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In biology , gonochorism is a sexual system where there are two sexes and each individual organism is either male or female . The term gonochorism is usually applied in animal species, the vast majority of which are gonochoric.

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79-439: The lancelets ( / ˈ l æ n s l ɪ t s , ˈ l ɑː n -/ LA(H)N -slits ), also known as amphioxi ( sg. : amphioxus / ˌ æ m f i ˈ ɒ k s ə s / AM -fee- OK -səs ), consist of 32 described species of "fish-like" benthic filter feeding chordates in the subphylum Cephalochordata , class Leptocardii , and family Branchiostomatidae . Lancelets diverged from other chordates during or prior to

158-425: A heart . There are no blood cells, and no hemoglobin . The excretory system consists of segmented "kidneys" containing protonephridia instead of nephrons , and quite unlike those of vertebrates. Also unlike vertebrates, there are numerous, segmented gonads . Lancelets became famous in the 1860s when Ernst Haeckel began promoting them as a model for the ancestor of all vertebrates. By 1900 lancelets had become

237-436: A model organism . By the mid-20th century they had fallen out of favor for a variety of reasons, including a decline of comparative anatomy and embryology, and due to the belief that lancelets were more derived than they appeared, e.g., the profound asymmetry in the larval stage. More recently, the fundamental symmetric and twisted development of vertebrates is the topic of the axial twist theory . According to this theory, there

316-424: A monomeric green fluorescent protein known as mNeonGreen , which is the brightest known monomeric green or yellow fluorescent protein. Lancelets are passive filter feeders , spending most of the time half-buried in sand with only their frontal part protruding. They eat a wide variety of small planktonic organisms, such as bacteria, fungi , diatoms , and zooplankton , and they will also take detritus . Little

395-599: A 2017 study involving 165 taxon groups, more evolutionary transitions from gonochorism to hermaphroditism were found than the reverse. The term is most often used with animals, in which the species are usually gonochoric. Gonochorism has been estimated to occur in 95% of animal species. It is very common in vertebrate species, 99% of which are gonochoric. 98% of fishes are gonochoric. Mammals (including humans ) and birds are solely gonochoric. Tardigrades are almost always gonochoric. 75% of snails are gonochoric. Most arthropods are gonochoric. For example

474-499: A cup-shaped pigment cell. The peak sensitivity of both cells is ~470 nm (blue). Both the Joseph cells and Hesse organs are in the neural tube, the Joseph cells forming a dorsal column, the Hesse organs in the ventral part along the length of the tube. The Joseph cells extend from the caudal end of the anterior vesicle (or cerebral vesicle) to the boundary between myomeres 3 and 4, where

553-400: A filter-feeder due to its bill proportions being similar to those of shoveler ducks . It is unique in being a large, flightless marine animal, unlike the smaller still volant flamingos and prions. Traditionally, Ctenochasmatoidea as a group has been listed as filter-feeders, due to their long, multiple slender teeth, clearly well adapted to trap prey. However, only Pterodaustro showcases

632-675: A food additive in dry form after being roasted in oil. When their gonads start to ripen in the spring it affects their flavor, making them taste bad during their breeding season. The lancelets were traditionally seen as the sister lineage to the vertebrates ; in turn, these two groups together (sometimes called Notochordata) were considered the sister group to the Tunicata (also called Urochordata and including sea squirts ). Consistent with this view, at least 10 morphological features are shared by lancelets and vertebrates, but not tunicates. Newer research suggests this pattern of evolutionary relationship

711-466: A highly effective source of fertilizer and animal feed In the U.S., researchers are investigating potential to model the use of shellfish and seaweed for nutrient mitigation in certain areas of Long Island Sound. Bivalves are also largely used as bioindicators to monitor the health of an aquatic environment, either fresh- or seawater. Their population status or structure, physiology, behaviour, or their content of certain elements or compounds can reveal

790-517: A kind of rudimentary filter feeding, using their long, slender teeth to trap small fish, though probably lacking the pumping mechanism of Pterodaustro . In essence, their foraging mechanism was similar to that of modern young Platanista " dolphins ". Filter feeding habits are conspicuously rare among Mesozoic marine reptiles , the main filter feeding niche being seemingly instead occupied by pachycormid fish. However, some sauropsids have been suggested to have engaged in filter feeding. Henodus

869-696: A majority of crustaceans are gonochoric. In animals, sex is most often genetically determined, but may be determined by other mechanisms. For example, alligators use temperature-dependent sex determination during egg incubation. Plants which have single-sex individuals are typically called dioecious ( vascular plants ) or dioicous ( bryophytes ) instead of gonochoric. In flowering plants , individual flowers may be hermaphroditic (i.e. with both stamens and ovaries) or dioecious (unisexual), having either no stamens (i.e. no male parts) or no ovaries (i.e. no female parts). Among flowering plants with unisexual flowers, some also produce hermaphrodite flowers, and

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948-438: A proper pumping mechanism, having up-turned jaws and powerful jaw and tongue musculature. Other ctenochasmatoids lack these, and are now instead thought to have been spoonbill -like catchers, using their specialised teeth simply to offer a larger surface area. Tellingly, these teeth, while small and numerous, are comparatively unspecialised to the baleen-like teeth of Pterodaustro . Boreopterids are thought to have relied on

1027-409: A small number of female gonads were observed within male individuals, typically ranging from 2 to 5 gonads out of a total of 45–50. An extraordinary occurrence of complete sex reversal was documented in B. belcheri, where a female amphioxus raised in laboratory conditions underwent a transformation into a male (Zhang et al., 2001). Their habitat preference reflects their feeding method: they only expose

1106-452: A speed of 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. Such a flow rate allows easy food capture by the collar cells. 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

1185-725: A swarm gulping, while lowering their tongue so that the head's ventral grooves expand and vastly increase the amount of water taken in. Baleen whales typically eat krill in polar or subpolar waters during summers, but can also take schooling fish, especially in the Northern Hemisphere. All baleen whales except the gray whale feed near the water surface, rarely diving deeper than 100 m (330 ft) or for extended periods. Gray whales live in shallow waters feeding primarily on bottom-living organisms such as amphipods . Bivalves are aquatic molluscs which have two-part shells . Typically both shells (or valves) are symmetrical along

1264-405: A true liver but a homolog of the vertebrate liver. Lancelets have no respiratory system, breathing solely through their skin, which consists of a simple epithelium . Despite the name, little if any respiration occurs in the "gill" slits, which are solely devoted to feeding. The circulatory system does resemble that of primitive fish in its general layout, but is much simpler, and does not include

1343-424: 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

1422-585: Is "an environmental management strategy by which nutrients are removed from an aquatic ecosystem through the harvest of enhanced biological production, including the aquaculture of suspension-feeding shellfish or algae". Nutrient removal by shellfish, which are then harvested from the system, has the potential to help address environmental issues including excess inputs of nutrients ( eutrophication ), low dissolved oxygen, reduced light availability and impacts on eelgrass, harmful algal blooms, and increases in incidence of paralytic shellfish poisoning (PSP). For example,

1501-513: Is a deep agreement between the vertebrates and cephalochordates, and even all chordates. With the advent of molecular genetics lancelets are once again regarded as a model of vertebrate ancestors, and are used again as a model organism. As a result of their use in science, methods of keeping and breeding lancelets in captivity have been developed for several of the species, initially the European Branchiostoma lanceolatum , but later also

1580-401: Is accomplished by atrial contraction. The remainder of the digestive system consists of a simple tube running from the pharynx to the anus. The hepatic caecum , a single blind-ending caecum , branches off from the underside of the gut, with a lining able to phagocytize the food particles, a feature not found in vertebrates. Although it performs many functions of a liver, it is not considered

1659-403: Is accomplished through filter feeding, using the krill's developed front legs, providing for a very efficient filtering apparatus: the six thoracopods form a very effective "feeding basket" used to collect phytoplankton from the open water. In the animation at the top of this page, the krill is hovering at a 55° angle on the spot. In lower food concentrations, the feeding basket is pushed through

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1738-405: Is derived from Greek gone 'generation' + chorizein 'to separate'. The term gonochorism originally came from German Gonochorismus . Gonochorism is also referred to as unisexualism or gonochory. Gonochorism has evolved independently multiple times. It is very evolutionarily stable in animals. Its stability and advantages have received little attention. Gonochorism owes its origin to

1817-539: Is incorrect. Extensive molecular phylogenetic analysis has shown convincingly that the Cephalochordata is the most basal subphylum of the chordates, with tunicates being the sister group of the vertebrates. This revised phylogeny of chordates suggests that tunicates have secondarily lost some of the morphological characters that were formerly considered to be synapomorphies (shared, derived characters) of vertebrates and lancelets. Lancelets have turned out to be among

1896-403: Is known about the diet of the lancelet larvae in the wild, but captive larvae of several species can be maintained on a diet of phytoplankton , although this apparently is not optimal for Asymmetron lucayanum . Lancelets have oral cirri, thin tentacle -like strands that hang in front of the mouth and act as sensory devices and as a filter for the water passing into the body. Water passes from

1975-780: Is likely that currently unrecognized cryptic species remain. The cladogram presented here illustrates the phylogeny (family tree) of lancelets, and follows a simplified version of the relationships found by Igawa, T.; M. Nozawa; D.G. Suzuki; J.D. Reimer; A.R. Morov; Y. Wang; Y. Henmi; K. Yasui (2017): Olfactores [REDACTED] † Cathaymyrus ? († 518 mya) [REDACTED] † Palaeobranchiostoma ? († 273.01 mya) [REDACTED] Asymmetron inferum Asymmetron lucayanum [REDACTED] Filter feeding Filter feeders are aquatic animals that acquire nutrients by feeding on organic matters , food particles or smaller organisms ( bacteria , microalgae and zooplanktons ) suspended in water, typically by having

2054-577: Is only reported in the Asymmetron genus. Multiple fluorescent protein genes have been recorded in lancelet species throughout the world. Branchiostoma floridae alone has 16 GFP-encoding genes. However, the GFP produced by lancelets is more similar to GFP produced by copepods than jellyfish ( Aequorea victoria ). It is suspected GFP plays multiple roles with lancelets such as attracting plankton towards their mouth. Considering that lancelets are filter feeders,

2133-426: Is present, so they are not especially good swimmers. While they do possess some cartilage material stiffening the gill slits , mouth, and tail, they have no true complex skeleton. In common with vertebrates, lancelets have a hollow nerve cord running along the back, pharyngeal slits and a tail that runs past the anus. Also like vertebrates, the muscles are arranged in blocks called myomeres . Unlike vertebrates,

2212-481: The Atlantic menhaden , a type of herring , lives on plankton caught in midwater. Adult menhaden can filter up to four gallons of water a minute and play an important role in clarifying ocean water. They are also a natural check to the deadly red tide . In addition to these bony fish, four types of cartilaginous fishes are also filter feeders. The whale shark sucks in a mouthful of water, closes its mouth and expels

2291-716: The Cambrian period. A number of fossil chordates have been suggested to be closely related to lancelets, including Pikaia and Cathaymyrus from the Cambrian and Palaeobranchiostoma from the Permian , but their close relationship to lancelets has been doubted by other authors. Molecular clock analysis suggests that modern lancelets probably diversified much more recently, during the Cretaceous or Cenozoic . Zoologists are interested in them because they provide evolutionary insight into

2370-471: The Cetacea (whales, dolphins, and porpoises), are characterized by having baleen plates for filtering food from water, rather than teeth. This distinguishes them from the other suborder of cetaceans, the toothed whales (Odontoceti). The suborder contains four families and fourteen species. Baleen whales typically seek out a concentration of zooplankton, swim through it, either open-mouthed or gulping, and filter

2449-416: The epipharyngeal groove , and passed back to the rest of the digestive tract. Having passed through the gill slits, the water enters an atrium surrounding the pharynx, then exits the body via the atriopore. Both adults and larvae exhibit a "cough" reflex to clear the mouth or throat of debris or items too large to swallow. In larvae the action is mediated by the pharyngeal muscles while in the adult animal it

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2528-425: The tegmental neuropil . The tegmental neuropil has been compared with locomotor control regions of the vertebrate hypothalamus , where paracrine release modulates locomotor patterns such as feeding and swimming. Lancelets naturally express green fluorescent proteins (GFP) inside their oral tentacles and near the eye spot. Depending on the species, it can also be expressed in the tail and gonads , though this

2607-554: The vertebrates . Nearly all tunicates are suspension feeders , capturing planktonic particles by filtering sea water through their bodies. Water is drawn into the body through the inhalant buccal siphon by the action of cilia lining the gill slits. The filtered water is then expelled through a separate exhalant siphon. To obtain enough food, a typical tunicate needs to process about one body-volume of water per second. Flamingos filter-feed on brine shrimp . Their oddly shaped beaks are specially adapted to separate mud and silt from

2686-485: The Bahamas lancelet, Asymmetron lucayanum The nerve cord is only slightly larger in the head region than in the rest of the body, so that lancelets do not appear to possess a true brain. However, developmental gene expression and transmission electron microscopy indicate the presence of a diencephalic forebrain , a possible midbrain , and a hindbrain . Recent studies involving a comparison with vertebrates indicate that

2765-524: The Hesse organs begin and continue nearly to the tail. The frontal eye consists of a pigment cup, a group of photoreceptor cells (termed Row 1 ), three rows of neurons ( Rows 2–4 ), and glial cells . The frontal eye, which expresses the PAX6 gene, has been proposed as the homolog of vertebrate paired eyes,or the pineal eye on vertebrates, the pigment cup as the homolog of the RPE ( retinal pigment epithelium ),

2844-627: The West Pacific Branchiostoma belcheri and Branchiostoma japonicum , the Gulf of Mexico and West Atlantic Branchiostoma floridae and the circumtropical (however, genetic evidence suggest the Atlantic and Indo-Pacific populations should be recognized as separate) Asymmetron lucayanum . They can reach an age of up to 7–8 years. The animals are edible and harvested in some parts of the world. They are eaten both fresh, tasting like herring, and as

2923-420: The average harvested mussel contains: 0.8–1.2% nitrogen and 0.06–0.08% phosphorus Removal of enhanced biomass can not only combat eutrophication and also support the local economy by providing product for animal feed or compost. In Sweden, environmental agencies utilize mussel farming as a management tool in improving water quality conditions, where mussel bioextraction efforts have been evaluated and shown to be

3002-495: The beating of cilia . Suspended food ( phytoplankton , zooplankton , algae and other water-borne nutrients and particles) are trapped in the mucus of a gill, and from there are transported to the mouth, where they are eaten, digested and expelled as feces or pseudofeces . Each oyster filters up to five litres of water per hour. Scientists believe that the Chesapeake Bay 's once-flourishing oyster population historically filtered

3081-926: The case of the buoy barnacle ) depending on the species and the niches they have evolved to occupy. Extant species that rely on such method of feeding encompass numerous phyla , including poriferans ( sponges ), cnidarians ( jellyfish , sea pens and corals ), arthropods ( krill , mysids and barnacles ), molluscs ( bivalves , such as clams , scallops and oysters ), echinoderms ( sea lilies ) and chordates ( lancelets , sea squirts and salps , as well as many marine vertebrates such as most species of forage fish , American paddlefish , silver and bighead carps , baleen whales , manta ray and three species of sharks —the whale shark , basking shark and megamouth shark ). Some water birds such as flamingos and certain duck species, though predominantly terrestrial, are also filter feeders when foraging . Most forage fish are filter feeders. For example,

3160-476: The cells lie outside of the pigment cup, while the cilia extend into the pigment cup before turning and exiting. The cells bear the opsin c-opsin 1 , except for a few which carry c-opsin 3 . The Row 2 cells are serotonergic neurons in direct contact with Row 1 cells. Row 3 and 4 cells are also neurons. Cells of all four rows have axons that project into the left and right ventrolateral nerves. For Row 2 neurons, axon projections have been traced to

3239-562: The contamination status of any aquatic ecosystem. They are useful as they are sessile, which means they are closely representative of the environment where they are sampled or placed (caging), and they breathe water all the time, exposing their gills and internal tissues: bioaccumulation . One of the most famous projects in that field is the Mussel Watch Programme in America. Sponges have no true circulatory system ; instead, they create

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3318-442: The dorsal nerve cord is not protected by bone but by a simpler notochord made up of a cylinder of cells that are closely packed in collagen fibers to form a toughened rod. The lancelet notochord, unlike the vertebrate spine , extends into the head. This gives the subphylum, Cephalochordata, its name ( κεφαλή , kephalē means 'head'). The fine structure of the notochord and the cellular basis of its adult growth are best known for

3397-453: The early evolution of vertebrates: by comparing genes from lancelets with the same genes in vertebrates, changes in gene expression, function and number as vertebrates evolved can be discovered. The genome of a few species in the genus Branchiostoma have been sequenced: B. floridae, B. belcheri , and B. lanceolatum . In Asia, lancelets are harvested commercially as food for humans. In Japan, amphioxus ( B. belcheri ) has been listed in

3476-428: The enlarged lower lip which fits onto the bowed upper jaw. As the right whale swims, a front gap between the two rows of baleen plates lets the water in together with the prey, while the baleens filter out the water. Rorquals such as the blue whale , in contrast, have smaller heads, are fast swimmers with short and broad baleen plates. To catch prey, they widely open their lower jaw — almost 90° — swim through

3555-461: The estuary's entire water volume of excess nutrients every three or four days. Today that process would take almost a year, and sediment, nutrients, and algae can cause problems in local waters. Oysters filter these pollutants, and either eat them or shape them into small packets that are deposited on the bottom where they are harmless. Bivalve shellfish recycle nutrients that enter waterways from human and agricultural sources. Nutrient bioextraction

3634-570: The evolution of anisogamy , but it is unclear if the evolution of anisogamy first led to hermaphroditism or gonochorism. Gonochorism is thought to be the ancestral state in polychaetes , hexacorallia , nematodes , and hermaphroditic fishes . Gonochorism is thought to be ancestral in hermaphroditic fishes because it is widespread in basal clades of fish and other vertebrate lineages. Two papers from 2008 have suggested that transitions between hermaphroditism and gonochorism or vice versa have occurred in animals between 10 and 20 times. In

3713-473: The exact species involved, the maximum length of lancelets is typically 2.5 to 8 cm (1.0–3.1 in). Branchiostoma belcheri and B. lanceolatum are among the largest. Except for the size, the species are very similar in general appearance, differing mainly in the number of myotomes and the pigmentation of their larvae. They have a translucent, somewhat fish-like body, but without any paired fins or other limbs. A relatively poorly developed tail fin

3792-507: The filter between the gill bars is swallowed. Whale sharks have been observed "coughing" and it is presumed that this is a method of clearing a build up of food particles in the gill rakers. The megamouth shark has luminous organs called photophores around its mouth. It is believed they may exist to lure plankton or small fish into its mouth. The basking shark is a passive filter feeder, filtering zooplankton , small fish, and invertebrates from up to 2,000 tons of water per hour. Unlike

3871-401: The first complete description of amphioxus embryos, while Schultze and Leuckart were the first to describe the larvae. Other important contributions to amphioxus embryonic anatomy were given by Hatschek, Conklin and later by Tung (experimental embryology). The larvae are extremely asymmetrical, with the mouth and anus on the left side, and the gill slits on the right side. Organs associated with

3950-411: The food they eat, and are uniquely used upside-down. The filtering of food items is assisted by hairy structures called lamellae which line the mandibles , and the large rough-surfaced tongue. Prions are specialised petrels with filter-feeding habits. Their name comes from their saw-like jaw edges, used to scope out small planktionic animals. The extinct swan Annakacygna is speculated to be

4029-798: The front end to the water and filter-feed on plankton by means of a branchial ciliary current that passes water through a mucous sheet. Branchiostoma floridae is capable of trapping particles from microbial to small phytoplankton size, while B. lanceolatum preferentially traps bigger particles (>4 μm). Lancelets are gonochoric animals, i.e. having two sexes, and they reproduce via external fertilization . They only reproduce during their spawning season, which varies slightly between species — usually corresponding to spring and summer months. All lancelets species spawn shortly after sunset, either synchronously (e.g. Branchiostoma floridae , about once every two weeks during spawning season) or asynchronously ( Branchiostoma lanceolatum , gradual spawning through

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4108-503: The genus Limax . It was not until 1834 that Gabriel Costa brought the phylogenetic position of the group closer to the agnathan vertebrates ( hagfish and lampreys ), including it in the new genus Branchiostoma (from the Greek, branchio = "gills", stoma = "mouth"). In 1836, Yarrell renamed the genus as Amphioxus (from the Greek: "pointed on both sides"), now considered an obsolete synonym of

4187-577: The genus Branchiostoma . Today, the term "amphioxus" is still used as a common name for the Amphioxiformes, along with "lancelet", especially in the English language. All living lancelets are all placed in the family Branchiostomatidae, class Leptocardii, and subphylum Cephalochordata. The family was first named by Charles Lucien Bonaparte in 1846, though he used the incorrect spelling "Branchiostomidae". One year previously, Johannes Müller had introduced

4266-448: The hinge line. The class has 30,000 species , including scallops , clams , oysters and mussels . Most bivalves are filter feeders (although some have taken up scavenging and predation), extracting organic matter from the sea in which they live. Nephridia , the shellfish version of kidneys , remove the waste material. Buried bivalves feed by extending a siphon to the surface. For example, oysters draw water in over their gills through

4345-413: The megamouth and whale sharks, the basking shark does not appear to actively seek its quarry; but it does possess large olfactory bulbs that may guide it in the right direction. Unlike the other large filter feeders, it relies only on the water that is pushed through the gills by swimming; the megamouth shark and whale shark can suck or pump water through their gills. Manta rays can time their arrival at

4424-459: The middle of the 19th century. First, the adult then the embryonic anatomy were described. Alexander Kowalevsky first described the key anatomical features of the adult amphioxus (hollow dorsal nerve tube, endostyle, segmented body, postanal tail). De Quatrefages first completely described the nervous system of amphioxus. Other important contributions to amphioxus adult anatomy were given by Heinrich Rathke and John Goodsir. Kowalevsky also released

4503-478: The most genetically diverse animals sequenced to date, due to high rates of genetic changes like exon shuffling and domain combination. Among the three extant (living) genera , Asymmetron is basal. Molecular clock studies have come to different conclusions on their divergence, with some suggesting that Asymmetron diverged from other lancelets more than 100 million years ago while others have suggested that it occurred about 46 million years ago. According to

4582-427: The mouth into the large pharynx , which is lined by numerous gill-slits. The ventral surface of the pharynx contains a groove called the endostyle , which, connected to a structure known as Hatschek's pit , produces a film of mucus . Ciliary action pushes the mucus in a film over the surface of the gill slits, trapping suspended food particles as it does so. The mucus is collected in a second, dorsal groove, known as

4661-424: The name Leptocardii as a subclass. Finally, the subphylum name Cephalochordata is attributed to Ernst Haeckel (1866). At the taxonomic rank of order, lancelets are sometimes placed in the order Amphioxi Bonaparte, 1846, Amphioxiformes Berg, 1937, or Branchiostomiformes Fowler, 1947. Another name sometimes used for high-ranked taxa for the lancelets is Acrania Haeckel, 1866. Observations of amphioxus anatomy began in

4740-475: The natural current would draw nearby plankton into the digestive tract. GFP is also expressed in larvae , signifying it may be used for photoprotection by converting higher energy blue light to less harmful green light. The fluorescent proteins from lancelets have been adapted for use in molecular biology and microscopy. The yellow fluorescent protein from Branchiostoma lanceolatum exhibits unusually high quantum yield (~0.95). It has been engineered into

4819-400: The neural tube, with the frontal eye at the front, followed by the lamellar body, the Joseph cells, and the Hesse organs. Joseph cells are bare photoreceptors surrounded by a band of microvilli . These cells bear the opsin melanopsin . The Hesse organs (also known as dorsal ocelli) consist of a photoreceptor cell surrounded by a band of microvilli and bearing melanopsin, but half enveloped by

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4898-502: The origins of vertebrates . Lancelets contain many organs and organ systems that are homologous to those of modern fish, but in a more primitive form. Therefore, they provide a number of examples of possible evolutionary exaptation . For example, the gill-slits of lancelets are used for feeding only, and not for respiration. The circulatory system carries food throughout their body, but does not have red blood cells or hemoglobin for transporting oxygen. Lancelet genomes hold clues about

4977-551: The pharynx are positioned either exclusively on the left or on the right side of the body. In addition, segmented muscle blocks and parts of the nervous system are asymmetrical. After metamorphosis the anatomy becomes more symmetrical, but some asymmetrical traits are still present also as adults, such as the nervous system and the location of the gonads which are found on the right side in Asymmetron and Epigonichthys (in Branchiostoma gonads develop on both sides of body). Depending on

5056-500: The prey from the water using their baleens. A baleen is a row of a large number of keratin plates attached to the upper jaw with a composition similar to those in human hair or fingernails. These plates are triangular in section with the largest, inward-facing side bearing fine hairs forming a filtering mat. Right whales are slow swimmers with large heads and mouths. Their baleen plates are narrow and very long — up to 4 m (13 ft) in bowheads  — and accommodated inside

5135-516: The putative photoreceptors as homologs of vertebrate rods and cones , and Row 2 neurons as homologs of the retinal ganglion cells . The pigment cup is oriented concave dorsally. Its cells contain the pigment melanin . The putative photoreceptor cells, Row 1, are arranged in two diagonal rows, one on either side of the pigment cup, symmetrically positioned with respect to the ventral midline. The cells are flask-shaped, with long, slender ciliary processes (one cilium per cell). The main bodies of

5214-492: The registry of "Endangered Animals of Japanese Marine and Fresh Water Organisms". Adult amphioxus typically inhabit the seafloor, burrowing into well-ventilated substrates characterized by a soft texture and minimal organic content. While various species have been observed in different types of substrate, such as fine sand, coarse sand, and shell deposits, most exhibit a distinct preference for coarse sand with low levels of fine particles. For instance, Branchiostoma nigeriense along

5293-416: The sea floor, constantly collecting particles with their filter basket. They are an important food source for herring , cod , flounder , and striped bass . Mysids have a high resistance to toxins in polluted areas, and may contribute to high toxin levels in their predators. Antarctic krill manages to directly utilize the minute phytoplankton cells, which no other higher animal of krill size can do. This

5372-439: The season). Nicholas and Linda Holland were the first researchers to describe a method of obtaining amphioxus embryos by induction of spawning in captivity and in vitro fertilization. Spawning can be artificially induced in the lab by electric or thermal shock. The first representative organism of the group to be described was Branchiostoma lanceolatum . It was described by Peter Simon Pallas in 1774 as molluscan slugs in

5451-431: The spawning of large shoals of fish and feed on the free-floating eggs and sperm. This stratagem is also employed by whale sharks. Like all arthropods, crustaceans are ecdysozoans , a clade without cilia . Cilia play an important role for many filter feeding animals, but because crustaceans don't have them, they need to use modified extremities for filter feeding instead. Mysidaceans live close to shore and hover above

5530-407: The sponge. The moon jellyfish has a grid of fibres which are slowly pulled through the water. The motion is so slow that copepods cannot sense it and do not react with an escape response . Other filter-feeding cnidarians include sea pens , sea fans , plumose anemones , and Xenia . Tunicates , such as ascidians , salps and sea squirts , are chordates which form a sister group to

5609-405: The three types may occur in different arrangements on the same or separate plants. Plant species can thus be hermaphrodite, monoecious , dioecious , trioecious , polygamomonoecious , polygamodioecious , andromonoecious , or gynomonoecious . Examples of species with gonochoric or dioecious pollination include hollies and kiwifruit . In these plants the male plant that supplies the pollen

5688-656: The unrelated Cenozoic Mourasuchus shares similar adaptations. Hupehsuchia is a lineage of bizarre Triassic reptiles adapted for suspension feeding. Some plesiosaurs might have had filter-feeding habits. Gonochorism Gonochorism contrasts with simultaneous hermaphroditism but it may be hard to tell if a species is gonochoric or sequentially hermaphroditic e.g. parrotfish , Patella ferruginea . However, in gonochoric species individuals remain either male or female throughout their lives. Species that reproduce by thelytokous parthenogenesis and do not have males can still be classified as gonochoric. The term

5767-422: The vertebrate thalamus , pretectum , and midbrain areas jointly correspond to a single, combined region in the amphioxus, which has been termed di-mesencephalic primordium (DiMes). Lancelets have four known kinds of light-sensing structures: Joseph cells, Hesse organs, an unpaired anterior eye and lamellar body, all of which utilize opsins as light receptors. All of these organs and structures are located in

5846-749: The water for over half a meter in an opened position, and then the algae are combed to the mouth opening with special setae on the inner side of the thoracopods. Porcelain crabs have feeding appendages covered with setae to filter food particles from the flowing water. Most species of barnacles are filter feeders, using their highly modified legs to sift plankton from the water. Also some insects with aquatic larvae or nymphs are filter feeders during their aquatic stage. Such as some species of mayfly nymphs, mosquito larvae, and black fly larvae. Instead of using modified limbs or mouthparts, some caddisfly larvae produce nets of silk used for filter feeding. The baleen whales (Mysticeti), one of two suborders of

5925-504: The water pass over or through a specialized filtering organ that sieves out and/or traps solids. Filter feeders can play an important role in condensing biomass and removing excess nutrients (such as nitrogen and phosphate ) from the local waterbody , and are therefore considered water-cleaning ecosystem engineers . They are also important in bioaccumulation and, as a result, as indicator organisms . Filter feeders can be sessile , planktonic , nektonic or even neustonic (in

6004-428: The water through its gills . During the slight delay between closing the mouth and opening the gill flaps, plankton is trapped against the dermal denticles which line its gill plates and pharynx . This fine sieve-like apparatus, which is a unique modification of the gill rakers, prevents the passage of anything but fluid out through the gills (anything above 2 to 3 mm in diameter is trapped). Any material caught in

6083-910: The west coast of Africa, Branchiostoma caribaeum in Mississippi Sound and along the coast from South Carolina to Georgia, B. senegalense in the offshore shelf region off North West Africa, and B. lanceolatum along the Mediterranean coast of southern France all demonstrate this preference (Webb and Hill, 1958; Webb, 1958; Boschung and Gunter, 1962; Cory and Pierce, 1967; Gosselck and Spittler, 1979; Caccavale et al., 2021b; Desdevises et al., 2011). However, Branchiostoma floridae from Tampa Bay, Florida, appears to be an exception to this trend, favoring fine sand bottoms instead (Stokes and Holland, 1996a; Stokes, 1996). All amphioxus species exhibit gonochorism , with only rare instances of hermaphroditism reported in Branchiostoma lanceolatum and B. belcheri. In these cases,

6162-443: The younger estimation, Branchiostoma and Epigonichthys have been estimated to have diverged from each other about 38.3 million years ago. Despite this deep separation, hybrids between Asymmetron lucayanum and Branchiostoma floridae are viable (among the deepest split species known to be able to produce such hybrids). The following are the species recognised by WoRMS . Other sources recognize about thirty species. It

6241-454: Was a placodont with unique baleen-like denticles and features of the hyoid and jaw musculature comparable to those of flamingos. Combined with its lacustrine environment, it might have occupied a similar ecological niche. In particular, it was probably a herbivore , filtering out algae and other small-sized flora from the substrates. Stomatosuchidae is a family of freshwater crocodylomorphs with rorqual-like jaws and minuscule teeth, and

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