Sponge 47271 225854622 2008-07-15T18:36:14Z SpongeSebastian 1568441 /* See also */ "SpongeBob SquarePants" capitalization ''{{Merge-multiple|pinacoderm|spongocoel|sclerocyte|osculum|date=August 2007}} {{Refimprove|date=January 2008}} {{Otheruses4|the aquatic animal|the porous cleaning tool|Sponge (tool)|other uses|Sponge (disambiguation)}} {{Paraphyletic group | name = Sponge | fossil_range = [[Ediacaran]] - Recent | image = Sponge.JPG | image_width = 250px | regnum = [[Animal]]ia | domain = [[Eukaryota]] | phylum = "'''Porifera'''" (Paraphyletic) | symetry= none | phylum_authority = [[Robert Edmund Grant|Grant]] in [[Robert Bentley Todd|Todd]], 1836 | included =[[Calcareous sponge|Calcarea]]<br />[[Hexactinellid]]a<br />[[Demosponges|Demospongiae]] }} The '''sponges ''' or '''poriferans''' (from [[Latin language|Latin]] ''[[:wikt:porus|porus]]'' "pore" and ''[[:wikt:ferre|ferre]]'' "to bear") are [[animal]]s of the [[phylum]] '''Porifera''' ({{pronEng|pɒˈrɪfərə}}). '''Porifera''' translates to "Pore-bearer". They are primitive, [[sessile]], mostly [[ocean|marine]], water dwelling [[filter feeder]]s that pump water through their bodies to filter out particles of food matter. Sponges represent the [[body plan|simplest]] of animals. With no true tissues ([[parazoa]]), they lack [[muscle]]s, [[nerve]]s, and internal [[organ (anatomy)|organ]]s. Their similarity to colonial [[choanoflagellates]] shows the probable evolutionary jump from [[unicellular]] to [[multicellular]] organisms. However, recent genomic studies suggest they are not the most ancient lineage of animals, but may instead be secondarily simplified.<ref name=class>{{cite journal|author=Dunn ''et al.'' |year=2008|title=Broad phylogenomic sampling improves resolution of the animal tree of life|journal=Nature | volume = 452 | pages = 745 | doi = 10.1038/nature06614 <!--Retrieved from CrossRef by DOI bot-->|unused_data=|06614}}</ref> There are over 5,000 modern species of sponges known, and they can be found attached to surfaces anywhere from the [[intertidal zone]] to as deep as 8,500 [[metre|m]] (29,000 [[foot (unit of length)|feet]]) or further. Though the [[fossil]] record of sponges dates back to the [[Neoproterozoic]] Era, new species are still commonly discovered. ==Anatomy and morphology== [[Image:Tide pools sponge.jpg|Sponge in [[Tide pool]]|thumb|250px]] [[Image:Aphrocallistes vastus.jpg|thumb|250px|right|''Aphrocallistes vastus'' (Cloud sponge) is a type of glass sponge. They are very fragile, as they are made out of tiny glass crystals ([[Hydrated silica|hydrated silica dioxide]]).<ref>[http://www.biology.ualberta.ca/faculty/sally_leys/uploads/personal/Glass%20sponge%20ecology.htm Department of Biological Sciences - Studies in Life Sciences, University of Alberta: Glass Sponge Ecology], accessed [[March 16]], [[2008]]</ref>]] Sponges have several cell types: *[[Archaeocytes]] (or [[amoebocytes]]) have many functions; they are [[totipotent]] cells which can transform into [[sclerocytes]], [[spongocytes]], or [[collencytes]]. They also have a role in nutrient transport and sexual reproduction. *Cells are arranged in a gelatinous non-cellular [[Matrix (biology)|matrix]] called mesophyll Sponges have three body types: [[asconoid]], [[syconoid]], and [[leuconoid]]. *[[Choanocyte]]s (also known as "collar cells") function as the sponge's [[digestive system]], and are remarkably similar to the [[protist]]an [[choanoflagellate]]s. The collars are composed of many [[Microvillus|microvilli]] and are used to filter particles out of the water. The beating of the choanocytes’ [[flagellum|flagella]] creates the sponge’s water current. *[[Collencytes]] secrete [[collagen]]. *[[Myocytes (sponge)|Myocytes]] are modified pinacocytes which control the size of the osculum and pore openings and thus the water flow. *[[Pinacocytes]] which form the [[pinacoderm]], the outer epidermal layer of cells. This is the closest approach to true tissue in sponges *[[Porocytes]] are tubular cells that make up the pores into the sponge body through the [[mesohyl]]. *[[Sclerocytes]] secrete [[calcareous]] siliceous spicules which reside in the mesohyl. *[[Spongocytes]] secrete [[spongin]], [[collagen]]-like fibers which make up the mesohyl. *[[Spicule]]s are stiffened rods or spikes made of [[calcium carbonate]] or [[Silicon dioxide|silica]] which are used for structure and [[antipredator adaptation|defense]]. ''Asconoid'' sponges are tubular with a central shaft called the [[spongocoel]]. The beating of flagella forces water into the spongocoel through pores in the body wall. Choanocytes line the spongocoel and filter nutrients out of the water. ''Leuconoid'' sponges lack a sperm and instead have flagellated chambers, containing choanocytes, which are led to and out of via canals. ''Syconoid'' sponges are similar to asconoids. They have a tubular body with a single osculum, but the body wall is thicker and more complex than that of asconoids and contains choanocyte-lined radial canals that empty into the spongocoel. Water enters through a large number of dermal ostia into incurrent canals and then filters through tiny openings called [[prosopyle]]s into the radial canals. Their food is ingested by the choanocytes. Syconoids do not usually form highly branched colonies as asconoids do. During their development, syconoid sponges pass through an asconoid stage. It should be noted that these 3 body grades are useful only in describing [[morphology (biology)|morphology]], and not in classifying sponge species, althought the asconoid and syconoid construction is present in Calcarea only<ref>[http://animaldiversity.ummz.umich.edu/site/accounts/information/Calcarea.html Animal Diversity Web - Class Calcarea]</ref>. ==Physiology== Sponges have no true [[circulatory system]]; instead, they create a water current which is used for circulation. Dissolved gases are brought to cells and enter the cells via simple [[diffusion]]. [[Metabolic wastes]] are also transferred to the water through diffusion. Sponges pump remarkable amounts of water. ''Leuconia'', for example, is a small leuconoid sponge about 10 cm tall and 1 cm in diameter. It is estimated that water enters through more than 80,000 incurrent canals at a speed of 6cm 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 3.6cm per hour.<ref>See Hickman and Roberts (2001) Integrated principles of zoology — 11th ed., p.247</ref> Such a flow rate allows easy food capture by the collar cells. All water is expelled through a single [[osculum]] at a velocity of about 8.5 cm/second: a jet force capable of carrying waste products some distance away from the sponge. Sponges of the family Cladorhizidae (order Poecilosclerida, class Demospongiae) are species usually found in deep water, but also in [[Sea caves|littoral caves]] in the [[Mediterranean Sea|Mediterranean]] (Asbestopluma hypogea), that have become [[carnivorous]], using a strategy that has much in common with what is found in carnivorous plants such as [[sundew]]. When small crustaceans comes in contact with their surface, they get captured by a sticky substance, or in the case of the Mediterranean species by spicules modified into raised hook-shaped spines, and then digested by migrating cells which soon covers the prey<ref>[http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=1464536 Predation on copepods by an Alaskan cladorhizid sponge]</ref>. This lifestyle has caused the loss of their aquiferous system and the choanocytes, resulting in forms like the ping-pong tree sponge (Chondrocladia lampadiglobus), which don't look like typical sponges<ref>[http://www.nature.com/nature/journal/v373/n6512/abs/373333a0.html Carnivorous sponges]</ref>. ==Taxonomy== [[Image:Sponge in papua new guinea.jpg|thumb|250px|A sponge in [[Papua New Guinea]]]] Sponges are traditionally divided into classes based on the type of [[spicules]] in their skeleton. The three classes of sponges are bony ([[Calcaerous sponges|Calcarea]]), glass ([[Hexactinellid|Hexactenellida]]), and spongin ([[Demosponges|Demospongiae]]). Some taxonomists have suggested a fourth class, [[Sclerosponges|Sclerospongiae]], of coralline sponges, but the modern consensus is that coralline sponges have arisen several times and are not closely related.<ref>{{cite book|author = R. C. Brusca and G. J. Brusca|year = 2003|title = Invertebrates. Second Edition|location= Sunderland, Mass. | publisher=Sinauer Associates}}</ref> In addition to these four, a fifth, extinct class has been proposed: [[Archaeocyatha]]. While these ancient animals have been phylogenetically vague for years, the current general consensus is that they were a type of sponge.{{Fact|date=January 2008}} Although 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. Sponge taxonomy is an area of active research, with molecular studies improving our understanding of their relationship with other animals. ===Basal lineage?=== Sponges are among the simplest [[animals]]. They lack [[Gastrulation|gastrulated embryos]], [[Gastrointestinal tract|extracellular digestive cavities]], [[nerve]]s, [[muscle]]s, [[tissue (biology)|tissue]]s, and obvious sensory structures, features possessed by all other animals. In addition, sponge [[choanocyte]]s (feeding cells) appear to be a homologous to [[choanoflagellate]]s, a group of unicellular and colonial protists that are believed to be the immediate precursors of animals. The traditional conclusion is that sponges are the basal lineage of the animals, and that features such as tissues developed after sponges and other animals diverged. Sponges were first assigned their own subkingdom, the [[Parazoa]], but more recent molecular studies suggested that the sponges were [[paraphyletic]] to other animals, with the eumetazoa as a sister group to the most derived:<ref name=Sperling2007>{{cite journal | author = Sperling, E.A. | coauthors = Peterson, K.J. | year = 2007 | title = Poriferan paraphyly and its implications for Precambrian paleobiology | journal = J Geol Soc London | url = http://sp.lyellcollection.org/cgi/content/abstract/286/1/355 | accessdate = 2008-04-07 | doi = 10.1144/SP286.25 | volume = 286 | pages = 355 }}</ref> {{phylogeny|most [[Demosponge]]s|[[Calcareous sponge]]s|[[Homoscleromorpha]] demosponges|[[Eumetazoa]]}} Either way, sponges have long been considered useful models of the earliest multicellular ancestors of animals. ===...or secondarily simplified?=== However, a phylogenomic study in 2008 of 150 genes in 21 genera<ref name=class>{{cite journal|author=Dunn ''et al.'' |year=2008|title=Broad phylogenomic sampling improves resolution of the animal tree of life|journal=Nature | volume = 452 | pages = 745 | doi = 10.1038/nature06614 <!--Retrieved from CrossRef by DOI bot-->|unused_data=|06614}}</ref> suggests that the [[ctenophora]] are the most basal lineage of the 21 taxa sampled, and that sponges—or at least those lines of sponges investigated so far—are not primitive, but secondarily simplified, having lost tissues and other eumetazoan characteristics from their common ancestor. ==Geological history== [[Image:Fossil Sponge Raphidonema.jpg|thumb|250px|right|Fossil sponge ''Raphidonema faringdonense'' from the Cretaceous of England]] [[Image:BoredEncrustedShell.JPG|thumb|250px|right|Sponge borings and encrusters on a modern bivalve shell, North Carolina.]] The [[fossil record]] of sponges is not abundant. Some fossil sponges have worldwide distribution, while others are restricted to certain areas. Sponge fossils such as ''[[Hydnoceras]]'' and ''[[Prismodictya]]'' are found in the [[Devonian]] rocks of [[New York]] state. In [[Europe]] the [[Jurassic]] [[limestone]] of the [[Swabian Alb]] are composed largely of sponge remains, some of which are well preserved. Many sponges are found in the [[Cretaceous]] Lower Greensand and [[Chalk Formation]]s of [[England]], and in rocks from the upper part of the Cretaceous period in [[France]]. A famous locality for fossil sponges is the Cretaceous Faringdon Sponge Gravels in [[Faringdon]], [[Oxfordshire]] in [[England]]. An older sponge is the [[Cambrian]] ''[[Vauxia]]''. Sponges have long been important agents of [[bioerosion]] in shells and carbonate rocks. Their borings extend back to the [[Ordovician]] in the fossil record. Fossil sponges differ in size from 1 cm (0.4 inches) to more than 1 meter (3.3 feet). They vary greatly in shape, being commonly vase-shapes (such as ''Ventriculites''), spherical (such as ''Porosphaera''), saucer-shaped (such as ''Astraeospongia''), pear-shaped (such as ''Siphonia''), leaf-shaped (such as ''Elasmostoma''), branching (such as ''Doryderma''), irregular or encrusting. Detailed identification of many fossil sponges relies on the study of thin sections. ==Ecology== Modern sponges are predominantly marine, with some 150 species adapted to freshwater environments. Their habitats range from the inter-tidal zone to depths of 6,000 metres (19,680 feet). Certain types of sponges are limited in the range of depths at which they are found. Sponges are worldwide in their distribution, and range from waters of the polar regions to the tropical regions. Sponges are most abundant in both numbers of individuals and species in warmer waters. Adult sponges are largely sessile, and live in an attached position. However, it has been noted that certain sponges can move slowly by directing their water current in a certain direction with myocytes. The greatest numbers of sponges are usually to be found where a firm means of fastening is provided, such as on a rocky ocean bottom. Some kinds of sponges are able to attach themselves to soft sediment by means of a root-like base. Sponges also live in quiet clear waters, because if the sediment is agitated by wave action or by currents, it tends to block the pores of the animal, lessening its ability to feed and survive. Recent evidence suggests that a new disease called [[Aplysina red band syndrome]] (ARBS) is threatening sponges in the Caribbean.[http://www.practicalfishkeeping.co.uk/pfk/pages/item.php?news=1113] Aplysina red band syndrome causes Aplysina to develop one or more rust-coloured leading edges to their structure, sometimes with a surrounding area of necrotic tissue so that the lesion causes a contiguous band around some or all of the sponge's branch. ===Reproduction=== Sponges can reproduce [[sexual reproduction|sexually]] or [[asexual reproduction|asexually]]. Asexual reproduction is through internal and external [[budding]]. External budding occurs when the parent sponge grows a bud on the outside of its body. This will either break away or stay connected. Internal budding occurs when archaeocytes collect in the mesohyl and become surrounded by [[spongin]]. The internal bud is called a [[gemmule]], and this is seen only in the freshwater sponge family, the Spongillidae. An asexually reproduced sponge has exactly the same genetic material as the parent. In sexual reproduction, [[sperm]] are dispersed by water currents and enter neighboring sponges. All sponges of a particular species release their sperm at approximately the same time.{{Fact|date=January 2008}} Fertilization occurs internally, in the mesohyl. Fertilized oocytes develop within the mesohyl. Cleavage stages are highly varied within and between groups, sometimes even within a single species. Larval development usually involves an odd type of morphogenetic movement termed an inversion of layers. When this occurs in some species (for example, in ''Sycon coactum ''), the larva flips into the choanocyte chamber, and then can emerge via the water canal system and out through the osculum. Although sponges are [[hermaphroditic]] (both male and female), they are not self-fertile. Most sponges are sequential hermaphrodites, capable of producing eggs or sperm, but not both at the same time. ==Use== ===By dolphins=== In 1997, use of sponges as a [[tool]] was described in [[Bottlenose Dolphins]] in [[Shark Bay]]. A dolphin will attach a marine sponge to its [[rostrum (anatomy)|rostrum]], which is presumably then used to protect it when searching for food in the sandy [[sea floor|sea bottom]].<ref name="Smolker 1997">{{cite journal | author=Smolker, R.A., ''et al.'' | title=Sponge-carrying by Indian Ocean bottlenose dolphins: Possible tool-use by a delphinid }| Journal=Ethology | Year=1997 | Volume=103 | Pages=454–465}}</ref> The behaviour, known as ''sponging'', has only been observed in this bay, and is almost exclusively shown by females. This is the only known case of tool use in [[marine mammal]]s outside of [[Sea Otter]]s. An elaborate study in 2005 showed that mothers most likely teach the behaviour to their daughters.<ref name="Krutzen 2005">{{cite journal | author=Krutzen M, Mann J, Heithaus MR, Connor RC, Bejder L, Sherwin WB | title=Cultural transmission of tool use in bottlenose dolphins | journal=[[Proceedings of the National Academy of Sciences]] | volume=102 | issue=25 | year=2005 | pages=8939–8943 | doi=10.1073/pnas.0500232102 | pmid=15947077}}</ref> ===By humans=== [[Image:SpongesTarponSprings.jpg|right|250px|thumb|Natural Sponges in Tarpon Springs, Florida]] ==== Skeleton as absorbent==== {{main|Sponge (tool)}} In common usage, the term '''sponge''' is applied to the skeleton of the animal, from which the tissue has been removed by [[maceration (bone)|maceration]] and washing, leaving just the [[spongin]] scaffolding. [[calcium|Calcareous]] and [[silicon dioxide|siliceous]] sponges are too harsh for similar use. Commercial sponges are derived from various species and come in many grades, from fine soft "lamb's wool" sponges to the coarse grades used for washing cars. The manufacture of [[rubber]]-, [[plastic]]- and [[cellulose]]-based synthetic sponges has significantly reduced the commercial sponge [[fishing]] industry in recent years. The [[luffa]] "sponge", also spelled ''loofah'', commonly sold for use in the kitchen or the shower, is not derived from an animal sponge, but from the [[locule]]s of a gourd ([[Cucurbitaceae]]). ====Antibiotic compounds==== Sponges have [[medicine|medicinal]] potential due to the presence of [[antimicrobial]] compounds in either the sponge itself or their microbial [[symbiosis|symbiont]]s.<ref>See e.g. Teeyapant R, Woerdenbag HJ, Kreis P, Hacker J, Wray V, Witte L, Proksch P. (1993) Antibiotic and cytotoxic activity of brominated compounds from the marine sponge Verongia aerophoba. ''Zeitschrift für Naturforschung. C, Journal of biosciences'' '''48''':939–45.</ref> ==Bibliography== {{sourcesstart}} *{{cite book | author=C. Hickman Jr., L. Roberts and A. Larson | year=2003 | title=Animal Diversity | edition=3rd | publisher=McGraw-Hill | location=New York | id=ISBN 0-07-234903-4 }} *[http://www.practicalfishkeeping.co.uk/pfk/pages/item.php?news=1113 New disease threatens sponges], Practical Fishkeeping {{sourcesend}} ==References== {{reflist}} ==Further reading== *[[Patricia Bergquist|Berguist, P. R.]] 1998. "The Porifera" (pp. 10-27), in D. T. Anderson (ed.) ''Invertebrate Zoology''. (A brief treatment) *Berguist, P. R. 1978. ''Sponges'' Hutchinson, London. == See also == *[[Aaptos kanuux]] *[[Sponge Reef Project]] *[[SpongeBob SquarePants]] ==External links== {{Commons}} *[http://www.wooster.edu/geology/Bioerosion/Bioerosion.html Bioerosion website at The College of Wooster] *[http://www.spongepage.info/ Carsten's Spongepage], Information on the ecology and the biotechnological potential of sponges and their associated bacteria. *[http://news.nationalgeographic.com/news/2005/06/0607_050607_dolphin_tools.html Dolphin Moms Teach Daughters to Use Tools][[National Geographic]] *[http://fcit.usf.edu/FLORIDA/lessons/tarpon/tarpon.htm History of [[Tarpon Springs, Florida]] sponge industry] *[http://www.porifera.org/a/cif1.htm The Sponge Reef Project] {{Wikispecies|Porifera}} {{Wikibookspar|Dichotomous Key|Porifera}} *[http://www.qm.qld.gov.au/organisation/sections/SessileMarineInvertebrates/index.asp Queensland Museum FAQ about sponges] *[http://www.qm.qld.gov.au/organisation/sections/SessileMarineInvertebrates/spong.pdf Sponge Guide from Queensland Museum], John Hooper *[http://www.marinespecies.org/porifera/ World Porifera database], the world list of extant sponges, includes a searchable database. [[Category:Poriferans| ]] <!-- interwiki --> [[ar:إسفنجيات]] [[bg:Водни гъби]] [[ca:Porífer]] [[cs:Houbovci]] [[cy:Sbwng]] [[da:Svampedyr]] [[de:Schwämme]] [[et:Käsnad]] [[el:Σπόγγος]] [[es:Porifera]] [[eo:Spongulo]] [[eu:Belaki]] [[fa:اسفنج دریایی]] [[fr:Porifera]] [[gl:Porifera]] [[ko:해면동물]] [[hr:Spužve]] [[io:Sponjo]] [[id:Porifera]] [[is:Svampdýr]] [[it:Porifera]] [[he:ספוגיים]] [[la:Porifera]] [[lb:Schwämm]] [[lt:Pintys]] [[hu:Szivacsok]] [[mk:Сунѓери]] [[nl:Sponsdieren]] [[ja:海綿]] [[no:Svamper]] [[nn:Svampar]] [[oc:Porifera]] [[pl:Gąbki]] [[pt:Porifera]] [[ro:Spongieri]] [[qu:Puqyala]] [[ru:Губки]] [[simple:Sponge]] [[sk:Hubky]] [[sl:Spužve]] [[sr:Сунђери]] [[sh:Spužve]] [[fi:Sienieläimet]] [[sv:Svampdjur]] [[te:స్పంజిక]] [[th:ฟองน้ำ]] [[tr:Süngerler]] [[uk:Губки]] [[zh:多孔动物门]]