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Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a
basal animal Marine life, sea life, or ocean life is the plants, animals and other organisms that live in the salt water of seas or oceans, or the brackish water of coastal estuaries. At a fundamental level, marine life affects the nature of the planet. ...
clade as a sister of the diploblasts. They are
multicellular 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 un ...
organisms that have bodies full of pores and channels allowing water to circulate through them, consisting of jelly-like
mesohyl The mesohyl, formerly known as mesenchyme or as mesoglea, is the gelatinous matrix within a sponge. It fills the space between the external pinacoderm and the internal choanoderm. The mesohyl resembles a type of connective tissue and contains s ...
sandwiched between two thin layers of cells. Sponges have unspecialized cells that can transform into other types and that often migrate between the main cell layers and the mesohyl in the process. Sponges do not have nervous, digestive or
circulatory system The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, tha ...
s. Instead, most rely on maintaining a constant water flow through their bodies to obtain food and oxygen and to remove wastes. Sponges were first to branch off the
evolutionary tree A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
from the
last common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...
of all animals, making them the sister group of all other animals.


Etymology

The term ''sponge'' derives from the
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic p ...
word ( 'sponge').


Overview

Sponges are similar to other animals in that they are
multicellular 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 un ...
,
heterotroph A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
ic, lack cell walls and produce sperm cells. Unlike other animals, they lack true tissues and
organs In biology, an organ is a collection of tissues joined in a structural unit to serve a common function. In the hierarchy of life, an organ lies between tissue and an organ system. Tissues are formed from same type cells to act together in a f ...
. Some of them are radially symmetrical, but most are asymmetrical. The shapes of their bodies are adapted for maximal efficiency of water flow through the central cavity, where the water deposits nutrients and then leaves through a hole called the
osculum The osculum (plural "oscula") is an excretory structure in the living sponge, a large opening to the outside through which the current of water exits after passing through the spongocoel. Wastes diffuse into the water and the water is pumped th ...
. Many sponges have internal skeletons of spicules (skeletal-like fragments of calcium carbonate or
silicon dioxide 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 one ...
), and/or
spongin Spongin, a modified type of collagen protein, forms the fibrous skeleton of most organisms among the phylum Porifera, the sponges. It is secreted by sponge cells known as spongocytes. Spongin gives a sponge its flexibility. True spongin is found ...
(a modified type of collagen protein). All adult sponges are
sessile Sessility, or sessile, may refer to: * Sessility (motility), organisms which are not able to move about * Sessility (botany), flowers or leaves that grow directly from the stem or peduncle of a plant * Sessility (medicine), tumors and polyps that ...
aquatic animals, meaning that they attach to an underwater surface and remain fixed in place (i.e., do not travel) while in
larval stage A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
of life they are
motile Motility is the ability of an organism to move independently, using metabolic energy. Definitions Motility, the ability of an organism to move independently, using metabolic energy, can be contrasted with sessility, the state of organisms th ...
. Although there are freshwater species, the great majority are marine (salt-water) species, ranging in habitat from tidal zones to depths exceeding . Although most of the approximately 5,000–10,000 known species of sponges feed on
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
and other microscopic food in the water, some host
photosynthesizing Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
microorganisms as
endosymbiont An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within ...
s, and these alliances often produce more food and oxygen than they consume. A few species of sponges that live in food-poor environments have evolved as
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other s ...
s that prey mainly on small
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group can ...
s. Most species use
sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote th ...
, releasing sperm cells into the water to fertilize
ova , abbreviated as OVA and sometimes as OAV (original animation video), are Japanese animated films and series made specially for release in home video formats without prior showings on television or in theaters, though the first part of an OVA s ...
that in some species are released and in others are retained by the "mother". The fertilized eggs develop into
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. ...
e, which swim off in search of places to settle. Sponges are known for regenerating from fragments that are broken off, although this only works if the fragments include the right types of cells. A few species reproduce by budding. When environmental conditions become less hospitable to the sponges, for example as temperatures drop, many freshwater species and a few marine ones produce
gemmule Gemmules are internal buds found in sponges and are involved in asexual reproduction. It is an asexually reproduced mass of cells, that is capable of developing into a new organism i.e., an adult sponge. Role in asexual reproduction Asexual ...
s, "survival pods" of unspecialized cells that remain dormant until conditions improve; they then either form completely new sponges or recolonize the skeletons of their parents. In most sponges, an internal gelatinous matrix called mesohyl functions as an
endoskeleton An endoskeleton (From Greek ἔνδον, éndon = "within", "inner" + σκελετός, skeletos = "skeleton") is an internal support structure of an animal, composed of mineralized tissue. Overview An endoskeleton is a skeleton that is on the ...
, and it is the only skeleton in soft sponges that encrust such hard surfaces as rocks. More commonly, the mesohyl is stiffened by
mineral In geology and mineralogy, a mineral or mineral species is, broadly speaking, a solid chemical compound with a fairly well-defined chemical composition and a specific crystal structure that occurs naturally in pure form.John P. Rafferty, ed. (2 ...
spicules, by spongin fibers, or both. Demosponges use spongin; many species have
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 ...
spicules, whereas some species have calcium carbonate
exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the ...
s.
Demosponges Demosponges (Demospongiae) are the most diverse class in the phylum Porifera. They include 76.2% of all species of sponges with nearly 8,800 species worldwide (World Porifera Database). They are sponges with a soft body that covers a hard, ...
constitute about 90% of all known sponge species, including all freshwater ones, and they have the widest range of habitats. Calcareous sponges, which have calcium carbonate spicules and, in some species, calcium carbonate exoskeletons, are restricted to relatively shallow marine waters where production of calcium carbonate is easiest. The fragile
glass sponge 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 consider ...
s, with "
scaffolding Scaffolding, also called scaffold or staging, is a temporary structure used to support a work crew and materials to aid in the construction, maintenance and repair of buildings, bridges and all other man-made structures. Scaffolds are widely use ...
" of silica spicules, are restricted to polar regions and the ocean depths where predators are rare. Fossils of all of these types have been found in rocks dated from . In addition
Archaeocyathid Archaeocyatha (or archaeocyathids 'ancient cups') is a taxon of extinct, sessile, reef-building marine sponges that lived in warm tropical and subtropical waters during the Cambrian Period. It is believed that the centre of the Archaeocyatha or ...
s, whose fossils are common in rocks from , are now regarded as a type of sponge. The single-celled choanoflagellates resemble the choanocyte cells of sponges which are used to drive their water flow systems and capture most of their food. This along with phylogenetic studies of ribosomal molecules have been used as morphological evidence to suggest sponges are the sister group to the rest of animals. Some studies have shown that sponges do not form a monophyletic group, in other words do not include ''all and only'' the descendants of a common ancestor. Recent phylogenetic analyses suggested that
comb jellies Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), and ...
rather than sponges are the sister group to the rest of animals. However reanalysis of the data showed that the computer algorithms used for analysis were misled by the presence of specific ctenophore genes that were markedly different from those of other species, leaving sponges as either the sister group to all other animals, or an ancestral paraphyletic grade. The few species of demosponge that have entirely soft fibrous skeletons with no hard elements have been used by humans over thousands of years for several purposes, including as padding and as cleaning tools. By the 1950s, though, these had been
overfished Overfishing is the removal of a species of fish (i.e. fishing) from a body of water at a rate greater than that the species can replenish its population naturally (i.e. the overexploitation of the fishery's existing fish stock), resulting in t ...
so heavily that the industry almost collapsed, and most sponge-like materials are now synthetic. Sponges and their microscopic endosymbionts are now being researched as possible sources of medicines for treating a wide range of diseases.
Dolphin A dolphin is an aquatic mammal within the infraorder Cetacea. Dolphin species belong to the families Delphinidae (the oceanic dolphins), Platanistidae (the Indian river dolphins), Iniidae (the New World river dolphins), Pontoporiidae (the ...
s have been observed using sponges as tools while
foraging Foraging is searching for wild food resources. It affects an animal's fitness because it plays an important role in an animal's ability to survive and reproduce. Foraging theory is a branch of behavioral ecology that studies the foraging behavi ...
.


Distinguishing features

Sponges constitute the phylum Porifera, and have been defined as
sessile Sessility, or sessile, may refer to: * Sessility (motility), organisms which are not able to move about * Sessility (botany), flowers or leaves that grow directly from the stem or peduncle of a plant * Sessility (medicine), tumors and polyps that ...
metazoa 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 ...
ns (multicelled immobile animals) that have water intake and outlet openings connected by chambers lined with
choanocyte Choanocytes (also known as "collar cells") are cells that line the interior of asconoid, syconoid and leuconoid body types of sponges that contain a central flagellum, or ''cilium,'' surrounded by a collar of microvilli which are connected by a ...
s, cells with whip-like flagella. However, a few carnivorous sponges have lost these water flow systems and the choanocytes. All known living sponges can remold their bodies, as most types of their cells can move within their bodies and a few can change from one type to another. Even if a few sponges are able to produce mucus – which acts as a microbial barrier in all other animals – no sponge with the ability to secrete a functional mucus layer has been recorded. Without such a mucus layer their living tissue is covered by a layer of microbial symbionts, which can contribute up to 40–50% of the sponge wet mass. This inability to prevent microbes from penetrating their porous tissue could be a major reason why they have never evolved a more complex anatomy. Like
cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that ...
ns (jellyfish, etc.) and
ctenophores Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), a ...
(comb jellies), and unlike all other known metazoans, sponges' bodies consist of a non-living jelly-like mass (
mesohyl The mesohyl, formerly known as mesenchyme or as mesoglea, is the gelatinous matrix within a sponge. It fills the space between the external pinacoderm and the internal choanoderm. The mesohyl resembles a type of connective tissue and contains s ...
) sandwiched between two main layers of cells. Cnidarians and ctenophores have simple nervous systems, and their cell layers are bound by internal connections and by being mounted on a basement membrane (thin fibrous mat, also known as " basal lamina"). Sponges have no nervous systems, their middle jelly-like layers have large and varied populations of cells, and some types of cells in their outer layers may move into the middle layer and change their functions.


Basic structure


Cell types

A sponge's body is hollow and is held in shape by the
mesohyl The mesohyl, formerly known as mesenchyme or as mesoglea, is the gelatinous matrix within a sponge. It fills the space between the external pinacoderm and the internal choanoderm. The mesohyl resembles a type of connective tissue and contains s ...
, a jelly-like substance made mainly of collagen and reinforced by a dense network of fibers also made of collagen. The inner surface is covered with
choanocyte Choanocytes (also known as "collar cells") are cells that line the interior of asconoid, syconoid and leuconoid body types of sponges that contain a central flagellum, or ''cilium,'' surrounded by a collar of microvilli which are connected by a ...
s, cells with cylindrical or conical collars surrounding one
flagellum A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...
per choanocyte. The wave-like motion of the whip-like flagella drives water through the sponge's body. All sponges have ostia, channels leading to the interior through the mesohyl, and in most sponges these are controlled by tube-like
porocytes Porocytes are tubular cells which make up the pores of a sponge known as ostia. Description Covering the sponge is a layer of cells known as the pinacoderm, which is composed of pinacocytes. In a sponge, pinacocytes are a thin, elastic layer whic ...
that form closable inlet valves. Pinacocytes, plate-like cells, form a single-layered external skin over all other parts of the mesohyl that are not covered by choanocytes, and the pinacocytes also digest food particles that are too large to enter the ostia, while those at the base of the animal are responsible for anchoring it. Other types of cell live and move within the mesohyl: * Lophocytes are
amoeba An amoeba (; less commonly spelled ameba or amœba; plural ''am(o)ebas'' or ''am(o)ebae'' ), often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and retracting pseudop ...
-like cells that move slowly through the mesohyl and secrete collagen fibres. * Collencytes are another type of collagen-producing cell. * Rhabdiferous cells secrete polysaccharides that also form part of the mesohyl. *
Oocyte An oocyte (, ), oöcyte, or ovocyte is a female gametocyte or germ cell involved in reproduction. In other words, it is an immature ovum, or egg cell. An oocyte is produced in a female fetus in the ovary during female gametogenesis. The femal ...
s and
spermatocyte Spermatocytes are a type of male gametocyte in animals. They derive from immature germ cells called spermatogonia. They are found in the testis, in a structure known as the seminiferous tubules. There are two types of spermatocytes, primary and ...
s are reproductive cells. * Sclerocytes secrete the mineralized spicules ("little spines") that form the skeletons of many sponges and in some species provide some defense against predators. * In addition to or instead of sclerocytes, demosponges have spongocytes that secrete a form of collagen that
polymer A polymer (; Greek '' poly-'', "many" + ''-mer'', "part") is a substance or material consisting of very large molecules called macromolecules, composed of many repeating subunits. Due to their broad spectrum of properties, both synthetic a ...
izes into
spongin Spongin, a modified type of collagen protein, forms the fibrous skeleton of most organisms among the phylum Porifera, the sponges. It is secreted by sponge cells known as spongocytes. Spongin gives a sponge its flexibility. True spongin is found ...
, a thick fibrous material that stiffens the mesohyl. *
Myocyte A muscle cell is also known as a myocyte when referring to either a cardiac muscle cell (cardiomyocyte), or a smooth muscle cell as these are both small cells. A skeletal muscle cell is long and threadlike with many nuclei and is called a mus ...
s ("muscle cells") conduct signals and cause parts of the animal to contract. * "Grey cells" act as sponges' equivalent of an
immune system The immune system is a network of biological processes that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to parasitic worms, as well as cancer cells and objects such as wood splint ...
. * Archaeocytes (or
amoebocytes An amebocyte or amoebocyte () is a mobile cell (moving like an amoeba) in the body of invertebrates including cnidaria, echinoderms, molluscs, tunicates, sponges and some chelicerates. They move by pseudopodia. Similarly to some of the white bloo ...
) are
amoeba An amoeba (; less commonly spelled ameba or amœba; plural ''am(o)ebas'' or ''am(o)ebae'' ), often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and retracting pseudop ...
-like cells that are
totipotent 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 other words each is capable of transformation into any other type of cell. They also have important roles in feeding and in clearing debris that block the ostia. Many larval sponges possess neuron-less eyes that are based on
cryptochrome Cryptochromes (from the Greek κρυπτός χρώμα, "hidden colour") are a class of flavoproteins found in plants and animals that are sensitive to blue light. They are involved in the circadian rhythms and the sensing of magnetic fields ...
s. They mediate phototaxic behavior.


Glass sponges' syncytia

Glass sponge 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 consider ...
s present a distinctive variation on this basic plan. Their spicules, which are made 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 ...
, form a
scaffolding Scaffolding, also called scaffold or staging, is a temporary structure used to support a work crew and materials to aid in the construction, maintenance and repair of buildings, bridges and all other man-made structures. Scaffolds are widely use ...
-like framework between whose rods the living tissue is suspended like a
cobweb A spider web, spiderweb, spider's web, or cobweb (from the archaic word '' coppe'', meaning "spider") is a structure created by a spider out of proteinaceous spider silk extruded from its spinnerets, generally meant to catch its prey. Spi ...
that contains most of the cell types. This tissue is a
syncytium 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 nucleu ...
that in some ways behaves like many cells that share a single external
membrane A membrane is a selective barrier; it allows some things to pass through but stops others. Such things may be molecules, ions, or other small particles. Membranes can be generally classified into synthetic membranes and biological membranes. ...
, and in others like a single cell with multiple nuclei. The mesohyl is absent or minimal. The syncytium's
cytoplasm In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. ...
, the soupy fluid that fills the interiors of cells, is organized into "rivers" that transport nuclei, organelles ("organs" within cells) and other substances. Instead of choanocytes, they have further syncytia, known as choanosyncytia, which form bell-shaped chambers where water enters via perforations. The insides of these chambers are lined with "collar bodies", each consisting of a collar and flagellum but without a nucleus of its own. The motion of the flagella sucks water through passages in the "cobweb" and expels it via the open ends of the bell-shaped chambers. Some types of cells have a single nucleus and membrane each, but are connected to other single-nucleus cells and to the main syncytium by "bridges" made of
cytoplasm In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. ...
. The sclerocytes that build spicules have multiple nuclei, and in glass sponge larvae they are connected to other tissues by cytoplasm bridges; such connections between sclerocytes have not so far been found in adults, but this may simply reflect the difficulty of investigating such small-scale features. The bridges are controlled by "plugged junctions" that apparently permit some substances to pass while blocking others.


Water flow and body structures

Most sponges work rather like chimneys: they take in water at the bottom and eject it from the
osculum The osculum (plural "oscula") is an excretory structure in the living sponge, a large opening to the outside through which the current of water exits after passing through the spongocoel. Wastes diffuse into the water and the water is pumped th ...
("little mouth") at the top. Since ambient currents are faster at the top, the suction effect that they produce by
Bernoulli's principle In fluid dynamics, Bernoulli's principle states that an increase in the speed of a fluid occurs simultaneously with a decrease in static pressure or a decrease in the fluid's potential energy. The principle is named after the Swiss mathematici ...
does some of the work for free. Sponges can control the water flow by various combinations of wholly or partially closing the osculum and ostia (the intake pores) and varying the beat of the flagella, and may shut it down if there is a lot of sand or silt in the water. Although the layers of pinacocytes and
choanocyte Choanocytes (also known as "collar cells") are cells that line the interior of asconoid, syconoid and leuconoid body types of sponges that contain a central flagellum, or ''cilium,'' surrounded by a collar of microvilli which are connected by a ...
s resemble the
epithelia Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellula ...
of more complex animals, they are not bound tightly by cell-to-cell connections or a basal lamina (thin fibrous sheet underneath). The flexibility of these layers and re-modeling of the mesohyl by lophocytes allow the animals to adjust their shapes throughout their lives to take maximum advantage of local water currents. The simplest body structure in sponges is a tube or vase shape known as "asconoid", but this severely limits the size of the animal. The body structure is characterized by a stalk-like spongocoel surrounded by a single layer of choanocytes. If it is simply scaled up, the ratio of its volume to surface area increases, because surface increases as the square of length or width while volume increases proportionally to the cube. The amount of tissue that needs food and
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as ...
is determined by the volume, but the pumping capacity that supplies food and oxygen depends on the area covered by choanocytes. Asconoid sponges seldom exceed in diameter. Some sponges overcome this limitation by adopting the "syconoid" structure, in which the body wall is pleated. The inner pockets of the pleats are lined with choanocytes, which connect to the outer pockets of the pleats by ostia. This increase in the number of choanocytes and hence in pumping capacity enables syconoid sponges to grow up to a few centimeters in diameter. The "leuconoid" pattern boosts pumping capacity further by filling the interior almost completely with mesohyl that contains a network of chambers lined with choanocytes and connected to each other and to the water intakes and outlet by tubes. Leuconid sponges grow to over in diameter, and the fact that growth in any direction increases the number of choanocyte chambers enables them to take a wider range of forms, for example "encrusting" sponges whose shapes follow those of the surfaces to which they attach. All freshwater and most shallow-water marine sponges have leuconid bodies. The networks of water passages in
glass sponge 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 consider ...
s are similar to the leuconid structure. In all three types of structure the cross-section area of the choanocyte-lined regions is much greater than that of the intake and outlet channels. This makes the flow slower near the choanocytes and thus makes it easier for them to trap food particles. For example, in '' Leuconia'', a small leuconoid sponge about tall and in diameter, water enters each of more than 80,000 intake canals at 6 cm per ''minute''. However, because ''Leuconia'' has more than 2 million flagellated chambers whose combined diameter is much greater than that of the canals, water flow through chambers slows to 3.6 cm per ''hour'', making it easy for choanocytes to capture food. All the water is expelled through a single
osculum The osculum (plural "oscula") is an excretory structure in the living sponge, a large opening to the outside through which the current of water exits after passing through the spongocoel. Wastes diffuse into the water and the water is pumped th ...
at about 8.5 cm per ''second'', fast enough to carry waste products some distance away.


Skeleton

In zoology a skeleton is any fairly rigid structure of an animal, irrespective of whether it has joints and irrespective of whether it is biomineralized. The mesohyl functions as an
endoskeleton An endoskeleton (From Greek ἔνδον, éndon = "within", "inner" + σκελετός, skeletos = "skeleton") is an internal support structure of an animal, composed of mineralized tissue. Overview An endoskeleton is a skeleton that is on the ...
in most sponges, and is the only skeleton in soft sponges that encrust hard surfaces such as rocks. More commonly the mesohyl is stiffened by mineral spicules, by spongin fibers or both. Spicules, which are present in most but not all species, may be made 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 ...
or calcium carbonate, and vary in shape from simple rods to three-dimensional "stars" with up to six rays. Spicules are produced by sclerocyte cells, and may be separate, connected by joints, or fused. Some sponges also secrete
exoskeleton An exoskeleton (from Greek ''éxō'' "outer" and ''skeletós'' "skeleton") is an external skeleton that supports and protects an animal's body, in contrast to an internal skeleton (endoskeleton) in for example, a human. In usage, some of the ...
s that lie completely outside their organic components. For example, sclerosponges ("hard sponges") have massive calcium carbonate exoskeletons over which the organic matter forms a thin layer with
choanocyte Choanocytes (also known as "collar cells") are cells that line the interior of asconoid, syconoid and leuconoid body types of sponges that contain a central flagellum, or ''cilium,'' surrounded by a collar of microvilli which are connected by a ...
chambers in pits in the mineral. These exoskeletons are secreted by the pinacocytes that form the animals' skins.


Vital functions


Movement

Although adult sponges are fundamentally
sessile Sessility, or sessile, may refer to: * Sessility (motility), organisms which are not able to move about * Sessility (botany), flowers or leaves that grow directly from the stem or peduncle of a plant * Sessility (medicine), tumors and polyps that ...
animals, some marine and freshwater species can move across the sea bed at speeds of per day, as a result of
amoeba An amoeba (; less commonly spelled ameba or amœba; plural ''am(o)ebas'' or ''am(o)ebae'' ), often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and retracting pseudop ...
-like movements of pinacocytes and other cells. A few species can contract their whole bodies, and many can close their oscula and ostia. Juveniles drift or swim freely, while adults are stationary.


Respiration, feeding and excretion

Sponges do not have distinct circulatory,
respiratory The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies gre ...
, digestive, and
excretory Excretion is a process in which metabolic waste is eliminated from an organism. In vertebrates this is primarily carried out by the lungs, kidneys, and skin. This is in contrast with secretion, where the substance may have specific tasks aft ...
systems – instead the water flow system supports all these functions. They
filter Filter, filtering or filters may refer to: Science and technology Computing * Filter (higher-order function), in functional programming * Filter (software), a computer program to process a data stream * Filter (video), a software component tha ...
food particles out of the water flowing through them. Particles larger than 50 micrometers cannot enter the ostia and pinacocytes consume them by
phagocytosis Phagocytosis () is the process by which a cell uses its plasma membrane to engulf a large particle (≥ 0.5 μm), giving rise to an internal compartment called the phagosome. It is one type of endocytosis. A cell that performs phagocytosis is ...
(engulfing and intracellular digestion). Particles from 0.5 μm to 50 μm are trapped in the ostia, which taper from the outer to inner ends. These particles are consumed by pinacocytes or by
archaeocyte Archaeocytes (from Greek '' archaios'' "beginning" and '' kytos'' "hollow vessel") or amoebocytes are amoeboid cells found in sponges. They are totipotent and have varied functions depending on the species. The structure of these cells match to ...
s which partially extrude themselves through the walls of the ostia. Bacteria-sized particles, below 0.5 micrometers, pass through the ostia and are caught and consumed by
choanocyte Choanocytes (also known as "collar cells") are cells that line the interior of asconoid, syconoid and leuconoid body types of sponges that contain a central flagellum, or ''cilium,'' surrounded by a collar of microvilli which are connected by a ...
s. Since the smallest particles are by far the most common, choanocytes typically capture 80% of a sponge's food supply. Archaeocytes transport food packaged in vesicles from cells that directly digest food to those that do not. At least one species of sponge has internal fibers that function as tracks for use by nutrient-carrying archaeocytes, and these tracks also move inert objects. It used to be claimed that
glass sponge 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 consider ...
s could live on nutrients dissolved in sea water and were very averse to silt. However, a study in 2007 found no evidence of this and concluded that they extract bacteria and other micro-organisms from water very efficiently (about 79%) and process suspended sediment grains to extract such prey. Collar bodies digest food and distribute it wrapped in vesicles that are transported by dynein "motor" molecules along bundles of microtubules that run throughout the
syncytium 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 nucleu ...
. Sponges' cells absorb oxygen by
diffusion Diffusion is the net movement of anything (for example, atoms, ions, molecules, energy) generally from a region of higher concentration to a region of lower concentration. Diffusion is driven by a gradient in Gibbs free energy or chemica ...
from water into cells as water flows through body, into which
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is trans ...
and other soluble waste products such as
ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous wa ...
also diffuse. Archeocytes remove mineral particles that threaten to block the ostia, transport them through the mesohyl and generally dump them into the outgoing water current, although some species incorporate them into their skeletons.


Carnivorous sponges

In waters where the supply of food particles is very poor, some species prey on crustaceans and other small animals. So far only 137 species have been discovered. Most belong to 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 ...
Cladorhizidae, but a few members of the Guitarridae and
Esperiopsidae Esperiopsidae is a family of marine demosponge Demosponges (Demospongiae) are the most diverse class in the phylum Porifera. They include 76.2% of all species of sponges with nearly 8,800 species worldwide (World Porifera Database). The ...
are also carnivores. In most cases little is known about how they actually capture prey, although some species are thought to use either sticky threads or hooked spicules. Most carnivorous sponges live in deep waters, up to , and the development of deep-ocean exploration techniques is expected to lead to the discovery of several more. However, one species has been found in
Mediterranean 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 Europe, Western and Southern Europe and Anatolia, on the south by North Africa ...
caves at depths of , alongside the more usual
filter feeding 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 ...
sponges. The cave-dwelling predators capture crustaceans under long by entangling them with fine threads, digest them by enveloping them with further threads over the course of a few days, and then return to their normal shape; there is no evidence that they use
venom Venom or zootoxin is a type of toxin produced by an animal that is actively delivered through a wound by means of a bite, sting, or similar action. The toxin is delivered through a specially evolved ''venom apparatus'', such as fangs or a st ...
. Most known carnivorous sponges have completely lost the water flow system and
choanocyte Choanocytes (also known as "collar cells") are cells that line the interior of asconoid, syconoid and leuconoid body types of sponges that contain a central flagellum, or ''cilium,'' surrounded by a collar of microvilli which are connected by a ...
s. However, the
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 ...
'' Chondrocladia'' uses a highly modified water flow system to inflate balloon-like structures that are used for capturing prey.


Endosymbionts

Freshwater sponges often host green algae as
endosymbiont An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within ...
s within
archaeocyte Archaeocytes (from Greek '' archaios'' "beginning" and '' kytos'' "hollow vessel") or amoebocytes are amoeboid cells found in sponges. They are totipotent and have varied functions depending on the species. The structure of these cells match to ...
s and other cells, and benefit from nutrients produced by the algae. Many marine species host other
photosynthesizing Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
organisms, most commonly cyanobacteria but in some cases dinoflagellates. Symbiotic cyanobacteria may form a third of the total mass of living tissue in some sponges, and some sponges gain 48% to 80% of their energy supply from these micro-organisms. In 2008 a University of Stuttgart team reported that spicules made 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 ...
conduct light into the
mesohyl The mesohyl, formerly known as mesenchyme or as mesoglea, is the gelatinous matrix within a sponge. It fills the space between the external pinacoderm and the internal choanoderm. The mesohyl resembles a type of connective tissue and contains s ...
, where the photosynthesizing endosymbionts live. Sponges that host photosynthesizing organisms are most common in waters with relatively poor supplies of food particles, and often have leafy shapes that maximize the amount of sunlight they collect. A recently discovered carnivorous sponge that lives near hydrothermal vents hosts methane-eating bacteria, and digests some of them.


"Immune" system

Sponges do not have the complex
immune system The immune system is a network of biological processes that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to parasitic worms, as well as cancer cells and objects such as wood splint ...
s of most other animals. However, they reject grafts from other species but accept them from other members of their own species. In a few marine species, gray cells play the leading role in rejection of foreign material. When invaded, they produce a chemical that stops movement of other cells in the affected area, thus preventing the intruder from using the sponge's internal transport systems. If the intrusion persists, the grey cells concentrate in the area and release toxins that kill all cells in the area. The "immune" system can stay in this activated state for up to three weeks.


Reproduction


Asexual

Sponges have three asexual methods of reproduction: after fragmentation, by
budding Budding or blastogenesis is a type of asexual reproduction in which a new organism develops from an outgrowth or bud due to cell division at one particular site. For example, the small bulb-like projection coming out from the yeast cell is kno ...
, and by producing
gemmule Gemmules are internal buds found in sponges and are involved in asexual reproduction. It is an asexually reproduced mass of cells, that is capable of developing into a new organism i.e., an adult sponge. Role in asexual reproduction Asexual ...
s. Fragments of sponges may be detached by currents or waves. They use the mobility of their pinacocytes and
choanocyte Choanocytes (also known as "collar cells") are cells that line the interior of asconoid, syconoid and leuconoid body types of sponges that contain a central flagellum, or ''cilium,'' surrounded by a collar of microvilli which are connected by a ...
s and reshaping of the
mesohyl The mesohyl, formerly known as mesenchyme or as mesoglea, is the gelatinous matrix within a sponge. It fills the space between the external pinacoderm and the internal choanoderm. The mesohyl resembles a type of connective tissue and contains s ...
to re-attach themselves to a suitable surface and then rebuild themselves as small but functional sponges over the course of several days. The same capabilities enable sponges that have been squeezed through a fine cloth to regenerate. A sponge fragment can only regenerate if it contains both collencytes to produce
mesohyl The mesohyl, formerly known as mesenchyme or as mesoglea, is the gelatinous matrix within a sponge. It fills the space between the external pinacoderm and the internal choanoderm. The mesohyl resembles a type of connective tissue and contains s ...
and archeocytes to produce all the other cell types. A very few species reproduce by budding. Gemmules are "survival pods" which a few marine sponges and many freshwater species produce by the thousands when dying and which some, mainly freshwater species, regularly produce in autumn. Spongocytes make gemmules by wrapping shells of spongin, often reinforced with spicules, round clusters of archeocytes that are full of nutrients. Freshwater gemmules may also include photosynthesizing symbionts. The gemmules then become dormant, and in this state can survive cold, drying out, lack of oxygen and extreme variations in salinity. Freshwater gemmules often do not revive until the temperature drops, stays cold for a few months and then reaches a near-"normal" level. When a gemmule germinates, the archeocytes round the outside of the cluster transform into pinacocytes, a membrane over a pore in the shell bursts, the cluster of cells slowly emerges, and most of the remaining archeocytes transform into other cell types needed to make a functioning sponge. Gemmules from the same species but different individuals can join forces to form one sponge. Some gemmules are retained within the parent sponge, and in spring it can be difficult to tell whether an old sponge has revived or been "recolonized" by its own gemmules.


Sexual

Most sponges are hermaphrodites (function as both sexes simultaneously), although sponges have no gonads (reproductive organs). Sperm are produced by
choanocyte Choanocytes (also known as "collar cells") are cells that line the interior of asconoid, syconoid and leuconoid body types of sponges that contain a central flagellum, or ''cilium,'' surrounded by a collar of microvilli which are connected by a ...
s or entire choanocyte chambers that sink into the
mesohyl The mesohyl, formerly known as mesenchyme or as mesoglea, is the gelatinous matrix within a sponge. It fills the space between the external pinacoderm and the internal choanoderm. The mesohyl resembles a type of connective tissue and contains s ...
and form spermatic
cyst A cyst is a closed sac, having a distinct envelope and division compared with the nearby tissue. Hence, it is a cluster of cells that have grouped together to form a sac (like the manner in which water molecules group together to form a bubble) ...
s while eggs are formed by transformation of archeocytes, or of choanocytes in some species. Each egg generally acquires a
yolk Among animals which produce eggs, the yolk (; also known as the vitellus) is the nutrient-bearing portion of the egg whose primary function is to supply food for the development of the embryo. Some types of egg contain no yolk, for example ...
by consuming "nurse cells". During spawning, sperm burst out of their cysts and are expelled via the
osculum The osculum (plural "oscula") is an excretory structure in the living sponge, a large opening to the outside through which the current of water exits after passing through the spongocoel. Wastes diffuse into the water and the water is pumped th ...
. If they contact another sponge of the same species, the water flow carries them to choanocytes that engulf them but, instead of digesting them, metamorphose to an
ameboid An amoeba (; less commonly spelled ameba or amœba; plural ''am(o)ebas'' or ''am(o)ebae'' ), often called an amoeboid, is a type of cell or unicellular organism with the ability to alter its shape, primarily by extending and retracting pseudopo ...
form and carry the sperm through the mesohyl to eggs, which in most cases engulf the carrier and its cargo. A few species release fertilized eggs into the water, but most retain the eggs until they hatch. There are four types of larvae, but all are balls of cells with an outer layer of cells whose flagellae or cilia enable the larvae to move. After swimming for a few days the larvae sink and crawl until they find a place to settle. Most of the cells transform into archeocytes and then into the types appropriate for their locations in a miniature adult sponge.
Glass sponge 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 consider ...
embryos start by dividing into separate cells, but once 32 cells have formed they rapidly transform into larvae that externally are ovoid with a band of cilia round the middle that they use for movement, but internally have the typical glass sponge structure of spicules with a cobweb-like main syncitium draped around and between them and choanosyncytia with multiple collar bodies in the center. The larvae then leave their parents' bodies.


Life cycle

Sponges in
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout ...
regions live for at most a few years, but some
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
species and perhaps some deep-ocean ones may live for 200 years or more. Some calcified demosponges grow by only per year and, if that rate is constant, specimens wide must be about 5,000 years old. Some sponges start sexual reproduction when only a few weeks old, while others wait until they are several years old.


Coordination of activities

Adult sponges lack
neuron 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. ...
s or any other kind of
nervous tissue Nervous tissue, also called neural tissue, is the main tissue component of the nervous system. The nervous system regulates and controls body functions and activity. It consists of two parts: the central nervous system (CNS) comprising the brain ...
. However, most species have the ability to perform movements that are coordinated all over their bodies, mainly contractions of the pinacocytes, squeezing the water channels and thus expelling excess sediment and other substances that may cause blockages. Some species can contract the
osculum The osculum (plural "oscula") is an excretory structure in the living sponge, a large opening to the outside through which the current of water exits after passing through the spongocoel. Wastes diffuse into the water and the water is pumped th ...
independently of the rest of the body. Sponges may also contract in order to reduce the area that is vulnerable to attack by predators. In cases where two sponges are fused, for example if there is a large but still unseparated bud, these contraction waves slowly become coordinated in both of the " Siamese twins". The coordinating mechanism is unknown, but may involve chemicals similar to neurotransmitters. However,
glass sponge 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 consider ...
s rapidly transmit electrical impulses through all parts of the
syncytium 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 nucleu ...
, and use this to halt the motion of their flagella if the incoming water contains toxins or excessive sediment.
Myocyte A muscle cell is also known as a myocyte when referring to either a cardiac muscle cell (cardiomyocyte), or a smooth muscle cell as these are both small cells. A skeletal muscle cell is long and threadlike with many nuclei and is called a mus ...
s are thought to be responsible for closing the osculum and for transmitting signals between different parts of the body. Sponges contain
gene In biology, the word gene (from , ; "... Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a b ...
s very similar to those that contain the "recipe" for the post- synaptic density, an important signal-receiving structure in the neurons of all other animals. However, in sponges these genes are only activated in "flask cells" that appear only in larvae and may provide some sensory capability while the larvae are swimming. This raises questions about whether flask cells represent the predecessors of true neurons or are evidence that sponges' ancestors had true neurons but lost them as they adapted to a sessile lifestyle.


Ecology


Habitats

Sponges are worldwide in their distribution, living in a wide range of ocean habitats, from the polar regions to the tropics. Most live in quiet, clear waters, because sediment stirred up by waves or currents would block their pores, making it difficult for them to feed and breathe. The greatest numbers of sponges are usually found on firm surfaces such as rocks, but some sponges can attach themselves to soft sediment by means of a root-like base. Sponges are more abundant but less diverse in temperate waters than in tropical waters, possibly because organisms that prey on sponges are more abundant in tropical waters.
Glass sponge 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 consider ...
s are the most common in polar waters and in the depths of temperate and tropical seas, as their very porous construction enables them to extract food from these resource-poor waters with the minimum of effort. Demosponges and calcareous sponges are abundant and diverse in shallower non-polar waters. The different classes of sponge live in different ranges of habitat: :


As primary producers

Sponges with
photosynthesizing Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...
endosymbiont An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within ...
s produce up to three times more
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as ...
than they consume, as well as more organic matter than they consume. Such contributions to their habitats' resources are significant along Australia's
Great Barrier Reef The Great Barrier Reef is the world's largest coral reef system composed of over 2,900 individual reefs and 900 islands stretching for over over an area of approximately . The reef is located in the Coral Sea, off the coast of Queensland, ...
but relatively minor in the Caribbean.


Defenses

Many sponges shed spicules, forming a dense carpet several meters deep that keeps away
echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the s ...
s which would otherwise prey on the sponges. They also produce toxins that prevent other sessile organisms such as
bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a ...
ns or
sea squirt Ascidiacea, commonly known as the ascidians, tunicates (in part), and sea squirts (in part), is a polyphyletic class in the subphylum Tunicata of sac-like marine invertebrate filter feeders. Ascidians are characterized by a tough outer "tunic" m ...
s from growing on or near them, making sponges very effective competitors for living space. One of many examples includes
ageliferin Ageliferin is a chemical compound produced by some sponges. It was first isolated from Caribbean and then Okinawan marine sponges in the genus ''Agelas''. It often co-exists with the related compound sceptrin and other similar compounds. It has ...
. A few species, the Caribbean fire sponge ''
Tedania ''Tedania'' is a genus of sea sponges in the family Tedaniidae. Species The following species are recognised in the genus Tedania: * Subgenus ''Tedania (Stylotedania)'' Van Soest, 2017 ** ''Tedania folium'' Van Soest, 2017 * Subgenus ''Tedania ...
ignis'', cause a severe rash in humans who handle them. Turtles and some fish feed mainly on sponges. It is often said that sponges produce
chemical defense Chemical defense is a life history strategy employed by many organisms to avoid consumption by producing toxic or repellent metabolites or chemical warnings which incite defensive behavioral changes. The production of defensive chemicals occurs in ...
s against such predators. However, experiments have been unable to establish a relationship between the toxicity of chemicals produced by sponges and how they taste to fish, which would diminish the usefulness of chemical defenses as deterrents. Predation by fish may even help to spread sponges by detaching fragments. However, some studies have shown fish showing a preference for non chemically defended sponges, and another study found that high levels of coral predation did predict the presence of chemically defended species.
Glass sponge 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 consider ...
s produce no toxic chemicals, and live in very deep water where predators are rare.


Predation

Sponge flies, also known as spongilla-flies ( Neuroptera,
Sisyridae Sisyridae, commonly known as spongeflies or spongillaflies, are a family of winged insects in the order Neuroptera. There are approximately 60 living species described, and several extinct species identified from the fossil record. Description ...
), are specialist predators of freshwater sponges. The female lays her eggs on vegetation overhanging water. The larvae hatch and drop into the water where they seek out sponges to feed on. They use their elongated mouthparts to pierce the sponge and suck the fluids within. The larvae of some species cling to the surface of the sponge while others take refuge in the sponge's internal cavities. The fully grown larvae leave the water and spin a cocoon in which to pupate.


Bioerosion

The Caribbean chicken-liver sponge ''
Chondrilla nucula ''Chondrilla nucula'', sometimes called the Caribbean chicken-liver sponge, is a species of sea sponge belonging to the family Chondrillidae. It is an amorphous shaped sponge that grows in flat, sometimes bulbous sheets in benthic communities. ...
'' secretes toxins that kill coral
polyps A polyp in zoology is one of two forms found in the phylum Cnidaria, the other being the medusa. Polyps are roughly cylindrical in shape and elongated at the axis of the vase-shaped body. In solitary polyps, the aboral (opposite to oral) end i ...
, allowing the sponges to grow over the coral skeletons. Others, especially in the family
Clionaidae Clionaidae is a family of demosponges which are found worldwide. This family is known for parasitically boring holes in calcareous material such as mollusc shells and corals, using both chemical and mechanical processes.Brusca, R.C. & Brusca, G. ...
, use corrosive substances secreted by their archeocytes to tunnel into rocks, corals and the shells of dead
mollusk Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is e ...
s. Sponges may remove up to per year from reefs, creating visible notches just below low-tide level.


Diseases

Caribbean sponges of the genus '' Aplysina'' suffer from Aplysina red band syndrome. This causes ''Aplysina'' to develop one or more rust-colored bands, sometimes with adjacent bands of
necrotic Necrosis () is a form of cell injury which results in the premature death of cells in living tissue by autolysis. Necrosis is caused by factors external to the cell or tissue, such as infection, or trauma which result in the unregulated dige ...
tissue. These lesions may completely encircle branches of the sponge. The disease appears to be contagious and impacts approximately 10 percent of ''A. cauliformis'' on Bahamian reefs. The rust-colored bands are caused by a
cyanobacterium Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, blu ...
, but it is unknown whether this organism actually causes the disease.


Collaboration with other organisms

In addition to hosting photosynthesizing endosymbionts, sponges are noted for their wide range of collaborations with other organisms. The relatively large encrusting sponge '' Lissodendoryx colombiensis'' is most common on rocky surfaces, but has extended its range into
seagrass meadow A seagrass meadow or seagrass bed is an underwater ecosystem formed by seagrasses. Seagrasses are marine (saltwater) plants found in shallow coastal waters and in the brackish waters of estuaries. Seagrasses are flowering plants with stems and ...
s by letting itself be surrounded or overgrown by seagrass sponges, which are distasteful to the local starfish and therefore protect ''Lissodendoryx'' against them; in return the seagrass sponges get higher positions away from the sea-floor sediment. Shrimps of the genus '' Synalpheus'' form colonies in sponges, and each shrimp species inhabits a different sponge species, making ''Synalpheus'' one of the most diverse crustacean genera. Specifically, Synalpheus regalis utilizes the sponge not only as a food source, but also as a defense against other shrimp and predators. As many as 16,000 individuals inhabit a single
loggerhead sponge ''Spheciospongia vesparium'', commonly known as the loggerhead sponge, is a species of sea sponge belonging to the family Clionaidae. While it is highly toxic to many fish, this sponge is eaten by certain angelfish and is known to form part of th ...
, feeding off the larger particles that collect on the sponge as it filters the ocean to feed itself. Other crustaceans such as hermit crabs commonly have a specific species of sponge, ''
Pseudospongosorites ''Pseudospongosorites'' is a genus of sea sponges belonging to the family Suberitidae. Currently, the genus is considered as monotypic, consisting of a single species ''Pseudospongosorites suberitoides''. It is found in the Caribbean Sea, the Gul ...
'', grow on them as both the sponge and crab occupy gastropod shells until the crab and sponge outgrow the shell, eventually resulting in the crab using the sponge's body as protection instead of the shell until the crab finds a suitable replacement shell.


Sponge loop

Most sponges are detritivores which filter organic debris particles and microscopic life forms from ocean water. In particular, sponges occupy an important role as detritivores in
coral reef food web Compared to terrestrial environments, marine environments have biomass pyramids which are inverted at the base. In particular, the biomass of consumers (copepods, krill, shrimp, forage fish) is larger than the biomass of primary producers. Thi ...
s by recycling detritus to higher
trophic level The trophic level of an organism is the position it occupies in a food web. A food chain is a succession of organisms that eat other organisms and may, in turn, be eaten themselves. The trophic level of an organism is the number of steps it ...
s. The hypothesis has been made that coral reef sponges facilitate the transfer of coral-derived organic matter to their associated detritivores via the production of sponge detritus, as shown in the diagram. Several sponge species are able to convert coral-derived DOM into sponge detritus, and transfer organic matter produced by corals further up the reef food web. Corals release organic matter as both dissolved and particulate mucus,Wild C, Huettel M, Klueter A, Kremb S, Rasheed M, Jorgensen B (2004) Coral mucus functions as an energy carrier and particle trap in the reef ecosystem. Nature 428: 66−70 as well as cellular material such as expelled ''
Symbiodinium : ''This is about the genus sometimes called Zoox. For the company, see Zoox (company)'' ''Symbiodinium'' is a genus of dinoflagellates that encompasses the largest and most prevalent group of endosymbiotic dinoflagellates known. These unicell ...
''.Hoegh-Guldberg O, McCloskey LR, Muscatine L (1987) Expulsion of zooxanthellae by symbiotic cnidarians from the Red Sea. Coral Reefs 5: 201−204 Organic matter could be transferred from corals to sponges by all these pathways, but DOM likely makes up the largest fraction, as the majority (56 to 80%) of coral mucus dissolves in the water column, and coral loss of fixed carbon due to expulsion of ''Symbiodinium'' is typically negligible (0.01%) compared with mucus release (up to ~40%). Coral-derived organic matter could also be indirectly transferred to sponges via bacteria, which can also consume coral mucus.


Sponge holobiont

Besides a one to one
symbiotic relationship Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasit ...
, it is possible for a host to become symbiotic with a
microbial consortium A microbial consortium or microbial community, is two or more bacterial or microbial groups living symbiotically. Consortiums can be endosymbiotic or ectosymbiotic, or occasionally may be both. The protist '' Mixotricha paradoxa'', itself an endosy ...
. Sponges are able to host a wide range of microbial communities that can also be very specific. The microbial communities that form a symbiotic relationship with the sponge can amount to as much as 35% of the biomass of its host. The term for this specific symbiotic relationship, where a microbial consortia pairs with a host is called a holobiotic relationship. The sponge as well as the microbial community associated with it will produce a large range of secondary metabolites that help protect it against predators through mechanisms such as
chemical defense Chemical defense is a life history strategy employed by many organisms to avoid consumption by producing toxic or repellent metabolites or chemical warnings which incite defensive behavioral changes. The production of defensive chemicals occurs in ...
. Some of these relationships include endosymbionts within bacteriocyte cells, and cyanobacteria or microalgae found below the pinacoderm cell layer where they are able to receive the highest amount of light, used for phototrophy. They can host over 50 different microbial phyla and candidate phyla, including Alphaprotoebacteria,
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 s ...
, Chloroflexota,
Nitrospirota Nitrospirota is a phylum of bacteria. It includes multiple genera, such as '' Nitrospira'', the largest. The first member of this phylum, '' Nitrospira marina'', was discovered in 1985. The second member, '' Nitrospira moscoviensis'', was discove ...
, " Cyanobacteria", the taxa Gamma-, the 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 Thaumarchaea.


Systematics and evolutionary history


Taxonomy

Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
, who classified most kinds of sessile animals as belonging to the order Zoophyta in the class
Vermes Vermes ("worms") is an obsolete taxon used by Carl Linnaeus and Jean-Baptiste Lamarck for non-arthropod invertebrate animals. Linnaeus In Linnaeus's ''Systema Naturae'', the Vermes had the rank of class, occupying the 6th (and last) slot of hi ...
, mistakenly identified the genus '' Spongia'' as plants in the order Algae. For a long time thereafter sponges were assigned to a separate subkingdom,
Parazoa Parazoa (Parazoa, gr. Παρα-, para, "next to", and ζωα, zoa, "animal") are a taxon with sub-kingdom category that is located at the base of the phylogenetic tree of the animal kingdom in opposition to the sub-kingdom Eumetazoa; they group ...
("beside the animals"), separate from the Eumetazoa which formed the rest of the
kingdom Kingdom commonly refers to: * A monarchy ruled by a king or queen * Kingdom (biology), a category in biological taxonomy Kingdom may also refer to: Arts and media Television * ''Kingdom'' (British TV series), a 2007 British television drama s ...
Animalia. They have been regarded as a paraphyletic phylum, from which the higher animals have evolved. Other research indicates Porifera is monophyletic. The phylum Porifera is further divided into classes mainly according to the composition of their skeletons: *
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 ...
(glass sponges) have silicate spicules, the largest of which have six rays and may be individual or fused. The main components of their bodies are
syncytia 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) ...
in which large numbers of cell share a single external
membrane A membrane is a selective barrier; it allows some things to pass through but stops others. Such things may be molecules, ions, or other small particles. Membranes can be generally classified into synthetic membranes and biological membranes. ...
. * Calcarea have skeletons made of calcite, a form of calcium carbonate, which may form separate spicules or large masses. All the cells have a single nucleus and membrane. * Most
Demospongiae Demosponges (Demospongiae) are the most diverse class in the phylum Porifera. They include 76.2% of all species of sponges with nearly 8,800 species worldwide (World Porifera Database). They are sponges with a soft body that covers a hard, ...
have silicate spicules or
spongin Spongin, a modified type of collagen protein, forms the fibrous skeleton of most organisms among the phylum Porifera, the sponges. It is secreted by sponge cells known as spongocytes. Spongin gives a sponge its flexibility. True spongin is found ...
fibers or both within their soft tissues. However, a few also have massive external skeletons made of
aragonite Aragonite is a carbonate mineral, one of the three most common naturally occurring crystal forms of calcium carbonate, (the other forms being the minerals calcite and vaterite). It is formed by biological and physical processes, including pre ...
, another form of calcium carbonate. All the cells have a single nucleus and membrane. * Archeocyatha are known only as fossils from the Cambrian period. In the 1970s, sponges with massive calcium carbonate skeletons were assigned to a separate class, Sclerospongiae, otherwise known as "coralline sponges". However, in the 1980s it was found that these were all members of either the Calcarea or the Demospongiae. So far scientific publications have identified about 9,000 poriferan species, of which: about 400 are glass sponges; about 500 are calcareous species; and the rest are demosponges. However, some types of habitat, vertical rock and cave walls and galleries in rock and coral boulders, have been investigated very little, even in shallow seas.


Classes

Sponges were traditionally distributed in three classes: calcareous sponges (Calcarea), glass sponges (Hexactinellida) and demosponges (Demospongiae). However, studies have shown that the
Homoscleromorpha Homosclerophorida is an order of marine sponges. It is the only order in the monotypic class Homoscleromorpha. The order is composed of two families: Plakinidae and Oscarellidae. Taxonomy Homoscleromorpha is phylogenetically well separated fr ...
, a group thought to belong to the
Demospongiae Demosponges (Demospongiae) are the most diverse class in the phylum Porifera. They include 76.2% of all species of sponges with nearly 8,800 species worldwide (World Porifera Database). They are sponges with a soft body that covers a hard, ...
, is actually phylogenetically well separated. Therefore, they have recently been recognized as the fourth class of sponges. Sponges are divided into classes mainly according to the composition of their skeletons: These are arranged in evolutionary order as shown below in ascending order of their evolution from top to bottom: :


Fossil record

Although
molecular clock The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleo ...
s and
biomarker In biomedical contexts, a biomarker, or biological marker, is a measurable indicator of some biological state or condition. Biomarkers are often measured and evaluated using blood, urine, or soft tissues to examine normal biological processes, p ...
s suggest sponges existed well before the Cambrian explosion of life,
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 ...
spicules like those of demosponges are absent from the fossil record until the Cambrian. An unsubstantiated 2002 report exists of spicules in rocks dated around . Well-preserved
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
sponges from about in the Ediacaran period have been found in the
Doushantuo Formation The Doushantuo Formation (formerly transcribed as Toushantuo or Toushantou, from ) is a geological formation in western Hubei, eastern Guizhou, southern Shaanxi, central Jiangxi, and other localities in China. It is known for the fossil Lagerst ...
. These fossils, which include: spicules; pinacocytes;
porocyte Porocytes are tubular cells which make up the pores of a sponge known as ostia. Description Covering the sponge is a layer of cells known as the pinacoderm, which is composed of pinacocytes. In a sponge, pinacocytes are a thin, elastic layer whic ...
s; archeocytes; sclerocytes; and the internal cavity, have been classified as demosponges. Fossils of
glass sponge 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 consider ...
s have been found from around in rocks in Australia, China, and Mongolia. Early Cambrian sponges from Mexico belonging to the genus ''Kiwetinokia'' show evidence of fusion of several smaller spicules to form a single large spicule. Calcium carbonate spicules of calcareous sponges have been found in Early Cambrian rocks from about in Australia. Other probable demosponges have been found in the Early Cambrian
Chengjiang fauna The Maotianshan Shales are a series of Early Cambrian deposits in the Chiungchussu Formation, famous for their '' Konservat Lagerstätten'', deposits known for the exceptional preservation of fossilized organisms or traces. The Maotianshan Shales ...
, from . Fossils found in the Canadian Northwest Territories dating to may be sponges; if this finding is confirmed, it suggests the first animals appeared before the Neoproterozoic oxygenation event. Freshwater sponges appear to be much younger, as the earliest known fossils date from the Mid-
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', " ...
period about . Although about 90% of modern sponges are
demosponges Demosponges (Demospongiae) are the most diverse class in the phylum Porifera. They include 76.2% of all species of sponges with nearly 8,800 species worldwide (World Porifera Database). They are sponges with a soft body that covers a hard, ...
, fossilized remains of this type are less common than those of other types because their skeletons are composed of relatively soft spongin that does not fossilize well. Earliest sponge symbionts are known from the early Silurian. A chemical tracer is 24-isopropyl cholestane, which is a stable derivative of 24-isopropyl
cholesterol Cholesterol is any of a class of certain organic molecules called lipids. It is a sterol (or modified steroid), a type of lipid. Cholesterol is biosynthesized by all animal cells and is an essential structural component of animal cell mem ...
, which is said to be produced by demosponges but not by eumetazoans ("true animals", i.e.
cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that ...
ns and bilaterians). Since choanoflagellates are thought to be animals' closest single-celled relatives, a team of scientists examined the
biochemistry Biochemistry or biological chemistry is the study of chemical processes within and relating to living organisms. A sub-discipline of both chemistry and biology, biochemistry may be divided into three fields: structural biology, enzymology and ...
and
gene In biology, the word gene (from , ; "... Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a b ...
s of one choanoflagellate species. They concluded that this species could not produce 24-isopropylcholesterol but that investigation of a wider range of choanoflagellates would be necessary in order to prove that the fossil 24-isopropylcholestane could only have been produced by demosponges. Although a previous publication reported traces of the chemical 24-isopropyl cholestane in ancient rocks dating to , recent research using a much more accurately dated rock series has revealed that these biomarkers only appear before the end of the
Marinoan glaciation The Marinoan glaciation, sometimes also known as the Varanger glaciation, was a period of worldwide glaciation that lasted from approximately 650 to 632.3 ± 5.9 Ma (million years ago) during the Cryogenian period. This glaciation possibly cover ...
approximately , and that "Biomarker analysis has yet to reveal any convincing evidence for ancient sponges pre-dating the first globally extensive Neoproterozoic glacial episode (the Sturtian, ~ in Oman)". While it has been argued that this 'sponge biomarker' could have originated from marine algae, recent research suggests that the algae's ability to produce this biomarker evolved only in the Carboniferous; as such, the biomarker remains strongly supportive of the presence of demosponges in the Cryogenian.
Archaeocyathid Archaeocyatha (or archaeocyathids 'ancient cups') is a taxon of extinct, sessile, reef-building marine sponges that lived in warm tropical and subtropical waters during the Cambrian Period. It is believed that the centre of the Archaeocyatha or ...
s, which some classify as a type of coralline sponge, are very common fossils in rocks from the Early Cambrian about , but apparently died out by the end of the Cambrian . It has been suggested that they were produced by: sponges;
cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that ...
ns; algae;
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 for catching food and other uses; and commonly ...
ns; a completely separate phylum of animals, Archaeocyatha; or even a completely separate
kingdom Kingdom commonly refers to: * A monarchy ruled by a king or queen * Kingdom (biology), a category in biological taxonomy Kingdom may also refer to: Arts and media Television * ''Kingdom'' (British TV series), a 2007 British television drama s ...
of life, labeled Archaeata or Inferibionta. Since the 1990s archaeocyathids have been regarded as a distinctive group of sponges. It is difficult to fit chancelloriids into classifications of sponges or more complex animals. An analysis in 1996 concluded that they were closely related to sponges on the grounds that the detailed structure of chancellorid sclerites ("armor plates") is similar to that of fibers of spongin, a collagen
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
, in modern keratose (horny) demosponges such as '' Darwinella''. However, another analysis in 2002 concluded that chancelloriids are not sponges and may be intermediate between sponges and more complex animals, among other reasons because their skins were thicker and more tightly connected than those of sponges. free text at In 2008 a detailed analysis of chancelloriids' sclerites concluded that they were very similar to those of
halkieriid The halkieriids are a group of fossil organisms from the Lower to Middle Cambrian. Their eponymous genus is ''Halkieria'' , which has been found on almost every continent in Lower to Mid Cambrian deposits, forming a large component of the smal ...
s, mobile
bilaterian The Bilateria or bilaterians are animals with bilateral symmetry as an embryo, i.e. having a left and a right side that are mirror images of each other. This also means they have a head and a tail (anterior-posterior axis) as well as a belly an ...
animals that looked like slugs in
chain mail Chain mail (properly called mail or maille but usually called chain mail or chainmail) is a type of armour consisting of small metal rings linked together in a pattern to form a mesh. It was in common military use between the 3rd century BC and ...
and whose fossils are found in rocks from the very Early Cambrian to the Mid Cambrian. If this is correct, it would create a dilemma, as it is extremely unlikely that totally unrelated organisms could have developed such similar sclerites independently, but the huge difference in the structures of their bodies makes it hard to see how they could be closely related.


Relationships to other animal groups

In the 1990s sponges were widely regarded as a monophyletic group, all of them having descended from a
common ancestor Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. All living beings are in fact descendants of a unique ancestor commonly referred to as the last universal comm ...
that was itself a sponge, and as the "sister-group" to all other
metazoans 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 ...
(multi-celled animals), which themselves form a monophyletic group. On the other hand, some 1990s analyses also revived the idea that animals' nearest evolutionary relatives are choanoflagellates, single-celled organisms very similar to sponges'
choanocytes Choanocytes (also known as "collar cells") are cells that line the interior of asconoid, syconoid and leuconoid body types of sponges that contain a central flagellum, or ''cilium,'' surrounded by a collar of microvilli which are connected by a ...
– which would imply that most Metazoa evolved from very sponge-like ancestors and therefore that sponges may not be monophyletic, as the same sponge-like ancestors may have given rise both to modern sponges and to non-sponge members of Metazoa. Analyses since 2001 have concluded that Eumetazoa (more complex than sponges) are more closely related to particular groups of sponges than to other sponge groups. Such conclusions imply that sponges are not monophyletic, because the
last common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...
of all sponges would also be a direct ancestor of the Eumetazoa, which are not sponges. A study in 2001 based on comparisons of ribosome DNA concluded that the most fundamental division within sponges was between
glass sponge 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 consider ...
s and the rest, and that Eumetazoa are more closely related to calcareous sponges (those with calcium carbonate spicules) than to other types of sponge. In 2007 one analysis based on comparisons of RNA and another based mainly on comparison of spicules concluded that demosponges and glass sponges are more closely related to each other than either is to the calcareous sponges, which in turn are more closely related to Eumetazoa. Other anatomical and biochemical evidence links the Eumetazoa with
Homoscleromorpha Homosclerophorida is an order of marine sponges. It is the only order in the monotypic class Homoscleromorpha. The order is composed of two families: Plakinidae and Oscarellidae. Taxonomy Homoscleromorpha is phylogenetically well separated fr ...
, a sub-group of demosponges. A comparison in 2007 of nuclear DNA, excluding glass sponges and
comb jellies Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), and ...
, concluded that: *
Homoscleromorpha Homosclerophorida is an order of marine sponges. It is the only order in the monotypic class Homoscleromorpha. The order is composed of two families: Plakinidae and Oscarellidae. Taxonomy Homoscleromorpha is phylogenetically well separated fr ...
are most closely related to Eumetazoa; * calcareous sponges are the next closest; * the other demosponges are evolutionary "aunts" of these groups; and * the chancelloriids, bag-like animals whose fossils are found in Cambrian rocks, may be sponges. The sperm of Homoscleromorpha share features with the sperm of Eumetazoa, that sperm of other sponges lack. In both Homoscleromorpha and Eumetazoa layers of cells are bound together by attachment to a carpet-like basal membrane composed mainly of "typ IV" collagen, a form of collagen not found in other sponges – although the
spongin Spongin, a modified type of collagen protein, forms the fibrous skeleton of most organisms among the phylum Porifera, the sponges. It is secreted by sponge cells known as spongocytes. Spongin gives a sponge its flexibility. True spongin is found ...
fibers that reinforce the mesohyl of all demosponges is similar to "type IV" collagen. The analyses described above concluded that sponges are closest to the ancestors of all Metazoa, of all multi-celled animals including both sponges and more complex groups. However, another comparison in 2008 of 150 genes in each of 21 genera, ranging from fungi to humans but including only two species of sponge, suggested that
comb jellies Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), and ...
(
ctenophora Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), and ...
) are the most basal lineage of the Metazoa included in the sample. If this is correct, either modern comb jellies developed their complex structures independently of other Metazoa, or sponges' ancestors were more complex and all known sponges are drastically simplified forms. The study recommended further analyses using a wider range of sponges and other simple Metazoa such as
Placozoa The Placozoa are a basal form of marine free-living (non-parasitic) multicellular organism. They are the simplest in structure of all animals. Three genera have been found: the classical ''Trichoplax adhaerens'', '' Hoilungia hongkongensis'', a ...
. The results of such an analysis, published in 2009, suggest that a return to the previous view may be warranted. 'Family trees' constructed using a combination of all available data – morphological, developmental and molecular – concluded that the sponges are in fact a monophyletic group, and with the cnidarians form the sister group to the bilaterians. A very large and internally consistent alignment of 1,719 proteins at the metazoan scale, published in 2017, showed that (i) sponges – represented by Homoscleromorpha, Calcarea, Hexactinellida, and Demospongiae – are monophyletic, (ii) sponges are sister-group to all other multicellular animals, (iii) ctenophores emerge as the second-earliest branching animal lineage, and (iv)  placozoans emerge as the third animal lineage, followed by cnidarians sister-group to bilaterians. In March 2021, scientists from Dublin found additional evidence that sponges are the sister group to all other animals.


Notable spongiologists

* Céline Allewaert * Patricia Bergquist *
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 ...
*
Maurice Burton Maurice Burton (28 March 1898 – 9 September 1992) was a British zoologist and popular science author, who produced many natural history encyclopedias and books including a skeptical treatment of the Loch Ness Monster. Early life and educatio ...
*
Henry John Carter Henry John Carter, FRS (18 August 1813 – 4 May 1895) was a surgeon working in Bombay, India, who carried out work in geology, paleontology, and zoology. He worked as an army surgeon in Bombay from 1859 on Her Majesty's Indian Service, Bom ...
*
Max Walker de Laubenfels Max Walker de Laubenfels (1894–1960) was an American spongiologist. He received his undergraduate degree from Oberlin College and his doctorate from Stanford University. He was among the most prolific identifiers of new species of Caribbean ...
*
Arthur Dendy Arthur Dendy (20 January 1865, in Manchester – 24 March 1925, in London) was an English zoologist known for his work on marine sponges and the terrestrial invertebrates of Victoria, Australia, notably including the "living fossil" '' Peripatus'' ...
* Édouard Placide Duchassaing de Fontbressin *
Randolph Kirkpatrick Randolph Kirkpatrick (1863 – 1950) was a British spongiologist, cnidariologist and bryozoologist. He was assistant keeper of lower invertebrates at the British Natural History Museum from 1886 until his retirement in 1927. Kirkpatrick ...
*
Robert J. Lendlmayer von Lendenfeld Robert J. Lendlmayer von Lendenfeld (1858-1913) was an Austrian people, Austrian zoologist, alpinist, and traveler. He was also a notable Spongiology, spongiologist. von Lendenfeld was one of a number of influential German-speaking residents suc ...
* Edward Alfred Minchin * Giovanni Domenico Nardo *
Eduard Oscar Schmidt Eduard Oscar Schmidt (21 February 1823, in Torgau – 17 January 1886, in Kappelrodeck) was a German zoologist and phycologist. Biography He initially studied mathematics and science at Halle, then continued his education in Berlin, where he c ...
* Émile Topsent


Use


By dolphins

A report in 1997 described use of sponges as a tool by
bottlenose dolphins Bottlenose dolphins are aquatic mammals in the genus ''Tursiops.'' They are common, cosmopolitan members of the family Delphinidae, the family of oceanic dolphins. Molecular studies show the genus definitively contains two species: the comm ...
in Shark Bay in Western Australia. A dolphin will attach a marine sponge to its
rostrum Rostrum may refer to: * Any kind of a platform for a speaker: **dais **pulpit * Rostrum (anatomy), a beak, or anatomical structure resembling a beak, as in the mouthparts of many sucking insects * Rostrum (ship), a form of bow on naval ships * Ros ...
, which is presumably then used to protect it when searching for food in the sandy sea bottom. The behavior, known as ''sponging'', has only been observed in this bay, and is almost exclusively shown by females. A study in 2005 concluded that mothers teach the behavior to their daughters, and that all the sponge-users are closely related, suggesting that it is a fairly recent innovation.


By humans


Skeleton

The calcium carbonate or
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 ...
spicules of most sponge genera make them too rough for most uses, but two genera, '' Hippospongia'' and '' Spongia'', have soft, entirely fibrous skeletons. Early Europeans used soft sponges for many purposes, including padding for helmets, portable drinking utensils and municipal water filters. Until the invention of synthetic sponges, they were used as cleaning tools, applicators for paints and
ceramic glaze Ceramic glaze is an impervious layer or coating of a vitreous substance which has been fused to a pottery body through firing. Glaze can serve to color, decorate or waterproof an item. Glazing renders earthenware vessels suitable for holding ...
s and discreet contraceptives. However, by the mid-20th century, over-fishing brought both the animals and the industry close to extinction. Many objects with sponge-like textures are now made of substances not derived from poriferans. Synthetic sponges include personal and household
cleaning tools Cleaning tools include the following: *Acoustic cleaning *Air blaster * Air knife *Besom *Broom *Brush * Building maintenance unit *Camel-hair brush *Carbon dioxide cleaning *Carpet beater * Carpet sweeper *Chamois leather * Cleret * Cyclone dust c ...
, breast implants, and
contraceptive sponge The contraceptive sponge combines barrier and spermicidal methods to prevent conception. Sponges work in two ways. First, the sponge is inserted into the vagina, so it can cover the cervix and prevent any sperm from entering the uterus. Secondly ...
s. Typical materials used are
cellulose Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell w ...
foam,
polyurethane Polyurethane (; often abbreviated PUR and PU) refers to a class of polymers composed of organic units joined by carbamate (urethane) links. In contrast to other common polymers such as polyethylene and polystyrene, polyurethane is produced from ...
foam, and less frequently, silicone foam. The
luffa ''Luffa'' is a genus of tropical and subtropical vines in the cucumber family (Cucurbitaceae). In everyday non-technical usage, the luffa, also spelled loofah, usually refers to the fruits of the species '' Luffa aegyptiaca'' and '' Luffa acuta ...
"sponge", also spelled ''loofah'', which is commonly sold for use in the kitchen or the shower, is not derived from an animal but mainly from the fibrous "skeleton" of the sponge gourd (''Luffa aegyptiaca'',
Cucurbitaceae The Cucurbitaceae, also called cucurbits or the gourd family, are a plant family consisting of about 965 species in around 95 genera, of which the most important to humans are: *''Cucurbita'' – squash, pumpkin, zucchini, some gourds *'' Lagen ...
).


Antibiotic compounds

Sponges have
medicinal Medicine is the science and practice of caring for a patient, managing the diagnosis, prognosis, prevention, treatment, palliation of their injury or disease, and promoting their health. Medicine encompasses a variety of health care practic ...
potential due to the presence in sponges themselves or their microbial symbionts of chemicals that may be used to control
virus A virus is a submicroscopic infectious agent that replicates only inside the living cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea. Since Dmitri Ivanovsk ...
es,
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
,
tumor A neoplasm () is a type of abnormal and excessive growth of tissue. The process that occurs to form or produce a neoplasm is called neoplasia. The growth of a neoplasm is uncoordinated with that of the normal surrounding tissue, and persists ...
s and fungi.


Other biologically active compounds

Lacking any protective shell or means of escape, sponges have evolved to synthesize a variety of unusual compounds. One such class is the oxidized fatty acid derivatives called oxylipins. Members of this family have been found to have anti-cancer, anti-bacterial and anti-fungal properties. One example isolated from the Okinawan ''plakortis'' sponges,
plakoridine A Plakoridine A is an alkaloid Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Som ...
, has shown potential as a cytotoxin to murine lymphoma cells.


See also

*
Lists of sponges The following are lists of 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 channe ...
*
Sponge Reef Project The Sponge Reef Project is a binational scientific project between Germany and Canada to study the sponge reefs off British Columbia, Canada, reefs formed by sponges of the Hexactinellid family. The project was started in 1999, following the dis ...
*
SpongeBob SquarePants ''SpongeBob SquarePants'' (or simply ''SpongeBob'') is an American animated comedy television series created by marine science educator and animator Stephen Hillenburg for Nickelodeon. It chronicles the adventures of the title character ...
* 3-Alkylpyridinium, compounds found in marine
Haplosclerida Haplosclerida is an order of demosponge Demosponges (Demospongiae) are the most diverse class in the phylum Porifera. They include 76.2% of all species of sponges with nearly 8,800 species worldwide (World Porifera Database). They are ...
sponges


References


Bibliography

* * * * * * * *


External links


Water flow and feeding in the phylum Porifera (sponges)
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animations of sponge body structures, water flow and feeding
Carsten's Spongepage
Information on the ecology and the biotechnological potential of sponges and their associated bacteria.

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Nature's 'fibre optics' experts



Queensland Museum information about sponges

Queensland Museum Sessile marine invertebrates collections

Queensland Museum Sessile marine invertebrates research

Sponge Guide for Britain and Ireland
Bernard Picton, Christine Morrow & Rob van Soest
World Porifera database
the world list of extant sponges, includes a searchable database.

// Food and Agriculture Organisation {{Authority control Aquatic animals Ediacaran first appearances Parazoa