Gromia

''Gromia'' is a genus of protists, closely related to foraminifera, which inhabit marine and freshwater environments. It is the only genus of the family Gromiidae. ''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'' were first discovered in shallow waters, with members of the best-characterized species ''Gromia oviformus'' often found inhabiting rock surfaces, sediments, or seaweed holdfasts. 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.

A recent study of the deep sea species '' Gromia sphaerica'' revealed that it produces traces on the seafloor which resemble fossil traces formerly attributed to early Bilateria (animals with bilateral symmetry); this now calls into question whether such fossil traces are reliable as documentation of early multicellular animal diversification in the Precambrian era.

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

History of study

''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. Initially, ''Gromia'' were regarded as members of Foraminifera or Filosea, as noted in a review by Cifelli (1990).

''Gromia'' became better characterized throughout the 1960s, when electron microscopy revealed more details on their morphology, including their honeycomb

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. Follow-up studies on this group placed ''Gromia'' within the Gromiidea class, again based on SSU rRNA genes
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. 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.

''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''. 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.

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'' is ''G. oviformis''. It inhabits intertidal zones and other regions of shallow waters; it is often found attached to rocks, kelp, weeds, Cladophora algae, or within sediments. ''G. oviformis'' has been shown to tolerate a temperature range of 0–30 °C.

Deep-sea gromiids have been found in the Arabian sea, off the coast of Antarctica and in the water of the Northwest Atlantic Ocean. They were often collected from the 1000–3,100 m range. Oxygen levels in gromiid habitats tend to exceed 0.2 mL/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. Their apertures face down on sediment surfaces and they use their pseudopodia to feed.

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

Description

''Gromia'' members are quite large, ranging from 0.4 mm to 30 mm. Their proteinaceous tests vary in shapes, from spherical (e.g. ''G. oviformis''), “sausage shaped”, “grape-shaped”, or pear-shaped (e.g. ''G. pyriforminis''). 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 (an opening in the test) allows the filose pseudopodia to extend out. The pseudopodia are non-granular, and can form connections to make net-like structures. Gromia use their pseudopodia to crawl along the surface of sediments. Waste pellets (“stercomata”) and mineral grains accumulate inside the cell — another characteristic feature of ''Gromia''.

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. 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. In addition, gromiids have been shown to store high levels of intracellular nitrate, suggesting a role for gromiids in denitrification.

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.

Scientific classification

Chromista, Harosa, Rhizaria, Cercozoa, Endomyxa, Grommidea, Gromia, Gromiidae, ''Gromia''.

List of species

Sources:

*''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 sphaerica'' Gooday, Bowser, Bett & Smith 2000

References

External links

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{{Taxonbar, from1=Q288010, from2=Q25368163

Cercozoa genera
Amoeboids