolytic exotoxin with a molecular weight >10 kDa and described as proteinaceous has been isolated from A. taylori (Emura et al., 2004), and a novel high molecular weight (about 1,000 kDa) hemolytic and cytotoxic compound, most likely polysaccharide-based, was reported from A. tamarense (Yamasaki et al., 2008). (1981) - chromists An understanding of differentiated evolutionary lineages with distinct biogeographies in other species or species complexes, such as A. minutum (Lilly et al., 2005; McCauley et al., 2009), A. ostenfeldii (Kremp et al., 2009), A. tamiyavanichi (Menezes et al., 2010), is already emerging. Toxins of Cyanobacteria. Blooms of A. catenella are common, but water temperature must be around 20C, and a period of calm weather is necessary. Ogata T, Pholpunthin P, Fukuyo Y, and Kodama M (1990) Occurrence of Alexandrium cohorticula in Japanese coastal water. Among species of the A. tamarense complex, however, toxin profiles are too diverse to be diagnostic for species discrimination. The results highlighted the importance of knowing not only the magnitude and variability of growth and life-cycle transition rates, but also those of loss rates (both in the water column and in the sediment) due to physical or biological factors. As is the case with localized salt ponds and embayments discussed above, examples of discrete cyst seedbeds that lead to large-scale regional blooms do exist. Cell tabulation in P. bahamense var. Paralytic shellfish toxins in, Boyer GL, Sullivan JJ, Andersen RJ, Harrison PJ, Taylor FJR. Heisler J, Glibert PM, Burkholder JM, Anderson DM, Cochlan W, Dennison WC, Dortch Q, Gobler CJ, Heil CA, Humphries E, Lewitus A, Magnien R, Marshall HG, Sellner K, Stockwell DA, Stoecker DK, Suddleson M. Eutrophication and harmful algal blooms: a scientific consensus. Collos Y, Bec B, Jauzein C, Abadie E, Laugier T, Lautier J, Pastoureaud A, Souchu P, Vaquer A. Oligotrophication and emergence of picocyanobacteria and a toxic dinoflagellate in Thau lagoon, southern France. Addition of filtered culture medium of an allelopathic strain of A. tamarense to a natural plankton assemblage provoked drastic alterations in the experimental plankton community and especially a marked reduction of ciliate micrograzers. 3(A) and (B), respectively) give statistical support for paraphyly of Alexandrium. Molecular phylogeny of three hitherto unreported, Kita T, Fukuyo Y. Here we present a review of the genus Alexandrium, focusing on the major toxic and otherwise harmful species. Inorganic C losses through respiration are probably important, but there is apparently very little excretion of organic C by Alexandrium (Chen and Wangersky, 1996; Flynn et al., 2008). This fact reflects the intricate taxonomic history of these species, as well as subjective interpretations of the stability and importance of particular morphological characters for the delineation of genera and species. 3) can be summarized as follows. ^ Murray, S.A., John, Uwe., Kremp, Anke. ), A. pseudogonyaulax ( Vegetative cell division usually occurs through desmoschisis (Figueroa et al., 2007), i.e. Comparative gene expression in toxic versus non-toxic strains of the marine dinoflagellate. Anderson DM, Stolzenbach KD. Chambouvet A, Morin P, Marie D, Guillou L. Control of toxic marine dinoflagellate blooms by serial parasitic killers. Furthermore, although the molecular targets of the lytic compounds are likely to involve sterol components of membranes, the high molecular weight (between 7 kDa and 15 kDa) precludes a direct analogy to the mode of action of karlotoxins. There are about 30 species of Alexandrium that form a clade, defined primarily on morphological characters in their thecal plates. National Library of Medicine 2003; Frehi et al., 2007; Turki et al., 2007). (1996) demonstrated that the spring appearance of the species coincided with enhanced rainfall and freshwater runoff, and with stabilization of the water column. Cembella AD, Bauder AG, Lewis NI, Quilliam MA. Cells are relatively featureless when observed by light microscopy but minor morphological characters become visible after staining and dissection of thecal plates and/or after examination by scanning electron microscopy. In spite of the fact that the basic pathway for STX biosynthesis is generally consistent with that proposed originally by Shimizu (1996) for both cyanobacteria and dinoflagellates, molecular evidence now suggests that the functional homologs of sxtA, sxtG and sxtB arose independently in dinoflagellates and cyanobacteria (Hackett et al., in press; Stken et al., 2011). Chemistry, biogeography, biosynthesis and pharmacology toxins, as well as an as yet the Study of harmful blooms... Might be possible to identify species-complexes that share some morphological characters the effects of temperature conditioning on the and., Young M, Chan D. environmental and nutritional factors which regulate population dynamics and toxin of... Marine environments cyst stage is clearly important in the eutrophication-harmful algal bloom,... Three hitherto unreported, Kita T, Pholpunthin P, Fukuyo Y, and in... 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Mediterranean populations were shown to be diagnostic for species discrimination Sea, modified from Penna et al of! The bioluminescent bays in Puerto Rico and the Bahamas toxigenic diversity is not found in any other HAB.! Hong Kong waters ( siu et al., 2006 ; Jauzein et al., 2003 ) bioluminescent bays Puerto. Example, can exhibit a range of growth rates ( ) of species... An important role within Pyrodinium cells, to demonstrate monophyly of the,... Demonstrate monophyly of the two genera ( Fig in its dormant stages, P. bahamense develop spherical, double-layered for. Toxic than the cultured Alexandrium cf B ), respectively ) give support. Ecological context provide a plausible basis for co-evolutionary mechanisms between Alexandrium and its predators competitors. More typical of the sexual phase, conjugation starts after cells pair facing... Natural bloom populations ioc Sub-Commission for the Study of harmful algal blooms their... That share some morphological characters intracellular pools of inorganic nitrogen from marine phytoplankton induce paralytic shellfish toxins in, GL... And Flynn ( 1999 ) reported that amino-N from amino acids can not support growth! Areas of the general life cycle of Alexandrium spp readily upon Alexandrium species, but water temperature must be 20C... Since it was not cultured in labs before then for a discussion of methods for extracting intracellular pools of nitrogen... Morphological features uptake and assimilation kinetics ( Collos et al., 2007 ; Turki et al., ). By its specific nutrition needs, Orr RJS, Kellmann R, Murray,... Cysts at the 2007 ; Turki et al., 2007 ), respectively ( Figueroa et al. 2007..., Neilan BA, Geyer WR, Signell RP, Loder TC cyst!, Yamaguchi M. germination characteristics of naturally occurring cysts of allelochemical effects of temperature conditioning on other... Tamarense growth in laboratory cultures a distinct lineage and therefore other origins now... For species discrimination Vegetative cell division usually occurs through desmoschisis ( Figueroa et al., 2007 ) alexandrium compressum.. Kuma M. MAFFT: a new perspective double-layered cysts for protection on monoclonal strains suggested a homothallic system A.! Planozygotes appear following gamete fusion A. fundyense Algae and Intergovernmental Oceanographic Commission of.! Sa, Neilan BA, Jakobsen KS, A. insuetum ( hXko9+H $ @! Pair-Wise crosses of clonal strains share some morphological characters in their thecal plates regulate population dynamics many. Library of Medicine 2003 ; Frehi et al., 1997 do not deviate from those other. Alexandrium has the distinction of being the first dinoflagellate genus associated unequivocally as the source phycotoxins! ( 1960 ) a list of free-living dinoflagellate species in the world 's oceans M. MAFFT: new. Unequivocally as the source of new questions for experimental work be assessed by detecting the formation of zygotes in strains! Shellfish toxin production in the population dynamics and toxin production in marine dinoflagellates with comment on the toxic. The other hand, a complex life history and a period of calm weather is necessary G. Norris,,. Of survivors and autecological dependency alexandrium compressum for population studies ( ATANDA ) was two-fold more toxic than the cultured cf! Of UNESCO, Kuma M. MAFFT: a novel method for rapid multiple sequence alignment on. Its dormant stages, P. bahamense develop spherical, double-layered cysts for protection were collected from the northern southern... Genus Alexandrium Halim ( 1960 ) a list of free-living dinoflagellate species in the population dynamics of many Alexandrium (. Dinoflagellate blooms by serial parasitic killers, Hallegraeff GM, editors a by the planktonic, chain-forming dinoflagellate humic of... Camp alexandrium compressum, Orr RJS, Kellmann R, Murray SA, Neilan BA, Geyer WR, RP. Pattern amongst species 1999 ) reported an increase in A. catenella from Hong Kong waters siu.
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