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Skeletonema costatum

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Skeletonema costatum
Scientific classification Edit this classification
Domain: Eukaryota
Clade: Diaphoretickes
Clade: SAR
Clade: Stramenopiles
Phylum: Gyrista
Subphylum: Ochrophytina
Class: Bacillariophyceae
Order: Thalassiosirales
Family: Skeletonemataceae
Genus: Skeletonema
Species:
S. costatum
Binomial name
Skeletonema costatum
(Greville) Cleve, 1866

Skeletonema costatum is a centric diatom that belongs to the genus Skeletonema[1].

Skeletonema costatum was first described by R. K. Greville in 1866, originally called Melosira costata[1], later renamed by Cleve in 1873[2], and more narrowly defined by Zingone et al.[3] and Sarno et al.[4][5]. Skeletonema costatum is the most well known species of Skeletonema among planktologists[4]. Skeletonema costatum is often one of the dominant species responsible for red tide events[6].

Structure and morphology

A single chain of Skeletonema costatum cells from a field sample from Long Island Sound.

S. costatum is a single-celled organism that exists in long chains of between 6 to 24 cells[7] (or up to 60 cells[3]), with between 3.5 to 11.5 μm between cells, as recorded by Castille (1995)[7]. Cells are connected by long straight fultoportula processes[8].

Each cell is between 8 to 12 μm in diameter[5] and contain up to 2 chloroplasts[7].

S. costatum have cylindrically shaped cells and a ring of long fultoportulae processes along the periphery of each valve[9].

Relative to the Skeletonema genus, S. costatum is unique in its "flattened IFPPs [intercalary fultoportula process] closed along their whole length, with no apparent external suture and with a hole at their base"[3]. Although Sarno et al., (2007)[5] suggests they have a longitudinal suture extending from the external hole at the base of their intercalary fultoportula processes to its tip.

S. costatum have terminal fultoportulae processes with claw-like tips [5]. Their intercalary fultoportula processes of adjacent valves are connected at a 1:2 junction, where each intercalary fultoportula process interlocks with two intercalary fultoportula processes, creating a "zigzag appearance"[5]. Their intercalary rimoportula are positioned marginally, they have long terminal rimoportula, and their cingular band features rows of pores[5]. Each valve has one of their fultoportulae replaced with a rimoportula, identified by its longer external process and the "spoutlike teapot" shaped tip of terminal rimoportulae[5].

Among the Skeletonema species, S. costatum is most morphologically similar to S. subsalsum, both exhibiting rows of small pores between parallel rows of transverse branching ribs on their girdle bands[5]. They are also the only two Skeletonema species with long intercalary rimoportula processes[5]. S. costatum can be identified by the persistent presence of a 1:2 junction, and the closed tubules of its intercalary fultoportulae processes[5].

Morphological variation

The morphological plasticity of S. costatum has been extensively studied[10][11][12][7]. Castillo (1995) attributes significant variations in morphological features, such as cell diameter, the number of cells per chain and the length of intercalary processes, with variations in environmental conditions, most notably, salinity[7]. If cultured in freshwater, S. costatum develops short intercellular processes[13], and at 1 psu, are observed with seemingly no space between sibling valves[5].

Taxonomy

As of 2021, 21 Skeletonema species were "identified and taxonomically accepted", Skeletonema costatum being one of them[14].

S. costatum was first described by R. K. Greville in 1866, originally called Melosira costata[1], and later renamed by Cleve in 1873[2]. S. costatum has since been more narrowly defined, with numerous species previously attributed to S. costatum identified as distinct species[3][15][4][5].

The species originally described by Greville in 1866 is often referred to as S. costatum sensu lato (s. l.)[16][14], representing multiple different species with similar morphological traits[5][3][4]. The species granted the original epithet, costatum, was the species more narrowly described by Zingone et al. (2005) after reexamination of the type materials of S. costatum using electron microscopy and molecular analysis of rDNA[3]. Zingone et al., (2005) identified two distinct morphologies within the type material, describing the less abundant morphology as S. grevillei and the more abundant morphology as the original epithet, costatum[3]. The latter was assigned the original epithet due to its closer similarity to the specimen originally described by Greville, which he had conveniently marked[3]. The two morphologies differed in their frustule ultrastructures; including "the shape of FPPs [fultoportula processes], the type of interlocking between IFPPs [intercalary fultoportula processes] of sibling valves, and the cingular band ornamentation"[3]. S. costatum sensu stricto (s. s.) can be used to describe the more narrowly defined S. costatum species to differentiate it from S. costatum sensu lato[16]. S. Costatum sensu stricto (s. s.) has also been referred to as S. costatum (Greville) Cleve emend. Zingone and Sarno[17][13].

Physiology

Distribution and habitat

Skeletonema costatum are found widely geographically distributed (apart from the Antarctic Ocean)[16], found around the world, including off the coasts of Hong Kong Island[1], Florida, USA, Uruguay, Brazil[5], Northern Queensland (Australia)[16], China, and the Sea of Japan[17][18].

S. costatum primarily reside in the neritic zone and are commonly found in brackish waters as opposed to the more oceanic, S. tropicum[7]. S. costatum are frequently the dominant phytoplankton species responsible for red tide events[6][19][20]. Although the dominant species responsible for red tide events in a given area can evolve over time, CITE

Red tide outbreaks in Yangtze River Estuary

S. costatum is one of the dominant species responsible for red tide outbreaks[6]. Of the red tide outbreaks in the Yangtze River Estuary between 1972 and 2009, S. costatum occurred during 20% of the 174 recorded outbreaks[6]. Over 50% of the outbreaks in this area occurred during May[6]. The frequency of red tide outbreaks in this area coincided with the concentration of dissolved inorganic nitrogen. and PO4-P concentrations in this area, both increasing after 2002[6]. After 2000, larger outbreaks of areal extent larger than 1000km2 became more common[6].

Growth and environmental conditions

Temperature

A temperature range of 10 to 30°C was observed to support the growth of S. costatum, but members of this species grow optimally at 25°C [13]. The strains of S. costatum from the Sea of Japan off the coast of Dokai Bay prefer warmer temperatures, only collected from water above 20°C[17]. At this temperature, their specific growth rates were measured as above 1.0 d-1[17].

Salinity

Although S. costatum has exhibited preference for lower salinities, it can grow in salinity conditions of 0 to 35 psu [13]. As such, S. costatum can thrive in a variety of ocean environments ranging from oceanic to marine estuary and even riverine environments [21][22]. The salinity tolerance of S. costatum is especially ideal in estuarian waters where salinities fluctuate, thus making this diatom one of its dominant planktonic species[13].

Light levels

At 20°C, S. costatum can grow under irradiance of 7 to 406 μmol/m2s [23]. Photoinhibition is observed at 700 μmol/m2s, at which point a decrease in photosynthetic ability of the diatom is detected [23].

Nutrients

S. costatum blooms in eutrophic waters that are often loaded with nitrogen, phosphorus, and other nutrients/minerals in both dissolved and particulate forms [24]. The eutrophic conditions that house S. costatum blooms are often limited in carbon dioxide, as there is heavy competition for this limiting resource.[25] It has been shown that nitrate enrichment and waters high in nitrate have the ability to stimulate S. costatum growth through the action of enhanced competitive photosynthetic activity in a CO2 limited environment.[26] Phosphate rich waters have also proven to have a stimulating effect on S. costatum growth rates in CO2-limited environments.[25] These studies( [26],[25]) also indicate that high concentrations of nitrates and phosphates increase the amount of inorganic carbon in the form of bicarbonate fixed by S. costatum.

Major viral pathogens

Skeletonema costatum-infecting virus (ScosV)

Skeletonema costatum-infecting virus (ScosV) is a novel virus that lyses S. costatum, specifically strain ME-SCM-1, upon infection [27]. It was first isolated in 2008 from seawater samples taken in Jaran Bay, South Korea, and was characterized in 2015 as having an icosahedral shape and a diameter of approximately 40 to 50 nm [27][28].

Ecological significance

Marine diatoms account for about 20% of our planet's primary production [29],

Human applications

Production of biofuels:

Production of natural products:

References

  1. ^ a b c d GREVILLE, R. K. (1866). "Descriptions of New and Rare Diatoms. Series XVIII". Transactions of The Microscopical Society & Journal. 14 (1): 1–9. doi:10.1111/j.1365-2818.1866.tb05059.x. ISSN 0962-7375.
  2. ^ a b Cleve, P. T. (1873). "Examination of diatoms found on the surface of the sea of Java". Bih. Kongl. Svenska Vetensk.-Akad. Handl. 11: 3–13. {{cite journal}}: line feed character in |title= at position 44 (help)
  3. ^ a b c d e f g h i Zingone, Adriana; Percopo, Isabella; Sims, Pat A.; Sarno, Diana (2005). "DIVERSITY IN THE GENUSSKELETONEMA(BACILLARIOPHYCEAE). I. A REEXAMINATION OF THE TYPE MATERIAL OFS. COSTATUMWITH THE DESCRIPTION OFS. GREVILLEISP. NOV". Journal of Phycology. 41 (1): 140–150. doi:10.1111/j.1529-8817.2005.04066.x. ISSN 0022-3646.
  4. ^ a b c d Sarno, Diana; Kooistra, Wiebe H. C. F.; Medlin, Linda K.; Percopo, Isabella; Zingone, Adriana (2005-02). "DIVERSITY IN THE GENUS SKELETONEMA (BACILLARIOPHYCEAE). II. AN ASSESSMENT OF THE TAXONOMY OF S. COSTATUM -LIKE SPECIES WITH THE DESCRIPTION OF FOUR NEW SPECIES". Journal of Phycology. 41 (1): 151–176. doi:10.1111/j.1529-8817.2005.04067.x. {{cite journal}}: Check date values in: |date= (help)
  5. ^ a b c d e f g h i j k l m n Sarno, Diana; Kooistra, Wiebe H. C. F.; Balzano, Sergio; Hargraves, Paul E.; Zingone, Adriana (2007). "DIVERSITY IN THE GENUSSKELETONEMA(BACILLARIOPHYCEAE): III. PHYLOGENETIC POSITION AND MORPHOLOGICAL VARIABILITY OF SKELETONEMA COSTATUM AND SKELETONEMA GREVILLEI, WITH THE DESCRIPTION OF SKELETONEMA ARDENS SP. NOV". Journal of Phycology. 43 (1): 156–170. doi:10.1111/j.1529-8817.2006.00305.x. ISSN 0022-3646.
  6. ^ a b c d e f g Liu, Lusan; Zhou, Juan; Zheng, Binghui; Cai, Wenqian; Lin, Kuixuan; Tang, Jingliang (2013). "Temporal and spatial distribution of red tide outbreaks in the Yangtze River Estuary and adjacent waters, China". Marine Pollution Bulletin. 72 (1): 213–221. doi:10.1016/j.marpolbul.2013.04.002. ISSN 0025-326X.
  7. ^ a b c d e f Castillo, J. Aké; Meave del Castillo, M.E.; Hernández-Becerril, D.U. (1995). "Morphology and distribution of species of the diatom genus Skeletonema in a tropical coastal lagoon". European Journal of Phycology. 30 (2): 107–115. doi:10.1080/09670269500650871. ISSN 0967-0262.
  8. ^ Shevchenko, Olga G.; Ponomareva, Anna A.; Shulgina, Maria A.; Orlova, Tatiana Yu. (2019). "Phytoplankton in the Coastal Waters of Russky Island, Peter the Great Bay, Sea of Japan". Botanica Pacifica. doi:10.17581/bp.2019.08112.
  9. ^ Zingone, Adriana; Percopo, Isabella; Sims, Pat A.; Sarno, Diana (2005). "DIVERSITY IN THE GENUSSKELETONEMA(BACILLARIOPHYCEAE). I. A REEXAMINATION OF THE TYPE MATERIAL OFS. COSTATUMWITH THE DESCRIPTION OFS. GREVILLEISP. NOV". Journal of Phycology. 41 (1): 140–150. doi:10.1111/j.1529-8817.2005.04066.x. ISSN 0022-3646.
  10. ^ Castellví, Josefina (1969). "Long polar fibres of Skeletonema costatum (Grev.) Cleve". Proc. Int. Seaweed Symp. 6: 85–88.
  11. ^ Hasle, G. R. (1973). "Morphology and taxonomy of Skeletonema costatum (Bacillariophyceae)". Norw. J. BoG. 20: 109–137.
  12. ^ Gallagher, Jane C. (1982). "PHYSIOLOGICAL VARIATION AND ELECTROPHORETIC BANDING PATTERNS OF GENETICALLY DIFFERENT SEASONAL POPULATIONS OF SKELETONEMA COSTATUM (BACILLARIOPHYCEAE)". Journal of Phycology. 18 (1): 148–162. doi:10.1111/j.1529-8817.1982.tb03169.x. ISSN 0022-3646.
  13. ^ a b c d e Balzano, S.; Sarno, D.; Kooistra, W. H. C. F. (2011). "Effects of salinity on the growth rate and morphology of ten Skeletonema strains". Journal of Plankton Research. 33 (6): 937–945. doi:10.1093/plankt/fbq150. ISSN 0142-7873.
  14. ^ a b Liu, Shuya; Wang, Yichao; Xu, Qing; Zhang, Mengjia; Chen, Nansheng (2021). "Comparative analysis of full-length mitochondrial genomes of five Skeletonema species reveals conserved genome organization and recent speciation". BMC Genomics. 22 (1): 1–14. doi:10.1186/s12864-021-07999-z. ISSN 1471-2164.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  15. ^ Medlin, Linda K.; Elwood, Hille J.; Stickel, Shawn; Sogin, Mitchell L. (1991). "MORPHOLOGICAL AND GENETIC VARIATION WITHIN THE DIATOM SKELETONEMA COSTATUM (BACILLARIOPHYTA): EVIDENCE FOR A NEW SPECIES, SKELETONEMA PSEUDOCOSTATUM1". Journal of Phycology. 27 (4): 514–524. doi:10.1111/j.0022-3646.1991.00514.x. ISSN 0022-3646.
  16. ^ a b c d Kooistra, Wiebe H.C.F.; Sarno, Diana; Balzano, Sergio; Gu, Haifeng; Andersen, Robert A.; Zingone, Adriana (2008). "Global Diversity and Biogeography of Skeletonema Species (Bacillariophyta)". Protist. 159 (2): 177–193. doi:10.1016/j.protis.2007.09.004.
  17. ^ a b c d Yamada, Machiko; Katsuki, Eri; Otsubo, Mayuko; Kawaguchi, Mayumi; Ichimi, Kazuhiko; Kaeriyama, Hideki; Tada, Kuninao; Harrison, Paul J. (2010). "Species diversity of the genus Skeletonema (Bacillariophyceae) in the industrial harbor Dokai Bay, Japan". Japan Journal of Oceanography. 66 (6): 755–771. doi:10.1007/s10872-010-0062-4. ISSN 0916-8370.
  18. ^ Shevchenko, Olga G.; Ponomareva, Anna A.; Shulgina, Maria A.; Orlova, Tatiana Yu. (2019). "Phytoplankton in the Coastal Waters of Russky Island, Peter the Great Bay, Sea of Japan". Botanica Pacifica. doi:10.17581/bp.2019.08112.
  19. ^ Baohong, Chen; Kang, Wang; Xu, Dong; Hui, Lin (2021). "Long-term changes in red tide outbreaks in Xiamen Bay in China from 1986 to 2017". Estuarine, Coastal and Shelf Science. 249: 107095. doi:10.1016/j.ecss.2020.107095. ISSN 0272-7714.
  20. ^ Li, Chunqiang; Zhu, Baibi; Chen, Hong; Liu, Zhixin; Cui, Baiming; Wu, Jingrui; Li, Bin; Yu, Haichuan; Peng, Ming (2009). "The Relationship between the Skeletonema costatum Red Tide and Environmental Factors in Hongsha Bay of Sanya, South China Sea". Journal of Coastal Research. 253: 651–658. doi:10.2112/07-0967.1. ISSN 0749-0208.
  21. ^ Tian, Yan; Mingjiang, Zhou; Peiyuan, Qian (2002-09-01). "Combined effects of temperature, irradiance and salinity on growth of diatomSkeletonema costatum". Chinese Journal of Oceanology and Limnology. 20 (3): 237–243. doi:10.1007/BF02848852. ISSN 1993-5005.
  22. ^ Castillo, J. Aké; Meave del Castillo, M.E.; Hernández-Becerril, D.U. (1995-05-01). "Morphology and distribution of species of the diatom genus Skeletonema in a tropical coastal lagoon". European Journal of Phycology. 30 (2): 107–115. doi:10.1080/09670269500650871. ISSN 0967-0262.
  23. ^ a b Kaeriyama, Hideki; Katsuki, Eri; Otsubo, Mayuko; Yamada, Machiko; Ichimi, Kazuhiko; Tada, Kuninao; Harrison, Paul J. (2011-05-01). "Effects of temperature and irradiance on growth of strains belonging to seven Skeletonema species isolated from Dokai Bay, southern Japan". European Journal of Phycology. 46 (2): 113–124. doi:10.1080/09670262.2011.565128. ISSN 0967-0262.
  24. ^ Huo, Wen Yi; Shu, Jian Jun (2005). "Outbreak of skeletonema costatum bloom and its relations to environmental factors in Jiaozhou Bay, China". {{cite journal}}: Cite journal requires |journal= (help)
  25. ^ a b c Gao, Guang; Xia, Jianrong; Yu, Jinlan; Fan, Jiale; Zeng, Xiaopeng (2018-08-17). "Regulation of inorganic carbon acquisition in a red tide alga (Skeletonema costatum): the importance of phosphorus availability". Biogeosciences. 15 (16): 4871–4882. doi:10.5194/bg-15-4871-2018. ISSN 1726-4170.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  26. ^ a b Gao, Guang; Xia, Jianrong; Yu, Jinlan; Zeng, Xiaopeng (2018-02-01). "Physiological response of a red tide alga (Skeletonema costatum) to nitrate enrichment, with special reference to inorganic carbon acquisition". Marine Environmental Research. 133: 15–23. doi:10.1016/j.marenvres.2017.11.003. ISSN 0141-1136.
  27. ^ a b Kim, JinJoo; Kim, Chang-Hoon; Youn, Seok-Hyun; Choi, Tae-Jin (2015-6). "Isolation and Physiological Characterization of a Novel Algicidal Virus Infecting the Marine Diatom Skeletonema costatum". The Plant Pathology Journal. 31 (2): 186–191. doi:10.5423/PPJ.NT.03.2015.0029. ISSN 1598-2254. PMC 4454000. PMID 26060438. {{cite journal}}: Check date values in: |date= (help)
  28. ^ Short, Steven M.; Staniewski, Michael A.; Chaban, Yuri V.; Long, Andrew M.; Wang, Donglin (2020). "Diversity of Viruses Infecting Eukaryotic Algae". Current Issues in Molecular Biology: 29–62. doi:10.21775/cimb.039.029.
  29. ^ Field, Christopher B.; Behrenfeld, Michael J.; Randerson, James T.; Falkowski, Paul (1998-07-10). "Primary Production of the Biosphere: Integrating Terrestrial and Oceanic Components". Science. 281 (5374): 237–240. doi:10.1126/science.281.5374.237. ISSN 0036-8075.
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