Acorn worm

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Acorn worms
Temporal range: MiaolingianPresent, 505–0  Ma
Enteropneusta.png
By Johann Wilhelm Spengel  [ de ], 1893
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Hemichordata
Class: Enteropneusta
Gegenbaur, 1870
Order: Enteropneusta
Families

The acorn worms or Enteropneusta are a hemichordate class of invertebrates consisting of one order of the same name. [2] The closest non-hemichordate relatives of the Enteropneusta are the echinoderms. [3] There are 111 known species of acorn worm in the world, [4] the main species for research being Saccoglossus kowalevskii . Two families—Harrimaniidae and Ptychoderidae—separated at least 370 million years ago. [5]

Contents

Until recently, it was thought that all species lived in the sediment on the seabed, subsisting as deposit feeders or suspension feeders. However, the early 21st century has seen the description of a new family, the Torquaratoridae, evidently limited to the deep sea, in which most of the species crawl on the surface of the ocean bottom and alternatively rise into the water column, evidently to drift to new foraging sites. [6] [7] [8] [9] [10] It is assumed that the ancestors of acorn worms used to live in tubes like their relatives Pterobranchia, but that they eventually started to live a safer and more sheltered existence in sediment burrows instead. [11] The body length normally range from 2 centimetres (0.79 in) to 2.5 metres (8 ft 2 in) ( Balanoglossus gigas ), [12] but one species, Meioglossus psammophilus , only reach 0.6 millimetres (0.024 in). [13] Due to secretions containing elements like iodine, the animals have an iodoform-like smell. [14]

Anatomy

Structure of branchial region - bc, coelom. tb, tongue-bars. ds, mesentery. pr, ridge. vv, vessel. gp, gill-pore. dn, dorsal nerve. dv, vessel. oe, oesophagus. vs, mesentery. vn, ventral nerve. Balanoglossus 2.png
Structure of branchial region bc, coelom. tb, tongue-bars. ds, mesentery. pr, ridge. vv, vessel. gp, gill-pore. dn, dorsal nerve. dv, vessel. œ, oesophagus. vs, mesentery. vn, ventral nerve.
Structure of anterior end - a, Arrow from proboscis-cavity (pc) passing to left of pericardium (per) and out through proboscis pore-canal. b , arrow from central canal of neurochord (cnc) passed out through anterior neuropore. b , ditto; through posterior neuropore. c, arrow intended to pass from 1st gill-pouch through collar pore-canal into collar-coelom (cc). cts, posterior limit of collar. dv, dorsal vessel passing into central sinus (bs). ev, efferent vessel passing into ventral vessel (vv). epr, epiphysial tubes. st, stomochord. vs, ventral septum of proboscis. sk, body of nuchal skeleton. m, mouth. th, throat. tb, tongue-bars. tc, trunk coelom. Balanoglossus 3.png
Structure of anterior end a, Arrow from proboscis-cavity (pc) passing to left of pericardium (per) and out through proboscis pore-canal. b , arrow from central canal of neurochord (cnc) passed out through anterior neuropore. b , ditto; through posterior neuropore. c, arrow intended to pass from 1st gill-pouch through collar pore-canal into collar-coelom (cc). cts, posterior limit of collar. dv, dorsal vessel passing into central sinus (bs). ev, efferent vessel passing into ventral vessel (vv). epr, epiphysial tubes. st, stomochord. vs, ventral septum of proboscis. sk, body of nuchal skeleton. m, mouth. th, throat. tb, tongue-bars. tc, trunk coelom.
Acorn worm on the ocean floor Expn7526 (38827990315).jpg
Acorn worm on the ocean floor

Most acorn worms range from 9 to 45 centimetres (3.5 to 17.7 in) in length, with the largest species, Balanoglossus gigas , reaching 1.5 metres (5 ft) or more. The body is made up of three main parts: an acorn-shaped proboscis, a short fleshy collar that lies behind it, and a long, worm-like trunk. The creature's mouth is located at the collar behind the proboscis. [16]

The skin is covered with cilia as well as glands that secrete mucus. Some produce a bromide compound that gives them a medicinal smell and might protect them from bacteria and predators. Acorn worms move only sluggishly, using ciliary action and peristalsis of the proboscis. [16]

Digestive system

Many acorn worms are detritus feeders, eating sand or mud and extracting organic detritus. Others feed on organic material suspended in the water, which they can draw into the mouth using the cilia on the gill bars. [17] Research indicates that the rate of feeding of acorn worms that are detritus feeders is dependent on food availability and flow rate. [18] A groove lined with cilia lies just in front of the mouth and directs suspended food into the mouth and may allow the animal to taste. [16]

The mouth cavity is tubular, with a narrow diverticulum or stomochord extending up into the proboscis. This diverticulum was once thought to be homologous with the notochord of chordates, hence the name "hemichordate" for the phylum. The mouth opens posteriorly into a pharynx with a row of gill slits along either side. The remainder of the digestive system consists of an oesophagus and intestine; there is no stomach. [16]

In some families there are openings in the dorsal surface of the oesophagus connecting to the external surface, through which water from the food can be squeezed, helping to concentrate it. Digestion occurs in the intestine, with food material being pulled through by cilia, rather than by muscular action. [16]

Acorn worms breathe by drawing in oxygenated water through their mouth. The water then flows out the animal's gills which are on its trunk. Thus, the acorn worm breathes about the same way as fish.

Circulatory system

Acorn worms have an open circulatory system, in which the blood flows through the tissues sinuses. A dorsal blood vessel in the mesentery above the gut delivers blood to a sinus in the proboscis that contains a muscular sac acting as a heart. Unlike the hearts of most other animals, however, this structure is a closed fluid-filled vesicle whose interior does not connect directly to the blood system. Nonetheless, it does regularly pulsate, helping to push blood through the surrounding sinuses. [16]

From the central sinus in the collar, blood flows to a complex series of sinuses and peritoneal folds in the proboscis. This set of structures is referred to as a glomerulus and may have an excretory function, since acorn worms otherwise have no defined excretory system. From the proboscis, blood flows into a single blood vessel running underneath the digestive tract, from which smaller sinuses supply blood to the trunk, and back into the dorsal vessel. [16]

The blood of acorn worms is colourless and acellular. [16]

Respiratory system

Acorn worms continually form new gill slits as they grow in size, with some older individuals of species like Balanoglossus aurantiacus having more than a hundred on each side. The microscopic species Meioglossus psammophilus has just a single gill slit. The gills in some acorn worms have cartilaginous support structures. [19] Each slit consists of a branchial chamber opening to the pharynx through a U-shaped cleft and to the exterior through a dorso-lateral pore (see diagram below). Cilia push water through the slits, maintaining a constant flow. The tissues surrounding the slits are well supplied with blood sinuses. [16]

Nervous system

A plexus of nerves lies underneath the skin, and is concentrated into both dorsal and ventral nerve cords. While the ventral cord runs only as far as the collar, the dorsal cord reaches into the proboscis, and is partially separated from the epidermis in that region. This part of the dorsal nerve cord is often hollow, and may well be homologous with the brain of vertebrates. In acorn worms, it seems to be primarily involved with coordinating muscular action of the body during burrowing and crawling. [16]

Acorn worms have no eyes, ears or other special sense organs, except for the ciliary organ in front of the mouth, which appears to be involved in filter feeding and perhaps taste (3). There are, however, numerous nerve endings throughout the skin. [16]

Skeletal system

Acorn worms have a Y-shaped nuchal skeleton that starts their proboscis and collar on their ventral side. The length of the horns of the nuchal skeleton varies between species. [20]

Similarities to chordates

Acorn worms have a circulatory system with a heart that also functions as a kidney.[ citation needed ] Acorn worms have gill-like structures that they use for breathing, similar to the gills of primitive fish. Therefore, acorn worms are sometimes said to be a link between classical invertebrates and vertebrates. Some also have a postanal tail which may be homologous to the post-anal tail of vertebrates. An interesting trait is that its three-section body plan is no longer present in the vertebrates, except for the anatomy of the frontal neural tube, later developed into a brain which is divided into three main parts. This means some of the original anatomy of the early chordate ancestors is still present even if it is not always visible.

One theory is that the three-part body originates from an early common ancestor of all the deuterostomes, and maybe even from a common bilateral ancestor of both the deuterostomes and protostomes.[ citation needed ] Studies have shown that the gene expression in the embryo share three of the same signaling centers that shape the brains of all vertebrates, but instead of taking part in the formation of their neural system, [21] they are controlling the development of the different body regions. [22]

Phylogeny

The internal relationships within the Enteropneusta are shown below. The tree is based on 16S +18S rRNA sequence data and phylogenomic studies from multiple sources. [23] [24]

Hemichordata

Lifestyle

Meioglossus psammophilus .

Acorn worms are rarely seen by humans because of their lifestyle. They live in U-shaped burrows on the sea-bed, from the shoreline down to a depth of 10,000 ft. (3,050 m). The worms lie there with the proboscis sticking out of one opening in the burrow. Acorn worms are generally slow burrowers.

To obtain food, many acorn worms swallow sand or mud that contains organic matter and microorganisms in the manner of earthworms (this is known as deposit feeding). At low tide, they stick out their rear ends at the surface and excrete coils of processed sediments (casts).

Another method that some acorn worms use to obtain food is to collect suspended particles of organic matter and microbes from the water. This is known as suspension feeding. [17]

Reproduction

Acorn worms are dioecious, having separate biological sexes, although at least some species are also capable of asexual reproduction in the form of fragmentation. [25] They have paired gonads, which lie close to the pharynx and release the gametes through a small pore near to the gill slits. The female lays a large number of eggs embedded in a gelatinous mass of mucus, which are then externally fertilized by the male before water currents break up the mass and disperse the individual eggs. [16]

Acorn worm life cycle by M. Singh AcornWormCycle.jpg
Acorn worm life cycle by M. Singh

In most species, the eggs hatch into planktonic larvae with elongated bodies covered in cilia. In some species, these develop directly into adults, but in others, there is a free-swimming intermediate stage referred to as a tornaria larva. These are very similar in appearance to the bipinnaria larvae of starfishes, with convoluted bands of cilia running around the body. Since the embryonic development of the blastula within the egg is also very similar to that of echinoderms, this suggests a close phylogenetic link between the two groups. [16]

After a number of days or weeks, a groove begins to form around the larval midsection, with the anterior portion eventually destined to become the proboscis, while the remainder forms the collar and trunk. The larvae eventually settle down and change into tiny adults to take on the burrowing lifestyle. A few species, such as Saccoglossus kowalevskii , lack even the planktonic larval stage, hatching directly as miniature adults. [16]

Related Research Articles

<span class="mw-page-title-main">Chordate</span> Phylum of animals having a dorsal nerve cord

A chordate is a deuterostomic bilaterial animal belonging to the phylum Chordata. All chordates possess, at some point during their larval or adult stages, five distinctive physical characteristics (synapomorphies) that distinguish them from other taxa. These five synapomorphies are a notochord, a hollow dorsal nerve cord, an endostyle or thyroid, pharyngeal slits, and a post-anal tail.

<span class="mw-page-title-main">Hemichordate</span> Phylum of marine deuterostome animals

Hemichordata is a phylum which consists of triploblastic, eucoelomate, and bilaterally symmetrical marine deuterostome animals, generally considered the sister group of the echinoderms. They appear in the Lower or Middle Cambrian and include two main classes: Enteropneusta, and Pterobranchia. A third class, Planctosphaeroidea, is known only from the larva of a single species, Planctosphaera pelagica. The class Graptolithina, formerly considered extinct, is now placed within the pterobranchs, represented by a single living genus Rhabdopleura.

<span class="mw-page-title-main">Vetulicolia</span> Extinct Cambrian group of animals

Vetulicolia is a possibly paraphyletic phylum of bilaterian animals encompassing several extinct species belonging to the Cambrian period. The phylum was created by Degan Shu and his research team in 2001, and named after Vetulicola cuneata, the first species of the phylum described in 1987.

<span class="mw-page-title-main">Graptolite</span> Subclass of Pterobranchs in the phylum Hemichordata

Graptolites are a group of colonial animals, members of the subclass Graptolithina within the class Pterobranchia. These filter-feeding organisms are known chiefly from fossils found from the Middle Cambrian through the Lower Carboniferous (Mississippian). A possible early graptolite, Chaunograptus, is known from the Middle Cambrian. Recent analyses have favored the idea that the living pterobranch Rhabdopleura represents an extant graptolite which diverged from the rest of the group in the Cambrian.

Pikaia gracilens is an extinct, primitive chordate animal known from the Middle Cambrian Burgess Shale of British Columbia. Described in 1911 by Charles Doolittle Walcott as an annelid, and in 1979 by Harry B. Whittington and Simon Conway Morris as a chordate, it became "the most famous early chordate fossil", or "famously known as the earliest described Cambrian chordate". It is estimated to have lived during the latter period of the Cambrian explosion. Since its initial discovery, more than a hundred specimens have been recovered.

<i>Balanoglossus</i> Genus of ocean-dwelling acorn worm

Balanoglossus is a genus of ocean-dwelling acorn worms. It has zoological importance because, being hemichordates, they are an "evolutionary link" between invertebrates and vertebrates. Balanoglossus specimens are deuterostomes, and resemble the sea squirts in that they possess branchial openings, or "gill slits". They have notochords and stomochords but no dorsal nerve cord.

<span class="mw-page-title-main">Pterobranchia</span> Class of hemichordates

Pterobranchia, members of which are often called pterobranchs, is a class of small worm-shaped animals. They belong to the Hemichordata, and live in secreted tubes on the ocean floor. Pterobranchia feed by filtering plankton out of the water with the help of cilia attached to tentacles. There are about 25 known living pterobranch species in three genera, which are Rhabdopleura, Cephalodiscus, and Atubaria. On the other hand, there are several hundred extinct genera, some of which date from the Cambrian Period.

<span class="mw-page-title-main">Pharyngeal slit</span> Repeated openings that appear along the pharynx of chordates

Pharyngeal slits are filter-feeding organs found among deuterostomes. Pharyngeal slits are repeated openings that appear along the pharynx caudal to the mouth. With this position, they allow for the movement of water in the mouth and out the pharyngeal slits. It is postulated that this is how pharyngeal slits first assisted in filter-feeding, and later, with the addition of gills along their walls, aided in respiration of aquatic chordates. These repeated segments are controlled by similar developmental mechanisms. Some hemichordate species can have as many as 200 gill slits. Pharyngeal clefts resembling gill slits are transiently present during the embryonic stages of tetrapod development. The presence of pharyngeal arches and clefts in the neck of the developing human embryo famously led Ernst Haeckel to postulate that "ontogeny recapitulates phylogeny"; this hypothesis, while false, contains elements of truth, as explored by Stephen Jay Gould in Ontogeny and Phylogeny. However, it is now accepted that it is the vertebrate pharyngeal pouches and not the neck slits that are homologous to the pharyngeal slits of invertebrate chordates. Pharyngeal arches, pouches, and clefts are, at some stage of life, found in all chordates. One theory of their origin is the fusion of nephridia which opened both on the outside and the gut, creating openings between the gut and the environment.

<i>Vetulicola</i> Fossil genus of marine animal

Vetulicola is an extinct genus of marine animal discovered from the Cambrian of China. It is the eponymous member of the enigmatic phylum Vetulicolia, which is of uncertain affinities but may belong to the deuterostomes. The name was derived from Vetulicola cuneata, the first species described by Hou Xian-guang in 1987 from the Lower Cambrian Chiungchussu Formation in Chengjiang, China.

<span class="mw-page-title-main">Deuterostome</span> Superphylum of bilateral animals

Deuterostomes are bilaterian animals of the superphylum Deuterostomia, typically characterized by their anus forming before the mouth during embryonic development. Deuterostomia is further divided into 4 phyla: Chordata, Echinodermata, Hemichordata, and the extinct Vetulicolia known from Cambrian fossils. The extinct clade Cambroernida is also thought to be a member of Deuterostomia.

<i>Saccoglossus</i> Genus of marine worm-like animals

Saccoglossus is a genus of acorn worm. It is the largest genus in the Enteropneusta class, with 20 species.

In evolutionary developmental biology, inversion refers to the hypothesis that during the course of animal evolution, the structures along the dorsoventral (DV) axis have taken on an orientation opposite that of the ancestral form.

The calcichordate hypothesis holds that each separate lineage of chordate evolved from its own lineage of mitrate, and thus the echinoderms and the chordates are sister groups, with the hemichordates as an out-group.

<span class="mw-page-title-main">Harrimaniidae</span> Family of marine worm-like animals

Harrimaniidae is a basal family of acorn worms. A taxonomic revision was undertaken in 2010, and a number of new genera and species found in the Eastern Pacific were described. In this family the development is direct without tornaria larva, and circular muscle fibers in their trunk is missing. There is some indication that Stereobalanus may be a separate basal acorn worm lineage, sister to all remaining acorn worms.

<i>Ooedigera</i> Ovoid Cambrian animal with a bulbous tail

Ooedigera peeli is an extinct vetulicolian from the Early Cambrian of North Greenland. The front body was flattened horizontally, oval-shaped, likely bearing a reticulated or anastomosing pattern, and had 5 evenly-spaced gill pouches along the midline. The tail was also bulbous and flattened horizontally, but was divided into 7 plates connected by flexible membranes, allowing movement. Ooedigera likely swam by moving side-to-side like a fish. It may have lived in an oxygen minimum zone alongside several predators in an ecosystem based on chemosynthetic microbial mats, and was possibly a deposit or filter feeder living near the seafloor.

Harrimania planktophilus is a marine acorn worm in the family Harrimaniidae. It lives in a burrow in sediment on the sea floor. It is only known from western Canada and was first described by Cameron in 2002. The species name is from the Greek and translates as "lover of plankton".

Yoda purpurata is a species of acorn worm discovered 2.5 km below the surface of the Atlantic ocean, and was the first of the genus Yoda found. Ranging from 12 to 19 cm in length, it was named after the fictional character Yoda from the Star Wars franchise. It was the first known hermaphroditic member within the phylum. The other known hermaphroditic member of the phylum is Yoda demiankoopi discovered in 2021.

<span class="mw-page-title-main">Torquaratoridae</span> Family of enteropneust in the hemichordate phylum

Torquaratoridae is a family of acorn worms (Hemichordata) that lives in deep waters between 350 and 4000 meters. They can grow up to three feet in length and have semitransparent gelatinous bodies, often brightly colored.

Saccoglossus bromophenolosus is a species of acorn worm occurring in the northwestern Atlantic Ocean and the northeastern Pacific Ocean. It grows to a length of about 20 cm (8 in) and lives in a burrow in soft sediment in the intertidal and subtidal zones. The scientific name refers to 2,4-dibromophenol, a secondary metabolite present in this worm.

The Cambrian chordates are an extinct group of animals belonging to the phylum Chordata that lived during the Cambrian, between 538 and 485 million years ago. The first Cambrian chordate known is Pikaia gracilens, a lancelet-like animal from the Burgess Shale in British Columbia, Canada. The discoverer, Charles Doolittle Walcott, described it as a kind of worm (annelid) in 1911, but it was later identified as a chordate. Subsequent discoveries of other Cambrian fossils from the Burgess Shale in 1991, and from the Chengjiang biota of China in 1991, which were later found to be of chordates, several Cambrian chordates are known, with some fossils considered as putative chordates.

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