Dinocyst
Dinocysts or dinoflagellate cysts are typically 15 to 100 µm in diameter and produced by around 15–20% [
History
The first person to recognize fossil dinoflagellates was
A first relation between dinoflagellate thecae and cysts was made through morphological comparison of both by Bill Evitt and Susan E. Davidson.[2] Further evidence came from detailed culture studies of dinoflagellate cysts by David Wall and Barrie Dale at Woods Hole Oceanographic Institution in the sixties.[3][4]
Types of cysts
Ontologically, the term cyst can apply to (1) a temporary resting state (pellicle, temporary or ecdysal cyst), (2) a dormant zygote (resting cysts or hypnozygotes) or (3) a coccoid condition in which the cells are still photosynthetically active.[5] For example, for this last special case, all cysts described from species of the order Phytodiniales (e.g. Cystodinium, Stylodinium, Hypnodinium, Tetradinium, Dinococcus, Gloeodinium), are coccoid stages.
Digestive cyst or digestion cysts denote pellicle cysts formed after feeding by phagocytosis as in Katodinium fungiforme.[6][7]
Division cysts refer to non-motile division stages wherein asexual reproduction takes place through division.[8] These are not pellicle or resting cysts since they are not dormant. Similarly, palmelloid or mucilage stages are not pellicle or resting cysts, but stages in which the monad loses its flagella and becomes enveloped in multilayered mucilage wherein division takes place.[9]
Taxonomy
Dinoflagellate cysts described in the literature have been linked to a particular motile stage through morphological similarities and/or co-occurrence in the same population/culture or through the technique of establishing the so-called cyst-theca relation by incubation of the cysts.
Size
Quaternary dinocysts are typically between 15 and 100 µm in diameter.[39] One of the smallest recent cysts is the cyst of Pentapharsodinium dalei, which can be as small as 19 µm in length.[40] One of the largest recent cysts is the cyst of Protoperidinium latissimum, which can be as large as 100 µm in length.[4]
Composition
The walls of organic-walled dinocysts are composed of the resistant biopolymer called
In addition to organic-walled cysts, there are also calcareous dinoflagellate cysts and siliceous dinoflagellate cysts.
Morphological terms
In pure morphological terms, a dinocyst can be described as the body formed by the cyst wall, as well as the space it encloses and all the spaces within it.[42] Cysts may develop their wall immediately within the theca, and such cysts are called proximate. Alternatively, the cyst may comprise a more or less spherical central body with processes or crests, and such cysts are termed chorate or proximochorate. Cysts may have a single-layered wall (autophragm), a two-layered wall (comprising an outer periphragm and an inner endophragm) or a three-layered wall (ectophragm, periphragm and endophragm if the outer wall is structurally supported, or otherwise periphragm, mesophragm and endophragm). Cysts with two or more wall layers that define a cavity are termed cavate. Excystment usually results in loss of part of, or an opening in, the cyst wall, termed archeopyle, the shape and position of which may indicate the position and/or shape of one or more thecal plates.[20]
Transmission electron microscopy (TEM) studies (e.g.[43]) suggest that endophragm and periphragm are not morphologically separable. Therefore, the use of the terms pedium and luxuria are suggested instead.[44] Within the cyst wall, a thick cellulose-like layer called the endospore is present which is birefringent under crossed nichols.[45] Cysts may be identified using the overall body shape but more often based on the characteristic furrows housing the flagella (cingulum and sulcus) or details of the patterns of plates covering many motiles (thecal tabulation). The one distinctive feature common to all cysts is the excystment opening (archaeopyle) through which the emerging new motile stage exits. In many cases this reflects a recognizable part of the tabulation (one or more plates). However, one large group of dinoflagellates (athecate - or naked dinoflagellates) do not have thecal plates and therefore produce cysts lacking all forms of reflected tabulation.[46]
Cyst ultrastructure
There have been very few ultrastructural studies of marine cysts with TEM, except for early on Hystrichosphaea bentorii, on Hystrichosphaeridium, Impletosphaeridium, Lingulodinium machaerophorum and Operculodinium centrocarpum and Bitectatodinium tepikiense[43][47][48] and more recent work on Lingulodinium machaerophorum[49] and Alexandrium.[50]
Some freshwater cysts have been investigated with TEM, such as Ceratium hirundinella.[51]
Relation to life cycle
Resting cysts are traditionally associated with the sexual cycle of dinoflagellates.[52] Induced by particular triggers such as changes in temperature, nutrients,[53] etc., dinoflagellates undergo gamete formation. The gametes fuse to form the planozygote and undergo encystment: they form cysts within the thecae of the planozygote. These rapidly sink to the sediment. Many species may spend longer periods resting in the sediment than active in the water column.[54] Resting stages also constitute a reservoir of genetic diversity, which increases the survival potential of the populations.[55] Thus, dinoflagellate cysts have great ecological importance and act as "seed banks", comparable to those found in terrestrial ecosystems. The encysted forms may remain viable for up to 100 years.[56] Sediment can be stored with live Lingulodinium cysts for at least 18 months.[57] Cysts often need triggers to germinate ('excyst'), such as changes in temperature, nutrients, etc. Some cysts, such as Scrippsiella acuminata, require light to germinate.[58]
Distribution and ecology of organic-walled dinocysts
Dinocyst distribution is mainly studied through studies of surface sediments.[59] Many studies are regional, such as the Iberian Margin[60] the North Sea,[61] Kiel bight,[62] Celtic Sea,[63] Norwegian Sea,[64] around Iceland,[65] the Southeast Pacific,[66] the Arctic,[67][68] Equatorial Atlantic,[69] South and Equatorial Atlantic,[70] off West Africa,[71] the Southern Ocean,[72] Benguela upwelling,[73] in the Mediterranean Sea,[74] Caspian Sea,[75] British Columbia,[76] The Northeastern Pacific,[77] Florida,[78] Mexico[79] and Barends Sea.[80]
Such surface sediment studies show that dinoflagellate cyst distribution is controlled by ranges of temperature, salinity and nutrients.[81] This often poses biogeographical boundaries, more particularly temperature.[82] Some species can be clearly related to cold waters.[83] Recent molecular work has shown the presence of such cold-water indicator, a life-stage of Islandinium sp. in Canadian sea-ice for the first time.[84] Other species are thermophilic, such as the "living fossil" Dapsilidinium pastielsii currently found in the Indo-Pacific Warm Pool only.[85]
Eutrophication can also be reflected in dinocyst assemblages.[86][87][88]
Cysts can be transported via ocean-currents, which can distort ecological signals. This has been documented for the warm water species Operculodinium israelianum and Polysphaeridium zoharyi which were interpreted to have been transported along the Southern coast of the United States.[59] Cyst are also often transported from the inner shelf to the outer shelf or slope.[59]
Another problem with cysts is that they also get transported with
Seasonality and fluxes are studied through sediment trap studies, which help to understand ecological signals.[90][91][92][93][94][95]
Palaeoecology of organic-walled dinocysts
The
Such reconstructions can be done via semi-quantitative techniques, such as ordination techniques,[46] which can indicate trends in environmental parameters.
A quantitative method is the use of transfer functions,[111][112][113][114][115] although these have been heavily debated.[116][117]
Another late Quaternary application is for environmental goals, more particularly the study of eutrophication[118][119][120] .[121]
An interval of particular interest during the late
Also during the Neogene, dinocysts have shown to be useful in the Miocene[127] and particularly the Messinian.[128] Also the paleoclimate of the Pliocene has been investigated.[129][130][131] Transfer functions have also been attempted during the Pliocene.[132] Some species have been suggested to have different environmental preferences during the Neogene.[133]
The
Morphological variation of organic-walled dinocysts
There is little known about how organic-walled dinocysts are formed except from culture experiments.[138] Cyst formation is suggested to happen through self-assembly processes.[139]
Organic-walled dinocyst morphology is shown to be controlled by changes in salinity and temperature in some species, more particularly process length variation. This is known to be the case for
The morphological variation can be applied for the reconstruction of salinity, in a semi-quantitative[147] or quantitative way.[142] Process length variation of Lingulodinium machaerophorum has been used to reconstruct Black Sea salinity variation.[148]
Biostratigraphy and evolution of organic-walled dinocysts
Organic-walled dinoflagellate cysts have a long geological record with lowest occurrences during the mid
The fossil record supports a major adaptive radiation of dinoflagellates during later Triassic and earlier Jurassic times. The majority of living thecate dinoflagellates can be interpreted as having either a peridinalean or gonyaulacalean tabulation, and that these tabulations, and hence the orders Gonyaulacales and Peridiniales, have been separate since at least the Early Jurassic.[20] The biostratigraphical application of dinoflagellate cysts has been thoroughly studied.[155][156] The Pliocene has been recently investigated[157][158] and also the Miocene.[159]
Palynological methods
Organic-walled dinoflagellate cysts are extracted using
Biological functions
Dinocysts are suggested to have a number of adaptive functions including survival during adverse conditions, bloom initiation and termination, dispersal in time, a seed bank for genetic diversity and dispersal in space.[168][169][170]
References
- ^ W.A.S. Sarjeant, 2002. 'As chimney-sweeps, come to dust': a history of palynology to 1970. pp. 273–327 In: Oldroyd, D. R. The earth inside and out: some major contributions to geology in the twentieth century. Geological Society (London) Special Publication no. 192.
- ^ Evitt, W.R. and Davidson, S.E. 1964. Dinoflagellate studies. 1. Dinoflagellate cysts and thecae. Stanford university publications X (1), pp. 3–12.
- S2CID 4267273.
- ^ JSTOR 1484690.
- ^ Pfiester L.A. & Anderson D.M. 1987. Dinoflagellate reproduction. In: The biology of dinoflagellates. Botanical monographs 21 (Ed. by F.J.R. Taylor), pp. 611–648., Blackwell Scientific Publications.
- JSTOR 1485525.
- S2CID 86726034.
- ^ BRAVO I., FIGUEROA R.I., GARCÉS E., FRAGA S. & MASSANET A. 2010. The intricacies of dinoflagellate pellicle cysts: the example of Alexandrium minutum cysts from a bloom-recurrent area (Bay of Baiona, NW Spain). Deep-Sea Research Part II: Topical Studies in Oceanography 57: 166–174.
- ^ POPOVSKÝ J. & PFIESTER L.A. 1990. Dinophyceae (Dinoflagellida). In: Süßwasserflora von Mitteleuropa. Begründet von A. Pascher. Band 6 (Ed. by H. Ettl,J. Gerloff,H. Heynig. & D. Mollenhauer). Gustav Fischer Verlag, Jena, 272 pp.
- S2CID 4267273.
- S2CID 84402741.
- .
- .
- .
- S2CID 85691284.
- S2CID 84378350.
- S2CID 82954522.
- ^ HEAD M.J. 1996. Modern dinoflagellate cysts and their biological affinities. In: Palynology: principles and applications (Ed. by J. Jansonius & D. C. McGregor), pp. 1197–1248. American Association of Stratigraphic Palynologists Foundation, Dallas, Texas.
- ^ S2CID 86845462.
- ^ a b c d Fensome, R.A.; Taylor, F.J.R.; Norris, G.; Sarjeant, W.A.S.; Wharton, D.I.; Williams, G.L. (1993). "A classification of living and fossil dinoflagellates". American Museum of Natural History, Micropaleontology, Special Publication. 7: 1–351.
- ^ Harland, R (1982). "A review of Recent and Quaternary organic-walled dinoflagellate cysts of the genus Protoperidinium". Palaeontology. 25: 369–397.
- ^ Schilling, A.J. (1891). "Die Süsswasser-Peridineen". Flora Oder Allgemeine Botanische Zeitung. 74: 220–299.
- S2CID 83927778.
- S2CID 86137363.
- ^ Evitt, W.R., Lentin, J.K., Millioud, M.E., Stover, L.E. and Williams, G.L., 1977. Dinoflagellate cyst terminology. Geological survey of Canada, Paper 76-24, 1-11.
- ^ Rochon, A., de Vernal, A., Turon, J.-L., Matthiessen, J., and Head, M.J., 1999. Distribution of recent dinoflagellate cysts in surface sediments from the North Atlantic Ocean and adjacent seas in relation to sea-surface parameters. AASP Contribution Series, 35, 146 pp.
- ^ MATSUOKA, K. & FUKUYO, Y. 2000. Technical guide for modern dinoflagellate cyst study. WESTPAC-HAB/WESTPAC/IOC, Japan Society of the Promotion Science, Tokyo, 29 pp.
- .
- .
- ^ Soliman, A., Head, M.J., and Louwye, S. In press. Morphology and distribution of the Miocene dinoflagellate cyst Operculodinium? borgerholtense Louwye 2001, emend. Palynology.
- ^ Head, M.J., 1999. The Late Pliocene St. Erth Beds of Cornwall: a review of the palynology and reappraisal of the dinoflagellates. In: Scource, J. and Furze, M.F.A. (eds.), The Quaternary of West Cornwall. Field Guide, Quaternary Research Association, Durham, U.K., p. 88–92.
- ^ Head, M.J. 2000. Geonettia waltonensis, a new goniodomacean dinoflagellate from the Pliocene of the North Atlantic region, and its evolutionary implications" Journal of Paleontology 74(5): 812–827, 6 pls.
- S2CID 131328767.
- S2CID 54847659.
- .
- .
- S2CID 83682507.
- .
- )
- .
- ^ Fensome, R.A., Taylor, F.J.R., Norris, G., Sarjeant, W.A.S., Wharton, D.I., and Williams, G.L., 1993. A classification of modern and fossil dinoflagellates, Sheridan Press, Hanover. .
- ^ DE, VERTEUIL L.; Norris, G. (1996). "Part 2. Homology and structure in dinoflagellate cyst terminology". Micropaleontology. S42: 83–172.
- ^ a b Jux, U (1968). "Über den feinbau der wandung bei Hystrichosphaera bentori Rossignol 1961". Palaeontographica Abteilung B. 123 (1–6): 147–152.
- JSTOR 1485937.
- .
- ^ a b Dale, B. & Dale, A.L. 2002. Environmental applications of dinoflagellate cysts and acritarchs . In Quaternary environmental micropalaeontology (Haslett, S.K., editor), 207-240. Arnold, London.
- ^ Jux, U (1971). "Über den feinbau der wandungen einiger Tertiärer Dinophyceen-zysten und Acritarcha Hystrichosphaeridium, Impletosphaeridium, Lingulodinium". Palaeontographica, Abt. B. 132 (5–6): 165–174.
- ^ Jux, U (1976). "Über den feinbau der wandungen bei Operculodinium centrocarpum (Deflandre & Cookson) Wall 1967 und Bitectatodinium tepikiense Wilson 1973". Palaeontographica, Abt. B. 155 (5–6): 149–156.
- .
- S2CID 84400812.
- S2CID 84164446.
- S2CID 9630941.
- ^ Pfiester, L. A. & Anderson, D. M. 1987. Dinoflagellate reproduction. In: The biology of dinoflagellates (ed. F. J. R. Taylor), pp. 611–648. - Blackwell, Oxford.
- ^ RENGEFORS K. 1998. Seasonal succession of dinoflagellates coupled to the benthic cyst dynamics in Lake Erken, Sweden. Archiv für Hydrobiologie, Special Issues, Advances in Limnology 51: 123–141.
- S2CID 28650477.
- PMID 21587228.
- .
- S2CID 4281882.
- ^ .
- S2CID 128824886.
- .
- .
- .
- ^ Matthießen, J. (1995) Distribution patterns of dinoflagellate cysts and other organic-walled microfossils in recent Norwegian-Greenland Sea sediments , Marine Micropaleontology
- .
- .
- .
- ^ Matthiessen, J., De Vernal, A., Head, M., Okolodkov, Y., Ángel, P., Zonneveld, K. and Harland, R. Modern organic-walled dinoflagellate cysts in Arctic marine environments and their (paleo-) environmental significance. Paläontologische Zeitschrift 79(1): 3-51.
- PMID 11134709.
- ^ Vink, A., Baumann, K-H., Böckel, B., Esper, O., Kinkel, H., Volbers, A., Willems, H., Zonneveld, K.A.F. Coccolithophorid and dinoflagellate synecology in the South and Equatorial Atlantic: Improving the paleoecological significance of phytoplankton microfossils. In: Wefer, G., Mulitza, S. and Ratmeyer, V. (eds.) The South Atlantic in the Late Quaternary: reconstruction of material budgets and current systems. Springer, Berlin: 121-142.
- .
- ^ Oliver Esper, Karin Zonneveld. The potential of organic-walled dinoflagellate cysts to reconstruct past sea-surface conditions in the Southern Ocean" Marine Micropaleontology 63 (3/4): 185-212.
- .
- ^ Elshanawany, R., Zonneveld, K.A.F., Ibrahim, M.I. and Kholeif, S.E.A. (2010). Distribution patterns of recent organic-walled dinoflagellate cysts in relation to environmental parameters in the Mediterranean Sea. Palynology
- .
- .
- .
- S2CID 82533706.
- .
- ^ Solignac, S.; Grøsfjeld, K.; Giraudeau, J.; de Vernal, A. (2009). "Distribution of recent dinocyst assemblages in the western Barents Sea". Norwegian Journal of Geology. 89 (1–2): 109–119.
- hdl:1854/LU-3226112.
- ^ Dale, B., 1996. Dinoflagellate cyst ecology: modelling and geological applications. In Jansonius, J. & McGregor, D.C. (eds.): Palynology: Principles and Applications, volume 3, 1249-1275, AASP Foundation, Dallas.
- ^ Head, M.J., Harland, R., and Matthiessen, J. 2001. Cold marine indicators of the late Quaternary: the new dinoflagellate cyst genus Islandinium and related morphotypes. Journal of Quaternary Science, 16(7): 621–636, 3 pls.
- S2CID 30085938.
- doi:10.1130/G35456.1.
- .
- .
- .
- .
- .
- .
- .
- .
- S2CID 83493908.
- .
- .
- S2CID 53454455.
- S2CID 129655535.
- .
- S2CID 52023457.
- .
- .
- S2CID 129877701.
- .
- PMID 10708791.
- .
- .
- .
- ^ de Vernal, A., Rochon, A., 2011. Dinocysts as tracers of sea-surface conditions and sea-ice cover in polar and subpolar environments, IOP Conference Series: Earth and Environmental Science, 14, 012007.
- ISBN 9781862391604.
- S2CID 140191345.
- .
- ^ Guiot, J., de Vernal, A., 2007. Transfer functions: methods for quantitative paleoceanography based on microfossils, In Hillaire-Marcel and de Vernal (eds.) Proxies in Late Cenozoic Paleoceanography, Elsevier, pp. 523–563.
- hdl:1885/52333.
- S2CID 54951606.
- ^ Telford, R.J., 2006. Limitations of dinoflagellate cyst transfer functions. Quaternary Science Reviews 25 : 1375-1382.
- .
- PMID 11213194.
- .
- PMID 22118910.
- .
- .
- S2CID 128814429.
- .
- ^ Van Nieuwenhove, N., Bauch, H.A., Matthiessen, J., 2008. Last Interglacial surface water conditions in the eastern Nordic Seas inferred from dinocyst
- S2CID 128434842.
- ^ Louwye, S., Foubert, A., Mertens, K.N., Van Rooij, D. & IODP Expedition 307 scientific party (2007). Integrated stratigraphy and palaeoecology of the Lower and Middle Miocene of the Porcupine Basin. Geological Magazine 145, 321-344.
- ^ Popescu, S.-M.; Dalesme, F.; Jouannic, G.; Escarguel, G.; Head, M.J.; Melinte-Dobrinescu, M.C.; Sütö-Szentai, M.; Bakrac, K.; Clauzon, G.; Suc, J.-P. "Galeacysta etrusca Corradini & Biffi 1988, dinoflagellate cyst marker of Paratethyan influxes into the Mediterranean Sea before and after the peak of the Messinian Salinity Crisis". Palynology.
- ^ Head, M.J. and Westphal, H. 1999. Palynology and paleoenvironments of a Pliocene carbonate platform: the Clino Core, Bahamas" Journal of Paleontology 73(1): 1–25, 8 pls.
- S2CID 128540933.
- ^ De Schepper S, Head MJ, Groeneveld J (2009) North Atlantic Current variability through marine isotope stage M2 (circa 3.3 Ma) during the mid-Pliocene. Paleoceanography 24:PA4206
- .
- .
- .
- S2CID 128843669.
- .
- S2CID 128763506.
- .
- .
- ^ Hallett, R.I., 1999. Consequences of environmental change on the growth and morphology of Lingulodinium polyedrum (Dinophyceae) in culture. PhD thesis. University of Westminster, 109 pp.
- .
- ^ S2CID 83160220.
- S2CID 128999348.
- S2CID 84740337.
- .
- .
- S2CID 129675846.
- ^ Mertens, K.N., Bradley, L.R., Takano, Y., Mudie, P.J., Marret, F., Aksu, A.E., Hiscott, R.N., Verleye, T.J., Mousing, E.A., Smyrnova, L.L., Bagheri, S., Mansor, M., Pospelova, V. & Matsuoka, K. (in press). Quantitative estimation of Holocene surface salinity variation in the Black Sea using dinoflagellate cyst process length. Quaternary Science Reviews
- doi:10.1139/b96-205.
- ^ Moldowan, J.M. and Talyzina, N.M., Biogeochemical evidence for dinoflagellate ancestors in the Early Cambrian. Science 281, 1168-1170.
- .
- ^ LeHerissé, A., Masure, E., Al Ruwaili, M., Massa, D., 2000. Revision of Arpylorus antiquus from the Silurian: the end of a myth. In: Wang, W., Quyang, S., Sun, X., Yu, G. (Eds.), Abstracts 10th International Palynological Congress, Nanjing. National Natural Science Foundation of China, p. 88.
- ^ Vozzhennikova, T.F., Shegeshova, L.I., 1989. Palaeodinophysis gen. et sp. N. from the Devonian of the Rudnyy Altay (a unique find of dinoflagellate fossils), Doklady Akademii Nauk SSSR 307, 442–445 (in Russian).
- .
- ^ Powell, A. J. (ed.), 1992: A Stratigraphic Index of Dinoflagellate Cysts. London: Chapman & Hall, 300 pp.
- ^ Williams, G.L., Stover, L.E., & Kidson, E.J., 1993: Morphology and stratigraphic ranges of selected Mesozoic-Cenozoic dinoflagellate taxa in the northern hemisphere. Geological Survey of Canada, Paper. 92-10 , 137 pp., 2 pl.
- S2CID 55267190.
- S2CID 131306285.
- .
- .
- S2CID 128773929.
- .
- .
- S2CID 86038878.
- ^ Stockmarr, J (1971). "Tablets with spores used in absolute pollen analysis". Pollen et Spores. 13: 615–621.
- .
- .
- .
- S2CID 85372383.
- ^ FRYXELL G.A. 1983. Introduction. In: Survival strategies of the algae (Ed. by A. Fryxell), pp. 1–22, Cambridge University Press, Cambridge, U.K.
External links
- AASP — THE PALYNOLOGICAL SOCIETY
- L'Association des Palynologues de Langue Française
- Canadian Association of Palynologists
- The Micropalaeontological Society, Palynology group
- Quaternary Dinoflagellate Cyst Association
- Dinoflaj2 Rob Fensome's and Graham Williams' Database on cysts
- Yasuo Fukuyo's site on motile stages and their cysts
- Dino6 meeting
- Dino8 meeting, Montreal Archived 15 October 2008 at the Wayback Machine
- Dino9 meeting, Liverpool
- Marine Micropaleontology (Journal)
- Review of Palaeobotany and Palynology (Journal)
- Journal of Micropalaeontology (Journal)
- Micropaleontology (Journal)