Gromia

Gromia is a genus of protists, closely related to foraminifera, which inhabit marine and freshwater environments. Gromia are ameboid, producing filose pseudopodia that extend out from the cell’s proteinaceous test through a gap enclosed by the cell’s oral capsule. The test, a shell made up of protein that encloses the cytoplasm, is made up of several layers of membrane, which resemble honeycombs in shape — a defining character of this genus.

Gromia
Gromia (1) and some foraminiferans (2-7)
Scientific classification
Domain:
Eukaryota
(unranked):
SAR
(unranked):
Rhizaria
Phylum:
Cercozoa
Class:
Gromiidea
Order:
Gromiida
Family:
Gromiidae
Genus:
Gromia

Dujardin, 1835

Gromia were first discovered in shallow waters, with members of the best-characterized species Gromia oviformus often found inhabiting rock surfaces, sediments or seaweed holdfasts [1]. However, research from the 1990s and early 2000s identified gromiids inhabiting depths up to 4,392 m, leading to several new deep-sea Gromia species being described and recognized [2]. A recent study of the deep sea species Gromia sphaerica revealed that it produces traces on the seafloor which resemble the fossils of early Bilateria (animals with bilateral symmetry), which called into question whether such fossils can serve as reliable documentation of early animal diversification in the Precambian era [3]

Deep-sea gromiids have also been shown to be important for carbon cycling [4] and denitrification [5].

History of Knowledge

Gromia were first described in the 1835, with G. oviformis gaining prominence because it was often found in the intertidal zones on the British coast [6]. Initially, Gromia were regarded as members of Foraminifera or Filosea, as noted in a review by Cifelli (1990) [7]. Gromia became better characterized throughout the 1960s, when electron microscopy revealed more details on their morphology, including the honeycomb membranes [8] [9]. The first molecular studies involving Gromia, which sampled G. oviformis, used small subunit (SSU) ribosomal RNA genes and concluded that Gromia were members of Cercozoa, a large group of amoebae with tests and filose pseudopodia [10]. Follow-up studies on this group placed Gromia within the Gromiidea class, again based on SSU rRNA genes [11] Eventually, when molecular studies combined data from several genes — actin, polyubiquitin, RNA polymerase II and small subunit rRNA genes —Gromia was shown to be a sister group to Foraminifera [12]. Moreover, within the genus Gromia, studies of the small subunit ribosomal RNA genes of various deep sea gromiids has revealed species diversity within Gromia, with molecular data tending to correlate with distinct morphologies of the various species’ tests [13].

Gromia were long thought to only inhabit shallow waters, until samples from the Arabian Sea from depths below 1,000 m revealed the first deep sea gromiid — Gromia spherica [14]. Additional species of deep sea Gromia protists were later described in waters from the Arabian sea, the European Arctic sea, and off the coast of Antarctica, among other locations, and characterized both morphologically and through molecular studies of their small subunit rRNA genes [15].

Habitat and Ecology

Gromiids inhabit sediments or surfaces of flora in both shallow waters and the deep sea. The best characterized species of shallow-water Gromia, G. oviformis, inhabits intertidal zones and other regions of shallow waters, often found attached to rocks, kelp, weeds, Cladophora algae, or within sediments [16]. G. oviformis has been shown to tolerate a temperature range of 0 - 30°C.

Deep-sea gromiids have been found in the Arabian sea [17], off the coast of Antarctica and in the water of the Northwest Atlantic ocean [18]. They were often collected from the 1000-3,100 m range [19]. Oxygen levels in gromiid habitats tend to exceed 0.2ml/l and are therefore not limiting to the organisms’ growth. The temperature tolerance of deep-sea Gromia is uncertain.

Gromia are thought to acquire nutrients from the organic matter in sediments on the sea floor, as they are often found in areas with abundant phytodetritus [20]. Their apertures face down on sediment surfaces and they use their pseudopodia to feed [21].

Gromiids found in the deep sea near Oman and Pakistan are often found with Foraminifera, filamentous prokaryotes and bacteria living on their cell surface [22]. Gromiids provide substrates and serve as a surface for attachment to their epibionts.

Description of Organism

Gromia members are quite large, ranging from 0.4 mm to 30 mm [23] [24]. Their proteinaceous tests vary in shapes, from spherical (e.g. G. oviformis), “sausage shaped,” “grape-shaped” or pear-shaped (e.g. G. pyriforminis) [25]. Test shape is often used for classifying Gromia species, and their morphology tends to align with the molecular data used to differentiate species. The interior of the test is layered with membranes with a honeycomb pattern. These honeycomb membranes are a unique feature of Gromia.

An oral complex containing an aperture, which is an opening in the test, allows the filose pseudopodia to extend out [26]. The pseudopodia are non-granular, and can form connections to make net-like structures [27]. Gromia use their pseudopodia to crawl along the surface of sediments [28]. Stercomata, or waste pellets, as well as mineral grains accumulate inside the cell — another characteristic feature of Gromia [29]).

Life cycle

Gromia have been observed to undergo both asexual and sexual reproduction. In sexual reproduction observed in G. oviformis, the shells of adult organisms fuse [30]. Gametogenesis and fertilization follow, after which the zygotes mature into amoebulae and exit the parental shells.

Practical Importance

Gromiids are hypothesized to be important for carbon cycling, as they are often found in carbon-rich sediments and feed on detritus [31]. In addition, gromiids have been shown to store high levels of intracellular nitrate, suggesting a role for gromiids in denitrification [32].

Gromiids have also enriched our understanding of evolutionary history. The ability of the giant, deep sea species G. sphaerica to produce tracks on the sea floor has been used to propose a re-evaluation of the use of fossils with similar traces as evidence for dating the origins of animals with bilateral symmetry [33]

Scientific Classification

Chromista, Harosa, Rhizaria, Cercozoa, Endomyxa, Grommidea, Gromia, Gromiidae, Gromia [34]

List of Species

[35] [36]

Gromia oviformis Dujardin, 1835

Gromia appendiculariae Brooks & Kellner, 1908

Gromia dubia Gruber, 1884

Gromia dujardinii Schultze, 1854

Gromia fluvialis Dujardin, 1837

Gromia granulata Schulze, 1875

Gromia solenopus Zarnik, 1907

Gromia granulata Schulze, 1875

Gromia hyalina Schlumberger, 1845

Gromia paludosa Cienkowski, 1876

Gromia schulzei Norman, 1892

Gromia terricola Leidy, 1874

Gromia sphaerica Gooday, Bowser, Bett & Smith 2000

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References
  1. Rothe, N., Gooday, A. J., Cedhagen, T., & Alan Hughes, J. (2011). Biodiversity and distribution of the genus Gromia (Protista, Rhizaria) in the deep Weddell Sea (Southern Ocean). Polar Biology, 34(1), 69–81. https://doi.org/10.1007/s00300-010-0859-z
  2. Rothe, N., Gooday, A. J., Cedhagen, T., & Alan Hughes, J. (2011). Biodiversity and distribution of the genus Gromia (Protista, Rhizaria) in the deep Weddell Sea (Southern Ocean). Polar Biology, 34(1), 69–81. https://doi.org/10.1007/s00300-010-0859-z
  3. Matz, M. V., Frank, T. M., Marshall, N. J., Widder, E. A., & Johnsen, S. (2008). Giant Deep-Sea Protist Produces Bilaterian-like Traces. Current Biology, 18(23), 1849–1854. https://doi.org/10.1016/j.cub.2008.10.028
  4. Rothe, N., Gooday, A. J., Cedhagen, T., & Alan Hughes, J. (2011). Biodiversity and distribution of the genus Gromia (Protista, Rhizaria) in the deep Weddell Sea (Southern Ocean). Polar Biology, 34(1), 69–81. https://doi.org/10.1007/s00300-010-0859-z
  5. Høgslund, S., Cedhagen, T., Bowser, S. S., & Risgaard-Petersen, N. (2017). Sinks and Sources of Intracellular Nitrate in Gromiids. Frontiers in Microbiology, 8(APR), 617. https://doi.org/10.3389/fmicb.2017.00617
  6. Hayward, B. W., Le Coze, F., & Gross, O. (2018). World Foraminifera Database. Gromia Dujardin. Retrieved February 26, 2020, from http://www.marinespecies.org/foraminifera/aphia.php?p=taxdetails&id=111984
  7. Cifelli, R. (1990). A History of the Classification of Foraminifera (1826-1933) Part 1 Foraminiferal Classification From D’Orbigny To Galloway. Cushman Foundation for Foraminiferal Research, 1(27), 106.
  8. Hedley, R. H., & Bertaud, W. S. (1962). Electron‐Microscopic Observations of Gromia oviformis (Sarcodina). The Journal of Protozoology, 9(1), 79–87.
  9. Rothe, N., Gooday, A. J., Cedhagen, T., & Alan Hughes, J. (2011). Biodiversity and distribution of the genus Gromia (Protista, Rhizaria) in the deep Weddell Sea (Southern Ocean). Polar Biology, 34(1), 69–81. https://doi.org/10.1007/s00300-010-0859-z
  10. Longet, D., Burki, F., Flakowski, J., Berney, C., Polet, S., Fahrni, J., & Pawlowski, J. (2004). Multigene evidence for close evolutionary relations between Gromia and foraminifera. Acta Protozoologica, 43(4), 303–311.
  11. Cavalier-Smith, T., & Chao, E. E.-Y. (2003). Phylogeny and Classification of Phylum Cercozoa (Protozoa). Protist, 154(3–4), 341–358. https://doi.org/10.1078/143446103322454112
  12. Longet, D., Burki, F., Flakowski, J., Berney, C., Polet, S., Fahrni, J., & Pawlowski, J. (2004). Multigene evidence for close evolutionary relations between Gromia and foraminifera. Acta Protozoologica, 43(4), 303–311.
  13. Aranda da Silva, A., Pawlowski, J., & Gooday, A. J. (2006). High diversity of deep-sea Gromia from the Arabian Sea revealed by small subunit rDNA sequence analysis. Marine Biology, 148(4), 769–777. https://doi.org/10.1007/s00227-005-0071-9
  14. Gooday, A. J., Bowser, S. S., Bett, B. J., & Smith, C. R. (2000). A large testate protist, Gromia sphaerica sp. nov. (Order Filosea), from the bathyal Arabian Sea. Deep-Sea Research Part II: Topical Studies in Oceanography, 47(1–2), 55–73. https://doi.org/10.1016/S0967-0645(99)00100-9
  15. Rothe, N., Gooday, A. J., Cedhagen, T., & Alan Hughes, J. (2011). Biodiversity and distribution of the genus Gromia (Protista, Rhizaria) in the deep Weddell Sea (Southern Ocean). Polar Biology, 34(1), 69–81. https://doi.org/10.1007/s00300-010-0859-z
  16. Aranda da Silva, A., Pawlowski, J., & Gooday, A. J. (2006). High diversity of deep-sea Gromia from the Arabian Sea revealed by small subunit rDNA sequence analysis. Marine Biology, 148(4), 769–777. https://doi.org/10.1007/s00227-005-0071-9
  17. Aranda da Silva, A., & Gooday, A. J. (2009). Large organic-walled Protista (Gromia) in the Arabian Sea: Density, diversity, distribution and ecology. Deep Sea Research Part II: Topical Studies in Oceanography, 56(6–7), 422–433. https://doi.org/10.1016/j.dsr2.2008.12.027
  18. Rothe, N., Gooday, A. J., Cedhagen, T., & Alan Hughes, J. (2011). Biodiversity and distribution of the genus Gromia (Protista, Rhizaria) in the deep Weddell Sea (Southern Ocean). Polar Biology, 34(1), 69–81. https://doi.org/10.1007/s00300-010-0859-z
  19. Rothe, N., Gooday, A. J., Cedhagen, T., & Alan Hughes, J. (2011). Biodiversity and distribution of the genus Gromia (Protista, Rhizaria) in the deep Weddell Sea (Southern Ocean). Polar Biology, 34(1), 69–81. https://doi.org/10.1007/s00300-010-0859-z
  20. Rothe, N., Gooday, A. J., Cedhagen, T., & Alan Hughes, J. (2011). Biodiversity and distribution of the genus Gromia (Protista, Rhizaria) in the deep Weddell Sea (Southern Ocean). Polar Biology, 34(1), 69–81. https://doi.org/10.1007/s00300-010-0859-z
  21. Da Silva, A.A. (2005) Benthic protozoan community attributes in relation to environmental gradients in the Arabian Sea. University of Southampton, Faculty of Engineering Science and Mathematics, School of Ocean and Earth Sciences, Doctoral Thesis, 197pp. a. Retrieved from http://eprints.soton.ac.uk/18664/
  22. Aranda da Silva, A., & Gooday, A. J. (2009). Large organic-walled Protista (Gromia) in the Arabian Sea: Density, diversity, distribution and ecology. Deep Sea Research Part II: Topical Studies in Oceanography, 56(6–7), 422–433. https://doi.org/10.1016/j.dsr2.2008.12.027
  23. Matz, M. V., Frank, T. M., Marshall, N. J., Widder, E. A., & Johnsen, S. (2008). Giant Deep-Sea Protist Produces Bilaterian-like Traces. Current Biology, 18(23), 1849–1854. https://doi.org/10.1016/j.cub.2008.10.028
  24. Rothe, N., Gooday, A. J., Cedhagen, T., & Alan Hughes, J. (2011). Biodiversity and distribution of the genus Gromia (Protista, Rhizaria) in the deep Weddell Sea (Southern Ocean). Polar Biology, 34(1), 69–81. https://doi.org/10.1007/s00300-010-0859-z
  25. Aranda da Silva, A., & Gooday, A. J. (2009). Large organic-walled Protista (Gromia) in the Arabian Sea: Density, diversity, distribution and ecology. Deep Sea Research Part II: Topical Studies in Oceanography, 56(6–7), 422–433. https://doi.org/10.1016/j.dsr2.2008.12.027
  26. Felts, W. J. L., & Harrison, R. J. (1968). International Review of General and Experimental Zoology. Journal of Anatomy, 103(Pt 2), 377.
  27. Longet, D., Burki, F., Flakowski, J., Berney, C., Polet, S., Fahrni, J., & Pawlowski, J. (2004). Multigene evidence for close evolutionary relations between Gromia and foraminifera. Acta Protozoologica, 43(4), 303–311.
  28. Matz, M. V., Frank, T. M., Marshall, N. J., Widder, E. A., & Johnsen, S. (2008). Giant Deep-Sea Protist Produces Bilaterian-like Traces. Current Biology, 18(23), 1849–1854. https://doi.org/10.1016/j.cub.2008.10.028
  29. Rothe, N., Gooday, A. J., Cedhagen, T., & Alan Hughes, J. (2011). Biodiversity and distribution of the genus Gromia (Protista, Rhizaria) in the deep Weddell Sea (Southern Ocean). Polar Biology, 34(1), 69–81. https://doi.org/10.1007/s00300-010-0859-z
  30. ARNOLD, Z. M. (1966). Observations on the Sexual Generation of Gromia oviformis Dujardin. The Journal of Protozoology, 13(1), 23–27. https://doi.org/10.1111/j.1550-7408.1966.tb01863.x
  31. Da Silva, A.A. (2005) Benthic protozoan community attributes in relation to environmental gradients in the Arabian Sea. University of Southampton, Faculty of Engineering Science and Mathematics, School of Ocean and Earth Sciences, Doctoral Thesis, 197pp. a. Retrieved from http://eprints.soton.ac.uk/18664/
  32. Høgslund, S., Cedhagen, T., Bowser, S. S., & Risgaard-Petersen, N. (2017). Sinks and Sources of Intracellular Nitrate in Gromiids. Frontiers in Microbiology, 8(APR), 617. https://doi.org/10.3389/fmicb.2017.00617
  33. Matz, M. V., Frank, T. M., Marshall, N. J., Widder, E. A., & Johnsen, S. (2008). Giant Deep-Sea Protist Produces Bilaterian-like Traces. Current Biology, 18(23), 1849–1854. https://doi.org/10.1016/j.cub.2008.10.028
  34. Hayward, B. W., Le Coze, F., & Gross, O. (2018). World Foraminifera Database. Gromia Dujardin. Retrieved February 26, 2020, from http://www.marinespecies.org/foraminifera/aphia.php?p=taxdetails&id=111984
  35. Hayward, B. W., Le Coze, F., & Gross, O. (2018). World Foraminifera Database. Gromia Dujardin. Retrieved February 26, 2020, from http://www.marinespecies.org/foraminifera/aphia.php?p=taxdetails&id=111984
  36. WoRMS - World Register of Marine Species. (2012). Retrieved March 30, 2020, from http://www.marinespecies.org/aphia.php?p=taxlist&tName=Gromia
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