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''Suberites'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of
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 th ...
s in the
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
Suberitidae. Sponges, known scientifically as ''Porifera'', are the oldest
metazoan Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
s and are used to elucidate the basics of
multicellular evolution A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially uni- ...
. These living fossils are ideal for studying the principal features of metazoans, such as extracellular matrix interactions, signal-receptor systems, nervous or sensory systems, and primitive immune systems. Thus, sponges are useful tools with which to study early animal evolution. They appeared approximately 580 million years ago, in the Ediacaran.


Evolutionary significance

As members of the oldest phylum of metazoans, ''Suberites'' serve as model organisms to elucidate features of the earliest animals. ''Suberites'' and their relatives are used to determine the structure of the first metazoans and have been studied to determine how
totipotency Pluripotency: These are the cells that can generate into any of the three Germ layers which imply Endodermal, Mesodermal, and Ectodermal cells except tissues like the placenta. According to Latin terms, Pluripotentia means the ability for many thin ...
has replaced by
pluripotency Pluripotency: These are the cells that can generate into any of the three Germ layers which imply Endodermal, Mesodermal, and Ectodermal cells except tissues like the placenta. According to Latin terms, Pluripotentia means the ability for many thin ...
in most higher animals. Among other things, ''Suberites'' show that tyrosine-phosphorylation machinery evolved in animals independently from other eukaryotes. ''Suberites'' are also used as models to elucidate the evolution of
transmembrane receptor Cell surface receptors (membrane receptors, transmembrane receptors) are receptors that are embedded in the plasma membrane of cells. They act in cell signaling by receiving (binding to) extracellular molecules. They are specialized integral me ...
s and cell-junction proteins. A combination of stem cell and apoptosis factors studies is used as a model for studies of development in higher animals.


Ecology

''Suberites'' are a global genus. One species, ''Suberites zeteki'', is found in Hawaii. ''S. zeteki'' associates with many fungi.G. Zheng, L. Binglin, Z. Chengchao, W. Guangyi, Molecular Detection of Fungal Communities in the Hawaiian Marine Sponges Suberites zeteki and Mycale armata. Applied & Environmental Microbiology 74, 6091 (2008). Another, ''S. japonicas'', is native to the waters around Japan.C. Tanaka, J. Tanaka, R. F. Bolland, G. Marriott, T. Higa, Seragamides A–F, new actin-targeting depsipeptides from the sponge Suberites japonicus Thiele. Tetrahedron 62, 3536 (2006). ''S. aurantiacus'' is found in the Caribbean sea.L. P. Ponomarenko, O. A. Vanteeva, S. A. Rod'kina, V. B. Krasokhin, S. S. Afiyatullov, Metabolites of the marine sponge Suberites cf. aurantiacus. Chemistry of Natural Compounds 46, 335 (2010). ''S.carnosus'' lives in the Indian Ocean and in the Mediterranean Sea and can also be found in Irish waters.B. Flemer et al., Diversity and antimicrobial activities of microbes from two Irish marine sponges, Suberites carnosus and Leucosolenia sp. Journal of Applied Microbiology 112, 289 (2012). ''S. diversicolor'' can be found in Indonesia. Due to ''Suberites’'' ability to efficiently filter water, many microbes, especially fungal species, are filtered through. If these microbes escape digestion, they can deposit on the sponge and reside there indefinitely. Symbiotic bacteria produce toxins, such as
okadaic acid Okadaic acid, C44H68O13, is a toxin produced by several species of dinoflagellates, and is known to accumulate in both marine sponges and shellfish. One of the primary causes of diarrhetic shellfish poisoning, okadaic acid is a potent inhibitor of ...
, which defend them from colonization by parasitic annelids.H. C. Schröder et al., Okadaic Acid, an Apoptogenic Toxin for Symbiotic/Parasitic Annelids in the Demosponge Suberites domuncula. Applied & Environmental Microbiology 72, 4907 (2006).W. E. G. Müller et al., Oxygen-Controlled Bacterial Growth in the Sponge Suberites domuncula: toward a Molecular Understanding of the Symbiotic Relationships between Sponge and Bacteria. Applied & Environmental Microbiology 70, 2332 (2004). Expression of various enzymes by ''Suberites'' influences the growth of their symbiotic bacteria. ''Suberites'' often live on the shells on the mollusk ''
Hexaplex trunculus ''Hexaplex trunculus'' (previously known as ''Murex trunculus'', ''Phyllonotus trunculus'', or the banded dye-murex) is a medium-sized sea snail, a marine gastropod mollusk in the family Muricidae, the murex shells or rock snails. It is included ...
''. ''Suberites'' have mechanisms of defense against predation, such as the toxic chemicals found below.


Physiology

''Suberites'' display
neuronal communication A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa. No ...
s, but
neuronal network A neural circuit is a population of neurons interconnected by synapses to carry out a specific function when activated. Neural circuits interconnect to one another to form large scale brain networks. Biological neural networks have inspired the ...
s are mysteriously missing. However, they do have many of the same sensory receptors and signals found in higher animals.X. Wang, X. Fan, H. Schröder, W. Müller, Flashing light in sponges through their siliceous fiber network: A new strategy of 'neuronal transmission' in animals. Chinese Science Bulletin 57, 3300 (2012). Researchers in China and Germany have found that sponge spicules contribute to their neural communication.W. E. G. Müller et al., Luciferase a light source for the silica-based optical waveguides (spicules) in the demosponge Suberites domuncula. Cellular and Molecular Life Sciences 66, 537 (2009). In effect, the silicaceous structures act as fiber optic cables to convey light signals generated from the protein luciferase. The sponges generate light from luciferin, after it is acted upon by luciferase.W. E. G. Müller et al., A cryptochrome-based photosensory system in the siliceous sponge Suberites domuncula (Demospongiae). FEBS Journal 277, 1182 (2010). ''Suberites'' have also been shown to produce light in response to tactile stimulation. ''Suberites'' consist mostly of cells, in contrast with other ''Porifera'' (such as the class
Hexactinellida Hexactinellid sponges are sponges with a skeleton made of four- and/or six-pointed siliceous spicules, often referred to as glass sponges. They are usually classified along with other sponges in the phylum Porifera, but some researchers con ...
) which are
syncytial A syncytium (; plural syncytia; from Greek: σύν ''syn'' "together" and κύτος ''kytos'' "box, i.e. cell") or symplasm is a multinucleate cell which can result from multiple cell fusions of uninuclear cells (i.e., cells with a single nucleus) ...
. As a result, ''Suberites'' have slower reaction times in their neural communication. ''Suberites'' utilized many Ras-like
GTPase GTPases are a large family of hydrolase enzymes that bind to the nucleotide guanosine triphosphate (GTP) and hydrolyze it to guanosine diphosphate (GDP). The GTP binding and hydrolysis takes place in the highly conserved P-loop "G domain", a pro ...
s which are used for signaling and affect development. According to comparative studies, ''Suberites'' have some of the most simple indicator proteins, such as collagen, of known animals. Like all sponges, ''Suberites'' are
filter-feeder Filter feeders are a sub-group of suspension feeding animals that feed by straining suspended matter and food particles from water, typically by passing the water over a specialized filtering structure. Some animals that use this method of feedin ...
s. They are extremely efficient and can process thousands of liters of water per day. ''S. domuncula'' has been used for study of graft rejection. Researchers have discovered that apoptotic factors are induced in the tissue that is rejected.


Development

''Suberites'' consist of many
telomerase Telomerase, also called terminal transferase, is a ribonucleoprotein that adds a species-dependent telomere repeat sequence to the 3' end of telomeres. A telomere is a region of repetitive sequences at each end of the chromosomes of most euka ...
-positive cells, which means the cells are essentially immortal, barring
cell death Cell death is the event of a biological cell ceasing to carry out its functions. This may be the result of the natural process of old cells dying and being replaced by new ones, as in programmed cell death, or may result from factors such as d ...
signal. In most cases, the signal is a lack of connection either to the
extracellular matrix In biology, the extracellular matrix (ECM), also called intercellular matrix, is a three-dimensional network consisting of extracellular macromolecules and minerals, such as collagen, enzymes, glycoproteins and hydroxyapatite that provide s ...
or other cells. Their apoptotic cells are similar to homologous to mammalian. However, maintenance of long-lived cells involves proteins such as SDLAGL that are highly similar to yeast and human homologs. Certain inorganic materials, such as iron and selenium, influence the growth of ''Suberites'', including the primmorph growth and spicule formation. ''Suberites'' undergo cell differentiation through a variety of mechanisms based on cell-cell communication.


Morphology

''Suberites'' are key examples of the importance of the extracellular matrix in animals. In sponges, it is mediated by proteoglycans.
Spicule 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 ( ...
formation is also important for ''Suberites''. Spicules are structural support of sponges, similar to skeletons in higher animals. They are normally hollow structures that are formed by lamellar growth.H. C. Schröder et al., Biosilica formation in spicules of the sponge Suberites domuncula: Synchronous expression of a gene cluster. Genomics 85, 666 (2005).H. C. Schröder et al., Apposition of silica lamellae during growth of spicules in the demosponge Suberites domuncula: Biological/biochemical studies and chemical/biomimetical confirmation. Journal of Structural Biology 159, 325 (2007). Whereas higher animal skeletons are largely calcium-based, sponge spicules consist mostly of
silica Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , most commonly found in nature as quartz and in various living organisms. In many parts of the world, silica is the major constituent of sand. Silica is ...
, a silicon dioxide polymer.W. Xiaohong et al., Evagination of Cells Controls Bio-Silica Formation and Maturation during Spicule Formation in Sponges. PLoS ONE 6, 1 (2011). These inorganic structures provide support for the animals.X. Wang et al., Silicateins, silicatein interactors and cellular interplay in sponge skeletogenesis: formation of glass fiber-like spicules. FEBS Journal 279, 1721 (2012). The spicules are biologically-formed silica structures, also known as biosilica. Silica deposition begins intracellularly and is carried out by the enzyme silicatein. Silicateins are modulated by a group of proteins called silintaphins. The process occurs in specialized cells known as
sclerocyte Sclerocytes are specialised cells that secrete the mineralized structures in the body wall of some invertebrates. In sponges they secrete calcareous or siliceous spicules which are found in the mesohyl The mesohyl, formerly known as mesenchyme or ...
s. Biosilica formation in ''Suberites'' differs from other species that utilize biosilica in this regard. Most other species, such as certain plants and
diatoms A diatom (New Latin, Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group com ...
, simply deposit a supersaturated biosilica solution. The network of silica found in sponges mediates much of the sponges’ neural communications.


Immunity and defense

''Suberites'' show the
cytokine Cytokines are a broad and loose category of small proteins (~5–25 kDa) important in cell signaling. Cytokines are peptides and cannot cross the lipid bilayer of cells to enter the cytoplasm. Cytokines have been shown to be involved in autocrin ...
-like molecule allograft inflammatory factor one (AIF-1), which is similar to vertebrate AIF-1.H. C. Schröder et al., Functional Molecular Biodiversity: Assessing the Immune Status of Two Sponge Populations ( Suberites domuncula) on the Molecular Level. Marine Ecology 25, 93 (2004). Immune response relies on phosphorylation cascades involving the p38 kinase. ''S. domuncula'' was the first demonstrated immune response of invertebrate species (1). These sponges also have similar graft-response inflammation to vertebrates. Their immune systems are much simpler than vertebrates; they consist of only innate immunity. Because they filter thousands of liters of water per day, and their environment contains a high concentration of bacteria and viruses, ''Suberites'' have developed a highly potent system of immunity. Despite the efficiency of their immune systems, ''Suberites'' can be susceptible to infection which often stimulates cell death through apoptotic pathways. ''Suberites'', namely ''S. domuncula'', defend themselves from macroscopic threats with a neurotoxin known as suberitine.L. Cariello, L. Zanetti, Suberitine, the toxic protein from the marine sponge suberites domuncula. Comparative Biochemistry and Physiology 64C, 15 (1979). It was the first known protein discovered in a sponge. The neurotoxic properties of suberitine arise from its ability to block action potentials.L. Cariello, E. Tosti, L. Zanetti, The hemolytic activity of suberitine. Comparative Biochemistry and Physiology 73C, 91 (1981). It additionally has hemolytic properties, which do not originate from phospholipase A activity. It has some antibacterial activity; however, the extent of the activity due solely to suberitine is not currently defined. The sponge itself neutralizes the toxin through a pathway that is not fully understood, but involves
retinal Retinal (also known as retinaldehyde) is a polyene chromophore. Retinal, bound to proteins called opsins, is the chemical basis of visual phototransduction, the light-detection stage of visual perception (vision). Some microorganisms use reti ...
, a β-carotene metabolite. ''S. japonicas'' also produces several cytotoxic compounds, seragamides A-F. The seragamides act by interfering with cytoskeleton activity, specifically the
actin Actin is a family of globular multi-functional proteins that form microfilaments in the cytoskeleton, and the thin filaments in muscle fibrils. It is found in essentially all eukaryotic cells, where it may be present at a concentration of ov ...
microfilaments. The activity of the seragamides is a possible route for
anti-cancer drugs Chemotherapy (often abbreviated to chemo and sometimes CTX or CTx) is a type of cancer treatment that uses one or more anti-cancer drugs (chemotherapeutic agents or alkylating agents) as part of a standardized chemotherapy regimen. Chemotherap ...
, similar to existing drugs which target microtubules. ''Suberites'' also produce cytotoxic compounds known as nakijinamines, which resemble other toxins found in ''Suberites'', but the role of the nakijinamines has not yet been found.Y. Takahashi et al., Heteroaromatic alkaloids, nakijinamines, from a sponge Suberites sp. Tetrahedron 68, 8545 (2012). Many of the bioactive compounds found on ''Suberites'' are microbial in nature.


Species

The following species are recognised in the genus ''Suberites'': * '' Suberites affinis'' Brøndsted, 1923 * '' Suberites anastomosus'' Brøndsted, 1924 * '' Suberites aurantiacus'' (Duchassaing & Michelotti, 1864) * '' Suberites australiensis'' Bergquist, 1968 * '' Suberites axiatus'' Ridley & Dendy, 1886 * '' Suberites axinelloides'' Brøndsted, 1924 * '' Suberites baffini'' Brøndsted, 1933 * ''
Suberites bengalensis ''Suberites'' is a genus of sea sponges in the family Suberitidae. Sponges, known scientifically as ''Porifera'', are the oldest metazoans and are used to elucidate the basics of multicellular evolution. These living fossils are ideal for studyi ...
'' Lévi, 1964 * '' Suberites caminatus'' Ridley & Dendy, 1886 * ''
Suberites carnosus ''Suberites'' is a genus of sea sponges in the family Suberitidae. Sponges, known scientifically as ''Porifera'', are the oldest metazoans and are used to elucidate the basics of multicellular evolution. These living fossils are ideal for studyi ...
'' (Johnston, 1842) * '' Suberites cebriones'' Morozov, Sabirov & Zimina, 2019 * '' Suberites clavatus'' Keller, 1891 * '' Suberites concinnus'' Lambe, 1895 * '' Suberites cranium'' Hajdu et al, 2013 * '' Suberites crelloides'' Marenzeller, 1886 * '' Suberites crispolobatus'' Lambe, 1895 * ''
Suberites cupuloides ''Suberites'' is a genus of sea sponges in the family Suberitidae. Sponges, known scientifically as ''Porifera'', are the oldest metazoans and are used to elucidate the basics of multicellular evolution. These living fossils are ideal for studyi ...
'' Bergquist, 1961 * '' Suberites dandelenae'' Samaai & Maduray, 2017 * '' Suberites difficilis'' Dendy, 1897 * '' Suberites distortus'' Schmidt, 1870 * '' Suberites diversicolor'' Becking & Lim 2009 * '' Suberites domuncula'' (Olivi, 1792) * '' Suberites excellens'' (Thiele, 1898) * ''
Suberites ficus ''Suberites ficus'' is a species of sponge in the family Suberitidae. It is sometimes known as the sea orange sponge. Sponges are primitive animals with little apparent internal organisation. They are composed of a jellylike mesohyl sandwiched ...
'' (Johnston, 1842) * '' Suberites flabellatus'' Carter, 1886 * '' Suberites gibbosiceps'' Topsent, 1904 * '' Suberites glaber'' Hansen, 1885 * '' Suberites glasenapii'' Merejkowski, 1879 * '' Suberites globosus'' Carter, 1886 * '' Suberites heros'' Schmidt, 1870 * '' Suberites hirsutus'' Topsent, 1927 * '' Suberites holgeri'' Van Soest & Hooper, 2020 * '' Suberites hystrix'' Schmidt, 1868 * '' Suberites insignis'' Carter, 1886 * '' Suberites japonicus'' Thiele, 1898 * '' Suberites kelleri'' Burton, 1930 * '' Suberites lambei'' Austin et al., 2014 * '' Suberites laticeps'' Topsent, 1904 * '' Suberites latus'' Lambe, 1893 * '' Suberites lobatus'' (Wilson, 1902) * '' Suberites luna'' Giraldes & Goodwin, 2020 * '' Suberites luridus'' Solé-Cava & Thorpe, 1986 * '' Suberites lutea'' Sole-Cava & Thorpe, 1986 * '' Suberites mammilaris'' Sim & Kim, 1994 * '' Suberites massa'' Nardo, 1847 * '' Suberites microstomus'' Ridley & Dendy, 1887 * '' Suberites mineri'' (de Laubenfels, 1935) * '' Suberites mollis'' Ridley & Dendy, 1886 * ''
Suberites montalbidus ''Suberites'' is a genus of sea sponges in the family Suberitidae. Sponges, known scientifically as ''Porifera'', are the oldest metazoans and are used to elucidate the basics of multicellular evolution. These living fossils are ideal for studyi ...
'' Carter, 1880 * '' Suberites pagurorum'' Solé-Cava & Thorpe, 1986 * '' Suberites paradoxus'' Wilson, 1931 * '' Suberites perfectus'' Ridley & Dendy, 1886 * ''
Suberites pisiformis ''Suberites'' is a genus of sea sponges in the family Suberitidae. Sponges, known scientifically as ''Porifera'', are the oldest metazoans and are used to elucidate the basics of multicellular evolution. These living fossils are ideal for studyi ...
'' Lévi, 1993 * '' Suberites placenta'' Thiele, 1898 * '' Suberites prototypus'' Czerniavsky, 1880 * '' Suberites puncturatus'' Thiele, 1905 * '' Suberites purpura'' Fortunato, Pérez & Lôbo-Hajdu, 2020 * '' Suberites radiatus'' Kieschnick, 1896 * ''
Suberites ramosus ''Suberites'' is a genus of sea sponges in the family Suberitidae. Sponges, known scientifically as ''Porifera'', are the oldest metazoans and are used to elucidate the basics of multicellular evolution. These living fossils are ideal for studyi ...
'' Brøndsted, 1924 * ''
Suberites rhaphidiophorus ''Suberites'' is a genus of sea sponges in the family Suberitidae. Sponges, known scientifically as ''Porifera'', are the oldest metazoans and are used to elucidate the basics of multicellular evolution. These living fossils are ideal for studyi ...
'' (Brøndsted, 1924) * '' Suberites ruber'' Thiele, 1905 * '' Suberites rubrus'' Sole-Cava & Thorpe, 1986 * '' Suberites senilis'' Ridley & Dendy, 1886 * '' Suberites sericeus'' Thiele, 1898 * '' Suberites spermatozoon'' (Schmidt, 1875) * '' Suberites spirastrelloides'' Dendy, 1897 * '' Suberites spongiosus'' Schmidt, 1868 * '' Suberites stilensis'' Burton, 1933 * '' Suberites strongylatus'' Sarà, 1978 * '' Suberites suberia'' (Montagu, 1818) * '' Suberites syringella'' (Schmidt, 1868) * '' Suberites topsenti'' (Burton, 1929) * '' Suberites tortuosus'' Lévi, 1959 * '' Suberites tylobtusus'' Lévi, 1958 * '' Suberites verrilli'' Van Soest & Hooper, 2020 * '' Suberites virgultosus'' (Johnston, 1842)


References

{{Taxonbar, from=Q3463146 Suberitidae