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''Geodia barretti'' is a massive deep-sea
sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through t ...
species found in the
boreal Boreal may refer to: Climatology and geography *Boreal (age), the first climatic phase of the Blytt-Sernander sequence of northern Europe, during the Holocene epoch *Boreal climate, a climate characterized by long winters and short, cool to mild ...
waters of the North Atlantic Ocean, and is fairly common on the coasts of Norway and Sweden. It is a dominant species in boreal sponge grounds. Supported by morphology and molecular data, this species is classified in the family Geodiidae.


Morphology


External morphology

Massive sponge, often irregularly lobate, whitish to light yellow color. The inside is light brown. Surface is smooth. Oscules are grouped in more or less shallow depressions (= preoscules) while pores are spread over the whole body. There is a distinct cortex about 0.5 mm thick, it is made of ball-shaped
spicules Spicules are any of various small needle-like anatomical structures occurring in organisms Spicule may also refer to: *Spicule (sponge), small skeletal elements of sea sponges *Spicule (nematode), reproductive structures found in male nematodes ( ...
called sterrasters.


Spicules


Megascleres.

* Oxeas (1000-5000 µm). * Dichotriaenes with rhabdomes up to 5000 µm long (more rarely orthotriaenes). * Anatriaenes. * Meso/protriaenes (rare).


Microscleres.

* Microxeas (190-900 µm), sometimes slightly centrotylote. * Sterrasters (50-130 µm). * Oxyasters I (30-80 µm), microspined branches, only found in some specimens living > 1000 m. * Oxyasters II (6-33 µm), microspined branches. * Strongylasters (3-11 µm), microspined branches.


Genetics


DNA barcodes

* Cytochrome oxidase subunit 1 gene (mitochondrial). Folmer fragment: 2
haplotype A haplotype ( haploid genotype) is a group of alleles in an organism that are inherited together from a single parent. Many organisms contain genetic material ( DNA) which is inherited from two parents. Normally these organisms have their DNA or ...
s with 1
base pair A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both DNA ...
difference. One haplotype
EU442194
is present throughout the North Atlantic. The second haplotype
KC574389
has so far only been found in
Flemish Cap The Flemish Cap is an area of shallow waters in the north Atlantic Ocean centered roughly at 47° north, 45° west or about 563 km (350 miles) east of St. John's, Newfoundland and Labrador. The shallow water is caused by a wide underwater ...
populations. *
28S rDNA 28S ribosomal RNA is the structural ribosomal RNA (rRNA) for the LSU rRNA, large subunit (LSU) of eukaryotic cytoplasmic ribosomes, and thus one of the basic components of all eukaryotic cells. It has a size of 25S in plants and 28S in mammals, ...
(nuclear). C1-D2 domains. Genbank number
HM592809EU552080
*
18S rDNA 18S ribosomal RNA (abbreviated 18S rRNA) is a part of the ribosomal RNA. The S in 18S represents Svedberg units. 18S rRNA is an SSU rRNA, a component of the eukaryotic ribosomal small subunit ( 40S). 18S rRNA is the structural RNA for the small ...
(nuclear). Genbank numbe
KC481224
ref name="Cárdenas 251–311">


Genomics

One meta-transcriptome was obtained by
pyrosequencing Pyrosequencing is a method of DNA sequencing (determining the order of nucleotides in DNA) based on the "sequencing by synthesis" principle, in which the sequencing is performed by detecting the nucleotide incorporated by a DNA polymerase. Pyrosequ ...
from a specimen collected south of
Bergen Bergen (), historically Bjørgvin, is a city and municipality in Vestland county on the west coast of Norway. , its population is roughly 285,900. Bergen is the second-largest city in Norway. The municipality covers and is on the peninsula of ...
(
Norway Norway, officially the Kingdom of Norway, is a Nordic country in Northern Europe, the mainland territory of which comprises the western and northernmost portion of the Scandinavian Peninsula. The remote Arctic island of Jan Mayen and t ...
) in 2007. Double-stranded
cDNA In genetics, complementary DNA (cDNA) is DNA synthesized from a single-stranded RNA (e.g., messenger RNA (mRNA) or microRNA (miRNA)) template in a reaction catalyzed by the enzyme reverse transcriptase. cDNA is often used to express a speci ...
was sequenced on a GF FLX sequencer (Roche Applied Sciences/
454 Life Sciences 454 Life Sciences was a biotechnology company based in Branford, Connecticut that specialized in high-throughput DNA sequencing. It was acquired by Roche in 2007 and shut down by Roche in 2013 when its technology became noncompetitive, although ...
, Barnford, CT).
rRNA Ribosomal ribonucleic acid (rRNA) is a type of non-coding RNA which is the primary component of ribosomes, essential to all cells. rRNA is a ribozyme which carries out protein synthesis in ribosomes. Ribosomal RNA is transcribed from ribosoma ...
ribo-tags revealed a microsymbiont community, dominated by group SAR202 of ''
Chloroflexota The Chloroflexota are a phylum of bacteria containing isolates with a diversity of phenotypes, including members that are aerobic thermophiles, which use oxygen and grow well in high temperatures; anoxygenic phototrophs, which use light for photo ...
'', candidate phylum "''
Poribacteria Poribacteria are a candidate phylum of bacteria originally discovered in the microbiome of marine sponges (''Porifera''). Poribacteria are Gram-negative primarily aerobic mixotrophs with the ability for oxidative phosphorylation, glycolysis, ...
''", and ''
Acidobacteriota Acidobacteriota is a phylum of Gram-negative bacteria. Its members are physiologically diverse and ubiquitous, especially in soils, but are under-represented in culture. Description Members of this phylum are physiologically diverse, and can be ...
''. The most abundant
mRNAs In molecular biology, messenger ribonucleic acid (mRNA) is a single-stranded molecule of RNA that corresponds to the genetic sequence of a gene, and is read by a ribosome in the process of synthesizing a protein. mRNA is created during the p ...
coded for key metabolic enzymes of
nitrification ''Nitrification'' is the biological oxidation of ammonia to nitrite followed by the oxidation of the nitrite to nitrate occurring through separate organisms or direct ammonia oxidation to nitrate in comammox bacteria. The transformation of amm ...
from ammonia-oxidizing
Archaea Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebac ...
as well as candidate genes involved in related processes.


Systematics


Etymology

This sponge was named in honor of one of its discoverers, the biologist
Lucas Barrett Lucas Barrett (14 November 1837 – 19 December 1862) was an English natural history, naturalist and geologist. He was the director of the Jamaican Geological Survey from 1859 to 1862. He was a young member of the Geological Society of London, Geo ...
(1837-1862).


Vernacular names

This sponge is called "
rutabaga Rutabaga (; North American English) or swede (British English and some Commonwealth English) is a root vegetable, a form of ''Brassica napus'' (which also includes rapeseed). Other names include Swedish turnip, neep (Scots), and turnip (Scott ...
sponge" in Norwegian (kålrabisvamp) and "football sponge" in Swedish (fotbollssvampdjur).


Type material

This sponge was collected by biologists Robert McAndrew and
Lucas Barrett Lucas Barrett (14 November 1837 – 19 December 1862) was an English natural history, naturalist and geologist. He was the director of the Jamaican Geological Survey from 1859 to 1862. He was a young member of the Geological Society of London, Geo ...
in 1855 on the South side of Vikna Island (formerly called Vigten or Vikten Island), North-Trøndelag, Norway, at a depth of 183 meters. It was later described by the British sponge taxonomist
James Scott Bowerbank James Scott Bowerbank (14 July 1797 – 8 March 1877) was a British naturalist and palaeontologist. Biography Bowerbank was born in Bishopsgate, London, and succeeded in conjunction with his brother to his father's distillery, in which he was a ...
in 1858. The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
is a dry specimen stored at the
Natural History Museum A natural history museum or museum of natural history is a scientific institution with natural history collections that include current and historical records of animals, plants, fungi, ecosystems, geology, paleontology, climatology, and more. ...
in London with the museum number BNHM 1877.5.21.1399. There are also slides of the holotype: BNHM 1877.5.21.1400 (one slide of surface and one spicule preparation) and BNHM 1877.5.21.1401 (slide of a section).


Phylogenetic relationships

According to morphology and molecular data, this species belongs to the family Geodiidae and to the genus '' Geodia''. It also seems to belong to a clade of '' Geodia'' species that all have their oscules in depressions (including '' Geodia megastrella'', '' Geodia hentscheli''). This '' Geodia'' clade was given a
PhyloCode The ''International Code of Phylogenetic Nomenclature'', known as the ''PhyloCode'' for short, is a formal set of rules governing phylogenetic nomenclature. Its current version is specifically designed to regulate the naming of clades, leaving the ...
name: ''Depressiogeodia''.


Distribution


Biogeography

''G. barretti'' is present throughout the
boreal Boreal may refer to: Climatology and geography *Boreal (age), the first climatic phase of the Blytt-Sernander sequence of northern Europe, during the Holocene epoch *Boreal climate, a climate characterized by long winters and short, cool to mild ...
waters of the
North Atlantic The Atlantic Ocean is the second-largest of the world's five oceans, with an area of about . It covers approximately 20% of Earth's surface and about 29% of its water surface area. It is known to separate the "Old World" of Africa, Europe and ...
, where water temperature is usually 3-9 degrees Celsius. This species is present on the Swedish west coast and all along the Norwegian coast and Svalbard. One single specimen was formally recorded from the
Mediterranean Sea The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the ea ...
, in the ‘Canyon des Moines’ (south
Corsica Corsica ( , Upper , Southern ; it, Corsica; ; french: Corse ; lij, Còrsega; sc, Còssiga) is an island in the Mediterranean Sea and one of the 18 regions of France. It is the fourth-largest island in the Mediterranean and lies southeast of ...
), at 167 m (water temperature was 13 °C). Deeper specimens (1100-1700 meter depths) were also recorded off the Atlantic coast of Morocco (Ibero-Moroccan Gulf). Several deep specimens were collected on the mid-Atlantic ridge at 1650 m depth.


Bathymetry

It is found from 30 to 2000 meter depths. Most North-East Atlantic records are from the
continental margin A continental margin is the outer edge of continental crust abutting oceanic crust under coastal waters. It is one of the three major zones of the ocean floor, the other two being deep-ocean basins and mid-ocean ridges. The continental margin ...
s, between 200 and 500 m. In the North-West Atlantic (Grand Banks, Flemish Cap, Nova Scotia, and Davis Strait), it has been recorded between 410 and 1852 meter depths. Shallow specimens are common along the Norwegian coast, especially in
fjord In physical geography, a fjord or fiord () is a long, narrow inlet with steep sides or cliffs, created by a glacier. Fjords exist on the coasts of Alaska, Antarctica, British Columbia, Chile, Denmark, Germany, Greenland, the Faroe Islands, Ice ...
s.


Biology


Reproduction

''G. barretti'' is an oviparous species with separated males and females. The reproductive cycle is annual with one or two events of
gamete A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce t ...
release per year, the major reproductive season being from late spring to early autumn. The larvae has never been observed. Asexual reproduction has never been formally described. The reproduction molecular toolkit of boreo-arctic '' Geodia'' species has been investigated, including in ''Geodia barretti'', using
proteomic Proteomics is the large-scale study of proteins. Proteins are vital parts of living organisms, with many functions such as the formation of structural fibers of muscle tissue, enzymatic digestion of food, or synthesis and replication of DNA. In ...
and trancriptomic data. Genes involved in the formation of the
gametes A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce t ...
appeared close to the ones found in the rest of the animals.


Predators

The sponge-feeding deep-sea
chiton Chitons () are marine molluscs of varying size in the class Polyplacophora (), formerly known as Amphineura. About 940 extant and 430 fossil species are recognized. They are also sometimes known as gumboots or sea cradles or coat-of-mail s ...
''
Hanleya ''Hanleya'' is a genus of polyplacophoran molluscs known from Oligocene and Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell ...
nagelfar'' is often found on ''G. barretti'' in the North-East Atlantic. The parasitic
foraminifera Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular Ectoplasm (cell biology), ectoplasm for catching food and ot ...
n '' Hyrrokkin sarcophaga'' is more rarely found living on ''G. barretti''.


Associated macrofauna

''G. barretti'' has less associated macrofauna than any of the other boreo-arctic ''Geodia'' species: only ten different species of epibionts were recorded vs. 62 for '' Geodia macandrewii''.


Microbiology

''G. barretti'' is a high microbial abundance (HMA) sponge, such that the mesohyl is essentially packed with microbes, and just a few sponge cells. It was calculated that the sponge itself (canals, choanocyte chambers and cells) represents only 5% of its total volume, the rest being microbes. Estimates of the number of
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
in ''G. barretti'' are between 2.9x1011 and 3.1x1011 microbes/cm3 sponge.


Isolation and study of specific bacteria

''
Streptomyces ''Streptomyces'' is the largest genus of Actinomycetota and the type genus of the family Streptomycetaceae. Over 500 species of ''Streptomyces'' bacteria have been described. As with the other Actinomycetota, streptomycetes are gram-positive, ...
'' sp. GBA 94-10, an
Actinomycetota The ''Actinomycetota'' (or ''Actinobacteria'') are a phylum of all gram-positive bacteria. They can be terrestrial or aquatic. They are of great economic importance to humans because agriculture and forests depend on their contributions to soi ...
, was isolated from a Norwegian ''G. barretti''. It was shown to be a close relative of the terrestrial ''Streptomyces albus'' J1074. The genome was fully sequenced (Genbank accession number
CM002271
plasmid pGBA
CM002272
. ''
Streptomyces ''Streptomyces'' is the largest genus of Actinomycetota and the type genus of the family Streptomycetaceae. Over 500 species of ''Streptomyces'' bacteria have been described. As with the other Actinomycetota, streptomycetes are gram-positive, ...
'' ''poriferorum'' (type strain P01-B04T), an
Actinomycetota The ''Actinomycetota'' (or ''Actinobacteria'') are a phylum of all gram-positive bacteria. They can be terrestrial or aquatic. They are of great economic importance to humans because agriculture and forests depend on their contributions to soi ...
, was isolated and described in two Norwegian sponges ('' Antho dichotoma'' and ''G. barretti''). This bacteria might provide their host with chemical defenses and might be involved in nitrogen metabolism. The whole genome was sequenced (Genbank accession numbe
JAELVH010000000
; the 16S rRNA gene sequence can also be found on Genbank
MW583039
. '' Actinoalloteichus fjordicus'', an
Actinomycetota The ''Actinomycetota'' (or ''Actinobacteria'') are a phylum of all gram-positive bacteria. They can be terrestrial or aquatic. They are of great economic importance to humans because agriculture and forests depend on their contributions to soi ...
, was isolated and described in two Norwegian sponges ('' Antho dichotoma'' and ''G. barretti''). This Actinomycetota genus has been found in terrestrial and marine habitats. This bacterium's genome was fully sequenced (Genbank accession number
CP016077
plasmi
CP016078
.


Microbiomes


Chemistry

The major compound of ''G. barretti'', called
barettin Barettin is a brominated alkaloid made of a dehydrogenated brominated derivative of tryptophan linked by two peptide bonds to an arginine residue, forming a 2,5-diketopiperazine nucleus. It is a cyclic dipeptide. Barettin is the major compound ...
, was isolated in 1986 at
Uppsala University Uppsala University ( sv, Uppsala universitet) is a public university, public research university in Uppsala, Sweden. Founded in 1477, it is the List of universities in Sweden, oldest university in Sweden and the Nordic countries still in opera ...
,
Sweden Sweden, formally the Kingdom of Sweden,The United Nations Group of Experts on Geographical Names states that the country's formal name is the Kingdom of SwedenUNGEGN World Geographical Names, Sweden./ref> is a Nordic country located on ...
; barettin has
antifouling Biofouling or biological fouling is the accumulation of microorganisms, plants, algae, or small animals where it is not wanted on surfaces such as ship and submarine hulls, devices such as water inlets, pipework, grates, ponds, and rivers that ...
activity. Since then, two more barettins were discovered, as well as other small molecules (see Table below). Peptides called barrettides were found and published in 2015. Using
metabolomics Metabolomics is the scientific study of chemical processes involving metabolites, the small molecule substrates, intermediates, and products of cell metabolism. Specifically, metabolomics is the "systematic study of the unique chemical fingerprin ...
on specimens of ''G. barretti'' from Sweden, most of these compounds could be found, in addition to
nucleotides Nucleotides are organic molecules consisting of a nucleoside and a phosphate. They serve as monomeric units of the nucleic acid polymers – deoxyribonucleic acid (DNA) and ribonucleic acid (RNA), both of which are essential biomolecules w ...
,
nucleosides Nucleosides are glycosylamines that can be thought of as nucleotides without a phosphate group. A nucleoside consists simply of a nucleobase (also termed a nitrogenous base) and a five-carbon sugar (ribose or 2'-deoxyribose) whereas a nucleotide ...
and nucleobases, as well as some
fatty acids In chemistry, particularly in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated or unsaturated. Most naturally occurring fatty acids have an unbranched chain of an even number of carbon atoms, f ...
.


Conservation


References

{{Taxonbar, from=Q946464 Tetractinellida Taxa named by James Scott Bowerbank Animals described in 1858 Fauna of the Atlantic Ocean