SPONGE

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This article is about the phylum of aquatic animal. For the porous cleaning tool, see Sponge
(tool). For other uses, see Sponge (disambiguation).
Porifera
Temporal range: Ediacaran–recent
PreꞒ
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O
S
D
C
P
T
J
K
Pg
N

A stove-pipe sponge
Scientific classification
Kingdom: Animalia
Phylum: Porifera
Grant, 1836
Classes
 Calcarea
 Hexactinellida
 Demospongiae
 Homoscleromorpha
 †Stromatoporoidea
  †Archaeocyatha

Synonyms
Parazoa/Ahistozoa (sans
Placozoa)[1]

Sponges, the members of the phylum Porifera (/pəˈrɪfərə/; meaning 'pore bearer'), are a basal
animal clade as a sister of the Diploblasts.[2][3][4][5][6] They are multicellular organisms that
have bodies full of pores and channels allowing water to circulate through them, consisting of
jelly-like mesohyl 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 systems. 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 from the last common ancestor of all animals, making them
the sister group of all other animals.[2]

Overview

Sponge biodiversity and morphotypes at the lip of a wall site in 60 feet (20 m) of water.
Included are the yellow tube sponge, Aplysina fistularis, the purple vase sponge, Niphates
digitalis, the red encrusting sponge, Spirastrella coccinea, and the gray rope sponge,
Callyspongia sp.

Sponges are similar to other animals in that they are multicellular, heterotrophic, lack cell
walls and produce sperm cells. Unlike other animals, they lack true tissues[8] and organs.[9]
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. Many sponges
have internal skeletons of spongin and/or spicules (skeletal-like fragments) of calcium
carbonate or silicon dioxide.[8] All sponges are sessile aquatic animals, meaning that they
attach to an underwater surface and remain fixed in place (i.e., do not travel). Although there
are freshwater species, the great majority are marine (salt-water) species, ranging in habitat
from tidal zones to depths exceeding 8,800 m (5.5 mi).

Although most of the approximately 5,000–10,000 known species of sponges feed on bacteria
and other microscopic food in the water, some host photosynthesizing microorganisms as
endosymbionts, 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 carnivores that
prey mainly on small crustaceans.[10]

Most species use sexual reproduction, releasing sperm cells into the water to fertilize ova that
in some species are released and in others are retained by the "mother". The fertilized eggs
develop into larvae, which swim off in search of places to settle. [11] 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 gemmules, "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. [12]

In most sponges, an internal gelatinous matrix called mesohyl functions as an endoskeleton,

and it is the only skeleton in soft sponges that encrust such hard surfaces as rocks. More
commonly, the mesohyl is stiffened by mineral spicules, by spongin fibers, or both.
Demosponges use spongin; many species have silica spicules, whereas some species have
calcium carbonate exoskeletons. Demosponges 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.[13] The fragile glass sponges, with "scaffolding" 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 580 million years ago. In addition
Archaeocyathids, whose fossils are common in rocks from 530 to 490 million years ago, are
now regarded as a type of sponge.

Cells of the protist choanoflagellate clade closely resemble sponge choanocyte cells. Beating
of choanocyte flagella draws water through the sponge so that nutrients can be extracted and
waste removed.[14]

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. [15] 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
rather than sponges are the sister group to the rest of animals. [16][17][18][19] 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.[20][21]

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 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. Dolphins have been observed using sponges
as tools while foraging.[22]

Distinguishing features
Further information: Cnidaria and Ctenophore

Sponges constitute the phylum Porifera, and have been defined as sessile metazoans
(multicelled immobile animals) that have water intake and outlet openings connected by
chambers lined with choanocytes, cells with whip-like flagella.[23] However, a few
carnivorous sponges have lost these water flow systems and the choanocytes. [24][25] 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.[25][26]

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. [27]

Like cnidarians (jellyfish, etc.) and ctenophores (comb jellies), and unlike all other known
metazoans, sponges' bodies consist of a non-living jelly- like mass (mesohyl) sandwiched
between two main layers of cells. [28][29] 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").[29] 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.[26]

Cnidarians and
Sponges [26][28]
ctenophores [29]
Nervous system No Yes, simple
No, except that Yes: inter-cell
Cells in each layer bound
Homoscleromorpha have basement connections; basement
together
membranes.[30] membranes
Number of cells in middle
Many Few
"jelly" layer
 Cells in outer layers can
move inwards and change Yes No
functions

Basic structure
Cell types

Mesohyl
Pinacocyte
Choanocyte
Lophocyte
Porocyte
Oocyte
Archeocyte
Sclerocyte
Spicule
Water flow

Main cell types of Porifera[31]

A sponge's body is hollow and is held in shape by the mesohyl, 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 choanocytes, cells with cylindrical or conical collars
surrounding one flagellum 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
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, [26][28] while those at
the base of the animal are responsible for anchoring it. [28]

Other types of cell live and move within the mesohyl:[26][28]

 Lophocytes are amoeba-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.
 Oocytes and spermatocytes 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 polymerizes into spongin, a thick fibrous material that stiffens
the mesohyl.
 Myocytes ("muscle cells") conduct signals and cause parts of the animal to contract.
 "Grey cells" act as sponges' equivalent of an immune system.
 Archaeocytes (or amoebocytes) are amoeba-like cells that are totipotent, 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 cryptochromes. They
mediate phototaxic behavior.[32]

Glass sponges' syncytia

Water flow
Main syncitium
Spicules
Choanosyncitium
and collar bodies
showing interior
The glass sponge Euplectella[33]

Glass sponges present a distinctive variation on this basic plan. Their spicules, which are
made of silica, form a scaffolding- like framework between whose rods the living tissue is
suspended like a cobweb that contains most of the cell types.[26] This tissue is a syncytium
that in some ways behaves like many cells that share a single external membrane, and in
others like a single cell with multiple nuclei. The mesohyl is absent or minimal. The
syncytium's cytoplasm, the soupy fluid that fills the interiors of cells, is organized into
"rivers" that transport nuclei, organelles ("organs" within cells) and other substances. [34]
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.[26]

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. 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. [34]

Water flow and body structures

Asconoid
Syconoid
Leuconoid
Pinacocytes
Choanocytes
Mesohyl
Water flow
Porifera body structures[35]

Most sponges work rather like chimneys: they take in water at the bottom and eject it from
the osculum ("little mouth") at the top. Since ambient currents are faster at the top, the
suction effect that they produce by Bernoulli's principle 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. [26]

Although the layers of pinacocytes and choanocytes resemble the epithelia 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. [36]

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 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
1 mm (0.039 in) in diameter.[26]

Diagram of a syconoid sponge

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 1 m (3.3 ft) 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 sponges are similar to the leuconid structure. [26] 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.[26] For example, in Leuconia, a small leuconoid
sponge about 10 centimetres (3.9 in) tall and 1 centimetre (0.39 in) 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 at about
8.5 cm per second, fast enough to carry waste products some distance away. [37]

P inacocyte
Choanocyte
Archeocytes and other cells in
mesohyl
Mesohyl
Spicules
Calcium carbonate
Seabed / rock
Water flow
Sponge with calcium carbonate skeleton[26]

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 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,[38] may be
made of silica 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,[26] and may be separate,
connected by joints, or fused.[25]

Some sponges also secrete exoskeletons 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 chambers in pits in the
mineral. These exoskeletons are secreted by the pinacocytes that form the animals' skins.[26]

Vital functions

Spongia officinalis, "the kitchen sponge", is dark grey when alive.

Movement

Although adult sponges are fundamentally sessile animals, some marine and freshwater
species can move across the sea bed at speeds of 1–4 mm (0.039–0.157 in) per day, as a
result of amoeba-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.[26]

Respiration, feeding and excretion

Euplectella aspergillum, a glass sponge known as "Venus' flower basket"

Sponges do not have distinct circulatory, respiratory, digestive, and excretory systems –
instead the water flow system supports all these functions. They filter food particles out of the
water flowing through them. Particles larger than 50 micrometers cannot enter the ostia and
pinacocytes consume them by phagocytosis (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 archaeocytes 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 choanocytes.[26] Since the smallest
particles are by far the most common, choanocytes typically capture 80% of a sponge's food
supply.[39] 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,[26] and these tracks also move inert
objects.[28]

It used to be claimed that glass sponges could live on nutrients dissolved in sea water and
were very averse to silt.[40] 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. [41] 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.[26]

Sponges' cells absorb oxygen by diffusion from water into cells as water flows through body,
into which carbon dioxide and other soluble waste products such as ammonia 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. [26]

Carnivorous sponges

The carnivorous ping-pong tree sponge, Chondrocladia lampadiglobus [42]

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. [43] Most belong to the
family Cladorhizidae, but a few members of the Guitarridae and Esperiopsidae are also
carnivores.[44] 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.[44][45] Most
carnivorous sponges live in deep waters, up to 8,840 m (5.49 mi),[46] and the development of
deep-ocean exploration techniques is expected to lead to the discovery of several more. [26][44]
However, one species has been found in Mediterranean caves at depths of 17–23 m (56–
75 ft), alongside the more usual filter feeding sponges. The cave-dwelling predators capture
crustaceans under 1 mm (0.039 in) 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.[46]

Most known carnivorous sponges have completely lost the water flow system and
choanocytes. However, the genus Chondrocladia uses a highly modified water flow system
to inflate balloon-like structures that are used for capturing prey. [44][47]

Endosymbionts

Freshwater sponges often host green algae as endosymbionts within archaeocytes and other
cells, and benefit from nutrients produced by the algae. Many marine species host other
photosynthesizing 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.[26] In 2008 a University of Stuttgart team reported that spicules made of silica
conduct light into the mesohyl, where the photosynthesizing endosymbionts live. [48] 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.[28]

A recently discovered carnivorous sponge that lives near hydrothermal vents hosts methane-
eating bacteria, and digests some of them. [28]

"Immune" system

Sponges do not have the complex immune systems 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. [28]

Reproduction

Asexual
The freshwater sponge Spongilla lacustris

Sponges have three asexual methods of reproduction: after fragmentation; by budding; and by
producing gemmules. Fragments of sponges may be detached by currents or waves. They use
the mobility of their pinacocytes and choanocytes and reshaping of the mesohyl 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. [49] A sponge fragment can only regenerate if it
contains both collencytes to produce mesohyl and archeocytes to produce all the other cell
types.[39] A very few species reproduce by budding. [50]

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.[51] Freshwater
gemmules may also include photosynthesizing symbionts. [52] The gemmules then become
dormant, and in this state can survive cold, drying out, lack of oxygen and extreme variations
in salinity.[26] Freshwater gemmules often do not revive until the temperature drops, stays
cold for a few months and then reaches a near-"normal" level.[52] 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.[53] 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.[52]

Sexual

Most sponges are hermaphrodites (function as both sexes simultaneously), although sponges
have no gonads (reproductive organs). Sperm are produced by choanocytes or entire
choanocyte chambers that sink into the mesohyl and form spermatic cysts while eggs are
formed by transformation of archeocytes, or of choanocytes in some species. Each egg
generally acquires a yolk by consuming "nurse cells". During spawning, sperm burst out of
their cysts and are expelled via the osculum. 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 form and carry the sperm through the mesohyl to eggs,
which in most cases engulf the carrier and its cargo. [54]

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. [54]

Glass sponge 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.[55]

Life cycle

Sponges in temperate regions live for at most a few years, but some tropical species and
perhaps some deep-ocean ones may live for 200 years or more. Some calcified demosponges
grow by only 0.2 mm (0.0079 in) per year and, if that rate is constant, specimens 1 m (3.3 ft)
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. [26]

Coordination of activities

Adult sponges lack neurons or any other kind of nervous tissue. 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 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.[56] However, glass sponges rapidly
transmit electrical impulses through all parts of the syncytium, and use this to halt the motion
of their flagella if the incoming water contains toxins or excessive sediment.[26] Myocytes are
thought to be responsible for closing the osculum and for transmitting signals between
different parts of the body.[28]

Sponges contain genes 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. [57]

Ecology
Habitats

See also: sponge ground and sponge reef

Sponges are worldwide in their distribution, living in a wide range of ocean habitats, from the
polar regions to the tropics.[39] 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. [40]
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.[58]

Euplectella aspergillum is a deep ocean glass sponge; seen here at a depth of 2,572 metres
(8,438 ft) off the coast of California

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. [59]
Glass sponges 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.[60]

The different classes of sponge live in different ranges of habitat:

Type of
Water type [28] Depth[28]
surface [28]
Calcarea Marine less than 100 m (330 ft) Hard
Glass Soft or firm
Marine Deep
sponges sediment
Marine, brackish; and Inter-tidal to abyssal;[28] a
Demosponges about 150 freshwater carnivorous demosponge has been Any
species[26] found at 8,840 m (5.49 mi)[46]

As primary producers

Sponges with photosynthesizing endosymbionts produce up to three times more oxygen 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 but relatively minor in
the Caribbean.[39]

Defenses
Holes made by clionaid sponge (producing the trace Entobia) after the death of a modern
bivalve shell of species Mercenaria mercenaria, from North Carolina

Close-up of the sponge boring Entobia in a modern oyster valve. Note the chambers which
are connected by short tunnels.

Many sponges shed spicules, forming a dense carpet several meters deep that keeps away
echinoderms which would otherwise prey on the sponges. [39] They also produce toxins that
prevent other sessile organisms such as bryozoans or sea squirts from growing on or near
them, making sponges very effective competitors for living space. One of many examples
includes ageliferin.

A few species, the Caribbean fire sponge Tedania ignis, cause a severe rash in humans who
handle them.[26] Turtles and some fish feed mainly on sponges. It is often said that sponges
produce chemical defenses against such predators.[26] 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. [28]
However, some studies have shown fish showing a preference for non chemically defended
sponges,[61] and another study found that high levels of coral predation did predict the
presence of chemically defended species. [62]

Glass sponges produce no toxic chemicals, and live in very deep water where predators are
rare.[40]

Predation

See also: Spongivore

Sponge flies, also known as spongilla- flies (Neuroptera, Sisyridae), 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. [63]

Bioerosion

The Caribbean chicken-liver sponge Chondrilla nucula secretes toxins that kill coral polyps,
allowing the sponges to grow over the coral skeletons. [26] Others, especially in the family
Clionaidae, use corrosive substances secreted by their archeocytes to tunnel into rocks, corals
and the shells of dead mollusks.[26] Sponges may remove up to 1 m (3.3 ft) per year from
reefs, creating visible notches just below low-tide level.[39]

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 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.[64] The rust-colored bands are caused by a cyanobacterium, but it is unknown
whether this organism actually causes the disease. [64][65]

Collaboration with other organisms

In addition to hosting photosynthesizing endosymbionts, [26] 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 meadows 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.[66]

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. [67] As many as 16,000 individuals inhabit a single
loggerhead sponge, feeding off the larger particles that collect on the sponge as it filters the
ocean to feed itself.[68] Other crustaceans such as hermit crabs commonly have a specific
species of sponge, Pseudospongosorites, 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.[69]
Bathymetrical range of some sponge species [70]
Demosponge Samus anonymus (up to 50 m), hexactinellid Scleroplegma lanterna (~100–600
m), hexactinellid Aulocalyx irregularis (~550–915 m), lithistid demosponge Neoaulaxinia
persicum (~500–1,700 m)
Generalised food web for sponge reefs [71]

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 webs by recycling detritus to higher trophic levels.[72]

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,[73][74] and transfer organic matter produced by corals further up the reef food web.
Corals release organic matter as both dissolved and particulate mucus, [75][76][77][78] as well as
cellular material such as expelled Symbiodinium.[79][80][72]

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,[76] and coral loss of fixed carbon due to expulsion of Symbiodinium is
typically negligible (0.01%)[79] compared with mucus release (up to ~40%).[81][82] Coral-
derived organic matter could also be indirectly transferred to sponges via bacteria, which can
also consume coral mucus.[83][84][85][72]
Sponge loop hypothesis
Steps of the sponge loop pathway: (1) corals and algae release exudates as dissolved organic
matter (DOM), (2) sponges take up DOM, (3) sponges release detrital particulate organic
matter (POM), (4) sponge detritus (POM) is taken up by sponge-associated and free-living
detritivores.[72][86][87]

The sponge holobiont

The sponge holobiont is an example of the concept of nested ecosystems. Key functions
carried out by the microbiome (colored arrows) influence holobiont functioning and, through
cascading effects, subsequently influence community structure and ecosystem functioning.
Environmental factors act at multiple scales to alter microbiome, holobiont, community, and
ecosystem scale processes. Thus, factors that alter microbiome functioning can lead to
changes at the holobiont, community, or even ecosystem level and vice versa, illustrating the
necessity of considering multiple scales when evaluating functioning in nested ecosystems.[88]
DOM: dissolved organic matter; POM: particulate organic matter
DIN: dissolved inorganic nitrogen
Sponge holobiont

Besides a one to one symbiotic relationship, it is possible for a host to become symbiotic with
a microbial consortia. 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.[89] 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.[90]

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, Actinobacteria,
Chloroflexi, Nitrospirae, Cyanobacteria, the taxa Gamma-, the candidate phylum
Poribacteria, and Thaumarchaea.[90]

Systematics and evolutionary history

Taxonomy

Linnaeus, who classified most kinds of sessile animals as belonging to the order Zoophyta in
the class Vermes, mistakenly identified the genus Spongia as plants in the order Algae.[91] For
a long time thereafter sponges were assigned to a separate subkingdom, Parazoa ("beside the
animals"), separate from the Eumetazoa which formed the rest of the kingdom Animalia.[92]
They have been regarded as a paraphyletic phylum, from which the higher animals have
evolved.[93] Other research indicates Porifera is monophyletic. [94]

The phylum Porifera is further divided into classes mainly according to the composition of
their skeletons:[25][39]

 Hexactinellida (glass sponges) have silicate spicules, the largest of which have six
rays and may be individual or fused. [25] The main components of their bodies are
syncytia in which large numbers of cell share a single external membrane.[39]
 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.[39]
 Most Demospongiae have silicate spicules or spongin fibers or both within their soft
tissues. However, a few also have massive external skeletons made of aragonite,
another form of calcium carbonate. [25][39] All the cells have a single nucleus and
membrane.[39]
 Archeocyatha are known only as fossils from the Cambrian period.[92]

In the 1970s, sponges with massive calcium carbonate skeletons were assigned to a separate
class, Sclerospongiae, otherwise known as "coralline sponges". [95] However, in the 1980s it
was found that these were all members of either the Calcarea or the Demospongiae. [96]
So far scientific publications have identified about 9,000 poriferan species, [39] of which: about
400 are glass sponges; about 500 are calcareous species; and the rest are demosponges.[26]
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.[39]

Classes

Sponges were traditionally distributed in three classes: calcareous sponges (Calcarea), glass
sponges (Hexactinellida) and demosponges (Demospongiae). However, studies have shown
that the Homoscleromorpha, a group thought to belong to the Demospongiae, is actually
phylogenetically well separated.[97] Therefore, they have recently been recognized as the
fourth class of sponges.[98][99]

Sponges are divided into classes mainly according to the composition of their skeletons:[28]
These are arranged in evolutionary order as shown below in ascending order of their
evolution from top to bottom:

Type of Spongin Massive

Spicules [28] Body form[28]
cells [28] fibers [28] exoskeleton[39]
Silica
Mostly
May be
Hexactinellida syncytia in Never Never Leuconoid
individual or
all species
fused
Single
In some species.
nucleus,
In many Made of
Demospongiae single Silica Leuconoid
species aragonite if
external
present.[25][39]
membrane
Single
Calcite Asconoid,
nucleus, Common.
May be syconoid,
Calcarea single Never Made of calcite
individual or leuconoid or
external if present.
large masses solenoid[100]
membrane
Single
nucleus,
In many Sylleibid or
Homoscleromorpha single Silica Never
species leuconoid
external
membrane

Fossil record

"Primitive Sponge" redirects here. Not to be confused with Sponge (material) § History.
Raphidonema faringdonense, a fossil sponge from the Cretaceous of England

1
2
3
4
5
6
7
1: Gap 2: Central cavity 3 Internal wall 4: Pore (all walls have pores) 5 Septum 6 Outer
wall 7 Holdfast

Archaeocyathid structure

Although molecular clocks and biomarkers suggest sponges existed well before the Cambrian
explosion of life, silica spicules like those of demosponges are absent from the fossil record
until the Cambrian.[101] An unsubstantiated 2002 report exists of spicules in rocks dated
around 750 million years ago.[102] Well-preserved fossil sponges from about 580 million
years ago in the Ediacaran period have been found in the Doushantuo Formation. These
fossils, which include spicules, pinacocytes, porocytes, archeocytes, sclerocytes and the
internal cavity, have been classified as demosponges. Fossils of glass sponges have been
found from around 540 million years ago in rocks in Australia, China and Mongolia. [103]
Early Cambrian sponges from Mexico belonging to the genus Kiwetinokia show evidence of
fusion of several smaller spicules to form a single large spicule.[104] Calcium carbonate
spicules of calcareous sponges have been found in Early Cambrian rocks from about
530 to 523 million years ago in Australia. Other probable demosponges have been found in
the Early Cambrian Chengjiang fauna, from 525 to 520 million years ago.[105] Fossils found
in the Canadian Northwest Territories dating to 890 million years ago may be sponges; if this
finding is confirmed, it suggests the first animals appeared before the Neoproterozoic
oxygenation event.[106]

Oxygen content of the atmosphere over the last billion years. If confirmed, the discovery of
fossilized sponges dating to 890 million years ago would predate the Neoproterozoic
Oxygenation Event (800-500 million years BP).

Freshwater sponges appear to be much younger, as the earliest known fossils date from the
Mid-Eocene period about 48 to 40 million years ago.[103] Although about 90% of modern
sponges are demosponges, 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.[107] Earliest sponge symbionts are known from the early Silurian.[108]

A chemical tracer is 24-isopropylcholestane, which is a stable derivative of 24-

isopropylcholesterol, which is said to be produced by demosponges but not by eumetazoans
("true animals", i.e. cnidarians and bilaterians). Since choanoflagellates are thought to be
animals' closest single-celled relatives, a team of scientists examined the biochemistry and
genes 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.[109] Although a previous publication reported traces of the
chemical 24-isopropylcholestane in ancient rocks dating to 1,800 million years ago,[110]
recent research using a much more accurately dated rock series has revealed that these
biomarkers only appear before the end of the Marinoan glaciation approximately 635 million
years ago,[111] 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, ~713 million years ago 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. [112][113][114]

Archaeocyathids, which some classify as a type of coralline sponge, are very common fossils
in rocks from the Early Cambrian about 530 to 520 million years ago, but apparently died out
by the end of the Cambrian 490 million years ago.[105] It has been suggested that they were
produced by: sponges; cnidarians; algae; foraminiferans; a completely separate phylum of
animals, Archaeocyatha; or even a completely separate kingdom of life, labeled Archaeata or
Inferibionta. Since the 1990s archaeocyathids have been regarded as a distinctive group of
sponges.[92]

= skin
= aragonite
= flesh

Halkieriid sclerite structure[115]

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, in modern keratose (horny) demosponges such as Darwinella.[116]
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. [117] In 2008 a detailed
analysis of chancelloriids' sclerites concluded that they were very similar to those of
halkieriids, mobile bilaterian animals that looked like slugs in chain mail 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. [115]

Relationships to other animal groups

A choanoflagellate
   
Fungi
   
    
Choanoflagellates
 
Metazoa  
Glass sponges
 
   
Demosponges
 
   
Calcareous sponges
   
  Eumetazoa  
Comb jellies
   
     
Placozoa
   
    Cnidaria
  (jellyfish, etc.)

Simplified family tree showing calcareous sponges

as closest to more complex animals [118]
   
Plants
 
 
Fungi
 
Metazoa  
Most demosponges
 
   
Calcareous sponges
   
   
Homoscleromorpha
   
  Eumetazoa   Cnidaria
    (jellyfish, etc.)
   
Other metazoans
 

Simplified family tree showing Homoscleromorpha

as closest to more complex animals [119]

In the 1990s sponges were widely regarded as a monophyletic group, all of them having
descended from a common ancestor that was itself a sponge, and as the "sister-group" to all
other metazoans (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
– 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.[118]

Analyses since 2001 have concluded that Eumetazoa (more complex than sponges) are more
closely related to particular groups of sponges than to the rest of the sponges. Such
conclusions imply that sponges are not monophyletic, because the last common ancestor 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 sponges and the rest, and that Eumetazoa are
more closely related to calcareous sponges, those with calcium carbonate spicules, than to
other types of sponge.[118] 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 calcareous sponges, which in turn are more
closely related to Eumetazoa.[103][120]

Other anatomical and biochemical evidence links the Eumetazoa with Homoscleromorpha, a
sub-group of demosponges. A comparison in 2007 of nuclear DNA, excluding glass sponges
and comb jellies, concluded that: Homoscleromorpha 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.[119] The sperm of Homoscleromorpha share with those of Eumetazoa
features that those 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 "type IV" collagen, a form of collagen not found in other sponges – although the spongin
fibers that reinforce the mesohyl of all demosponges is similar to "type IV" collagen. [30]

A comb jelly

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)
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.[16] 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.[121][122]

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.[4]

In March 2021, scientists from Dublin found additional evidence that sponges are the sister
group to all other animals.[123]

Notable spongiologists
 Céline Allewaert
 Patricia Bergquist
 James Scott Bowerbank
 Maurice Burton
 Henry John Carter
 Max Walker de Laubenfels
 Arthur Dendy
 Édouard Placide Duchassaing de Fontbressin
 Randolph Kirkpatrick
 Robert J. Lendlmayer von Lendenfeld
 Edward Alfred Minchin
 Giovanni Domenico Nardo
 Eduard Oscar Schmidt
 Émile Topsent

Use

Sponges made of sponge gourd for sale alongside sponges of animal origin (Spice Bazaar at
Istanbul, Turkey).

By dolphins

A report in 1997 described use of sponges as a tool by bottlenose dolphins in Shark Bay in
Western Australia. A dolphin will attach a marine sponge to its rostrum, which is presumably
then used to protect it when searching for food in the sandy sea bottom.[124] 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. [22]

By humans

Main article: Sea sponge aquaculture

Natural sponges in Tarpon Springs, Florida

Display of natural sponges for sale on Kalymnos in Greece

Skeleton

Main article: Sponge (material)

The calcium carbonate or silica spicules of most sponge genera make them too rough for
most uses, but two genera, Hippospongia and Spongia, have soft, entirely fibrous
skeletons.[125] 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 glazes
and discreet contraceptives. However, by the mid-20th century, over-fishing brought both the
animals and the industry close to extinction. [126] See also sponge diving.

Many objects with sponge-like textures are now made of substances not derived from
poriferans. Synthetic sponges include personal and household cleaning tools, breast
implants,[127] and contraceptive sponges.[128] Typical materials used are cellulose foam,
polyurethane foam, and less frequently, silicone foam.

The luffa "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).[129]

Antibiotic compounds
Sponges have medicinal potential due to the presence in sponges themselves or their
microbial symbionts of chemicals that may be used to control viruses, bacteria, tumors and
fungi.[130][131]

Other biologically active compounds

Main article: Sponge isolates

Halichondria produces the eribulin precursor halichondrin B

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, has shown potential as a cytotoxin to murine lymphoma cells. [132][133]

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