Hemichordata
Encyclopedia
Hemichordata is a phylum of marine deuterostome
animal
s, generally considered the sister group of the echinoderm
s. They date back to the Lower or Middle Cambrian
and include two main classes: Enteropneusta (acorn worms), and Pterobranchia
. A third class, Planctosphaeroidea, is known only from the larva of a single species. The extinct class Graptolithina is closely related to the pterobranchs.
Acorn worms are solitary worm-shaped organisms. They generally live in burrows and are deposit feeders, but some species are pharyngeal filter feeders. Pterobranchs are colonial filter-feeding organisms that live in a collagenous tubular structure called a coenecium.
is divided into three parts: the anterior prosome, the intermediate mesosome, and the posterior metasome.
The body of acorn worms is worm-shaped and divided into an anterior proboscis, an intermediate collar, and a posterior trunk. The proboscis is a muscular and ciliated organ used in locomotion and in the collection and transport of food particles. The mouth is located between the proboscis and the collar. The trunk is the longest part of the animal. It contains the pharynx, which is perforated with gill slits (or pharyngeal slits), the esophagus, a long intestine, and a terminal anus. It also contains the gonads.
The prosome of pterobranchs is specialized into a muscular and ciliated cephalic shield used in locomotion and in secreting the coenecium. The mesosome extends into one pair (in the genus Rhabdopleura) or several pairs (in the genus Cephalodiscus) of tentaculated arms used in filter feeding. The metasome, or trunk, contains a looped digestive tract, gonads, and extends into a contractile stalk that connects individuals to the other members of the colony, produced by asexual budding. In the genus Cephalodiscus, asexually produced individuals stay attached to the contractile stalk of the parent individual until completing their development. In the genus Rhabdopleura, zooids are permanently connected to the rest of the colony via a common stolon system.
They have a diverticulum
of the foregut called a stomochord
, previously thought to be related with the chordate notochord
, but this is most likely the result of convergent evolution
rather than homology
. A hollow neural tube
exists among some species (at least in early life), probably a primitive trait they share with the common ancestor of chordata
and the rest of the deuterostomes.
s both in the vegetal and the animal pole. The fourth division occurs mainly in blastomeres in the animal pole, which divide transversally as well as equally to make eight blastomeres. The four vegetal blastomeres divide equatorially but unequally and they give rise to four big macromeres and four smaller micromeres. Once this fourth division has occurred the embryo has reached a 16 cell stage. P. flava has a 16 cell embryo with four vegetal micromeres, eight animal mesomeres and 4 larger macromeres. Further divisions occur until P. flava finishes the blastula
stage and goes on to gastrulation
. The animal mesomeres of P. flava go on to give rise to the larva’s ectoderm
, animal blastomere
s also appear to give rise to these structures though the exact contribution varies from embryo to embryo. The macromeres give rise to the posterior larval ectoderm and the vegetal micromeres give rise to the internal endomesodermal tissues. Studies done on the potential of the embryo at different stages have shown that at both the two and four cell stage of development P. flava blastomeres can go on to give rise to a tornaria larvae, so fates of these embryonic cells don’t seem to be established till after this stage.
, commonly called acorn worms, and the Pterobranchia
, which may include the graptolite
s. A third class, Planctosphaeroidea, is proposed based on a single species known only from larvae. The phylum contains about 80 living species. Hemichordata appears to be sister to the Echinodermata as Ambulacraria; Xenoturbellida may be basal to that grouping. Pterobranchia may be derived from within Enteropneusta, making Enteropneusta paraphyletic.
Deuterostome
Deuterostomes are a superphylum of animals. They are a subtaxon of the Bilateria branch of the subregnum Eumetazoa, and are opposed to the protostomes...
animal
Animal
Animals are a major group of multicellular, eukaryotic organisms of the kingdom Animalia or Metazoa. Their body plan eventually becomes fixed as they develop, although some undergo a process of metamorphosis later on in their life. Most animals are motile, meaning they can move spontaneously and...
s, generally considered the sister group of the echinoderm
Echinoderm
Echinoderms are a phylum of marine animals. Echinoderms are found at every ocean depth, from the intertidal zone to the abyssal zone....
s. They date back to the Lower or Middle Cambrian
Cambrian
The Cambrian is the first geological period of the Paleozoic Era, lasting from Mya ; it is succeeded by the Ordovician. Its subdivisions, and indeed its base, are somewhat in flux. The period was established by Adam Sedgwick, who named it after Cambria, the Latin name for Wales, where Britain's...
and include two main classes: Enteropneusta (acorn worms), and Pterobranchia
Pterobranchia
Pterobranchia is a clade 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 30 known living species in the group.The...
. A third class, Planctosphaeroidea, is known only from the larva of a single species. The extinct class Graptolithina is closely related to the pterobranchs.
Acorn worms are solitary worm-shaped organisms. They generally live in burrows and are deposit feeders, but some species are pharyngeal filter feeders. Pterobranchs are colonial filter-feeding organisms that live in a collagenous tubular structure called a coenecium.
Structure
The body plan of hemichordates is characterized by a tripartite organization. The anteroposterior axisAnatomical terms of location
Standard anatomical terms of location are designations employed in science that deal with the anatomy of animals to avoid ambiguities that might otherwise arise. They are not language-specific, and thus require no translation...
is divided into three parts: the anterior prosome, the intermediate mesosome, and the posterior metasome.
The body of acorn worms is worm-shaped and divided into an anterior proboscis, an intermediate collar, and a posterior trunk. The proboscis is a muscular and ciliated organ used in locomotion and in the collection and transport of food particles. The mouth is located between the proboscis and the collar. The trunk is the longest part of the animal. It contains the pharynx, which is perforated with gill slits (or pharyngeal slits), the esophagus, a long intestine, and a terminal anus. It also contains the gonads.
The prosome of pterobranchs is specialized into a muscular and ciliated cephalic shield used in locomotion and in secreting the coenecium. The mesosome extends into one pair (in the genus Rhabdopleura) or several pairs (in the genus Cephalodiscus) of tentaculated arms used in filter feeding. The metasome, or trunk, contains a looped digestive tract, gonads, and extends into a contractile stalk that connects individuals to the other members of the colony, produced by asexual budding. In the genus Cephalodiscus, asexually produced individuals stay attached to the contractile stalk of the parent individual until completing their development. In the genus Rhabdopleura, zooids are permanently connected to the rest of the colony via a common stolon system.
They have a diverticulum
Diverticulum
A diverticulum is medical or biological term for an outpouching of a hollow structure in the body. Depending upon which layers of the structure are involved, they are described as being either true or false....
of the foregut called a stomochord
Stomochord
The stomochord is a flexible, hollow tube found in hemichordates. Stomochords were initially considered a variant of a primitive notochord, but are now recognized to not share histological composition to that of the notochord found in chordates....
, previously thought to be related with the chordate notochord
Notochord
The notochord is a flexible, rod-shaped body found in embryos of all chordates. It is composed of cells derived from the mesoderm and defines the primitive axis of the embryo. In some chordates, it persists throughout life as the main axial support of the body, while in most vertebrates it becomes...
, but this is most likely the result of convergent evolution
Convergent evolution
Convergent evolution describes the acquisition of the same biological trait in unrelated lineages.The wing is a classic example of convergent evolution in action. Although their last common ancestor did not have wings, both birds and bats do, and are capable of powered flight. The wings are...
rather than homology
Homology (biology)
Homology forms the basis of organization for comparative biology. In 1843, Richard Owen defined homology as "the same organ in different animals under every variety of form and function". Organs as different as a bat's wing, a seal's flipper, a cat's paw and a human hand have a common underlying...
. A hollow neural tube
Neural tube
In the developing vertebrate, the neural tube is the embryo's precursor to the central nervous system, which comprises the brain and spinal cord...
exists among some species (at least in early life), probably a primitive trait they share with the common ancestor of chordata
Chordate
Chordates are animals which are either vertebrates or one of several closely related invertebrates. They are united by having, for at least some period of their life cycle, a notochord, a hollow dorsal nerve cord, pharyngeal slits, an endostyle, and a post-anal tail...
and the rest of the deuterostomes.
Development
The hemichordates give us the closest extant phylogenetic relative between the chordates and other invertebrates. Thus these marine worms, described to be the sister group of such animals as sea urchins, are of great importance to the scientific community interested in knowing the origins of chordate development. There are several species classified as hemichordates and there exists a moderate diversity of embryological development between these species. Hemichordates are classically known to develop in two ways both directly and indirectly. Hemichordates are a phylum composed of two classes the enteropneusts and the pterobranchs, both are forms of marine worm. The enteropneusts have two developmental strategies direct and indirect development. The indirect way of development is known to end in an extended pelagic plankotrophic tornaria larval stage, which means that this hemichordate exists in a larval stage that feeds on plankton before turning into an adult worm. Those species that are direct developing bypass this prolonged larval stage and develop directly into an adult worm. The following details the development of two popularly studied species of the hemichordata phylum Saccoglossus kowalevskii and Ptychodera flava. Saccoglossus kowalevskii is a direct developer and Ptychodera flava is an indirect developer. Most of what has been detailed in Hemichordate development has come from hemichordates that develop directly.Ptychodera flava
P. flava’s early cleavage pattern is similar to that of S. kowalevskii. The first and second cleavages from the single cell zygote of P. flava are equal cleavages, are orthogonal to each other and both include the animal and vegetal poembryo has four blastomereBlastomere
A blastomere is a type of cell produced by division of the egg after fertilization.- References :* "Blastomere." Stedman's Medical Dictionary, 27th ed. . ISBN 0-683-40007-X...
s both in the vegetal and the animal pole. The fourth division occurs mainly in blastomeres in the animal pole, which divide transversally as well as equally to make eight blastomeres. The four vegetal blastomeres divide equatorially but unequally and they give rise to four big macromeres and four smaller micromeres. Once this fourth division has occurred the embryo has reached a 16 cell stage. P. flava has a 16 cell embryo with four vegetal micromeres, eight animal mesomeres and 4 larger macromeres. Further divisions occur until P. flava finishes the blastula
Blastula
The blastula is a hollow sphere of cells formed during an early stage of embryonic development in animals . The blastula is created when the zygote undergoes the cell division process known as cleavage. The blastula is preceded by the morula and is followed by the gastrula in the developmental...
stage and goes on to gastrulation
Gastrulation
Gastrulation is a phase early in the embryonic development of most animals, during which the single-layered blastula is reorganized into a trilaminar structure known as the gastrula. These three germ layers are known as the ectoderm, mesoderm, and endoderm.Gastrulation takes place after cleavage...
. The animal mesomeres of P. flava go on to give rise to the larva’s ectoderm
Ectoderm
The "ectoderm" is one of the three primary germ cell layers in the very early embryo. The other two layers are the mesoderm and endoderm , with the ectoderm as the most exterior layer...
, animal blastomere
Blastomere
A blastomere is a type of cell produced by division of the egg after fertilization.- References :* "Blastomere." Stedman's Medical Dictionary, 27th ed. . ISBN 0-683-40007-X...
s also appear to give rise to these structures though the exact contribution varies from embryo to embryo. The macromeres give rise to the posterior larval ectoderm and the vegetal micromeres give rise to the internal endomesodermal tissues. Studies done on the potential of the embryo at different stages have shown that at both the two and four cell stage of development P. flava blastomeres can go on to give rise to a tornaria larvae, so fates of these embryonic cells don’t seem to be established till after this stage.
Saccoglossus kowalevskii
Eggs of S. kowalevskii are oval in shape and become spherical in shape after fertilization. The first cleavage occurs from the animal to the vegetal pole and usually is equal though very often can also be unequal. The second cleavage to reach the embryos four cell stage also occurs from the animal to the vegetal pole in an approximately equal fashion though like the first cleavage it’s possible to have an unequal division. The eight cell stage cleavage is latitudinal; so that each cell from the four cell stage goes on to make two cells. The fourth division occurs first in the cells of the animal pole which end up making 8 blastomeres(mesomeres) that are not radially symmetric, then the four vegetal pole blastomeres divide to make a level of four large blastomeres (macromeres) and four very small blastomeres (micromeres). The fifth cleavage occurs first in the animal cells and then in the vegetal cells to give a 32 cell blastomere. The sixth cleavage occurs in a similar order and completes a 64 cell stage, finally the seventh cleavage marks the end of the cleavage stage with a blastula with 128 blastomeres. This structure goes on to go thru gastrulation movements which will determine the body plan of the resulting gill slit larva, this larva will ultimately give rise to the marine acorn wormGenetic control of dorsal-ventral hemichordate patterning
Much of the genetic work done on hemichordates has been done to make comparison with chordates, so it is obvious that many of the genetic markers identified in this group are also found in chordates or are homologous to chordates in some way. Studies of this nature have been done particularly on S. kowalevskii, and like chordates S. kowalevskii has dorsalizing bmp-like factors such as bmp 2/4 which is homologous to Drosophila’s decapentaplegic dpp. The expression of bmp2/4 begins at the onset of gastrulation on the ectodermal side of the embryo, and as grastulation progresses its expression is narrowed down to the dorsal midline but is not expressed in the post anal tail. The bmp antagonist chordin is also expressed in the endoderm of gastrulating S. kowalevskii. Besides these well known dorsalizing factors, further molecules known to be involved in dorsal ventral patterning are also present in S. kowalevskii, such as a netrin that groups with netrin gene class 1 and 2. Netrin is important in patterning of the neural system in chordates, as well as is the molecule Shh, but S. kowalevskii was only found to have one hh gene and it appears to be expressed in a region that is uncommon to where it is usually expressed in developing chordates along the ventral midline.Classification
Hemichordata are divided into two classes: the EnteropneustaAcorn worm
The Acorn worms or Enteropneusta are a hemichordate class of invertebrates, closely related to the echinoderms. There are about 90 species of acorn worm in the world, the main species for research being Saccoglossus kowaleskii....
, commonly called acorn worms, and the Pterobranchia
Pterobranchia
Pterobranchia is a clade 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 30 known living species in the group.The...
, which may include the graptolite
Graptolite
Graptolithina is a class in the animal phylum Hemichordata, the members of which are known as Graptolites. Graptolites are fossil colonial animals known chiefly from the Upper Cambrian through the Lower Carboniferous...
s. A third class, Planctosphaeroidea, is proposed based on a single species known only from larvae. The phylum contains about 80 living species. Hemichordata appears to be sister to the Echinodermata as Ambulacraria; Xenoturbellida may be basal to that grouping. Pterobranchia may be derived from within Enteropneusta, making Enteropneusta paraphyletic.
External links
- Cameron, C. B. Evolution and classification of Hemichordata
- Dr. Billie J. Swalla on the WEB "Development and Evolution of Hemichordates"
- Dr. Ken Halanych "Phylogeny and Evolution of Hemichordates"
- Dr. Chris Lowe "Genomics and Development of Saccoglossus kowalevskii"
- At the Tree of Life