Difference between revisions of "Hemichordata" - New World Encyclopedia

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'''Hemichordata''' is a small phylum of worm-shaped, marine [[invertebrate]]s.  
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'''Hemichordata''' is a small phylum of worm-shaped, marine [[invertebrate]]s. Hemichordates are [[symmetry (biology)|bilaterially symmetrical]] and their bodies are divided into three sections (Protosome or proboscis, collar, and trunk). The musculature in their gut is very poorly developed, and food is mostly transported through it by using the cilia that covers the inside.
  
 +
Hemichordates are considered a sister group of the [[chordate]]s and [[echinoderm]]s. They were once considered part of Chordata, but lack a true notochord. A hollow nerve cord, or notochord, is found in all chordates, even tunicates, and some hemichordates also seem to have a primitive form of notochord (at least in early life). In the embryonic stage, this tubular nerve cord looks like the hollow nerve cord of chordates. Both Hemichordata and Chordata have gill slits, and primitive [[fossil]] echinoderms also show signs of gill slits.
  
As with Phylum Chordata, Echinodermata, and Hemichordata, Hemichordata are deuterostomes. As deuterostomes, they
+
As with the Chordata and Echinodermata phyla, Hemichordata are ''deuterostomes''. As deuterostomes, they have true coeloms (body cavities), with the coelom forming through enterocoely (the mesoderm forms as evaginations of the developed gut that pinch off, forming the coelom); the first opening becomes the anus rather than the mouth as in protostomes; and the early divisions of the zygote occurs parallel or perpendicular to the polar axis (radial cleavage).
  
They are deuterostomes, having a true coelom that has
+
Hemichordata includes about 100 living species. These are widely distributed, being found both in shallow costal waters and in the deeper sea, and are benthic (live on the sea floor), at least in their adult form (Ramel 2005). Hemichordates range from only a few millimeters long to the ''Balanoglossus gigas'', which reaches 1.5 meters (4.7 feet) (Ramel 2005).
  
Note: the first opening (the blastopore) becomes the anus, while in protostomes it becomes the mouth.
+
==Hemichordata classes==
  
Note: In deuterostomes the mesoderm forms as evaginations of the developed gut that pinch off, forming the coelom. This is called enterocoely.
+
Hemichordates date back to the Lower or Middle [[Cambrian]] and include an important class of [[fossil]]s called graptolites, most of which became extinct in the [[Carboniferous]].
  
Note: Both the Hemichordata and Chordata have gill slits, and primitive fossil echinoderms also show signs of gill slits. A hollow nerve cord is found in all chordates, even tunicates (even if it disappears in the adults). Some hemichordates also have a tubular nerve cord. In early embryonic stage it looks like the hollow nerve cord of chordates
+
Hemichordata is divided into two classes: the [[#Enteropneusta (Acorr worms)|Enteropneusta]], commonly called acorn worms, and the [[#Pterobranchia|Pterobranchia]], which may include the graptolites.  
  
Note: In both deuterostomes and protostomes, a zygote first develops into a hollow ball of cells, called a blastula. In deuterostomes, the early divisions occur parallel or perpendicular to the polar axis. This is called radial cleavage
+
A third class, Planctosphaeroidea, is proposed based on a single species known only from larvae.
  
Note: Phylum Chordata (vertebrates and their kin)
+
One of the suggestions are that the pterobranchs are more basal deuterostomes, while the enteropneusts are an early offshoot of the lineage that lead to [[Chordata]].
Phylum Echinodermata (starfishes, sea urchins, sea cucumbers, etc.)
 
Phylum Hemichordata (acorn worms and possibly graptolites)
 
Phylum Xenoturbellida (2 species of worm-like animals from the Baltic Sea)
 
 
 
 
 
[[deuterostome]] [[animal]]s, generally considered the sister group of the [[chordate]]s.
 
 
 
They date back to the Lower or Middle [[Cambrian]] and include an important class of [[fossil]]s called [[graptolite]]s, most of which became extinct in the [[Carboniferous]]. They seem to have a primitive form of [[notochord]], but this is most likely the result of [[convergent evolution]]. 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. The musculature in their gut is very poorly developed, and food is mostly transported through it by using the cilia that covers its inside. They are tradtionally considered as a sister group to the [[echinoderms]].
 
 
 
Hemichordata are divided into two classes: the [[Enteropneusta]], commonly called acorn worms, and the [[Pterobranchia]], which may include the graptolites. A third class, [[Planctosphaeroidea]], is proposed based on a single species known only from larvae. The phylum contains about 100 living species. The exact taxonomic position of hemichordata and whether the group is [[monophyletic]] is currently under debate. One of the suggestions are that the pterobranchs are more basal deuterostomes, while the enteropneusts are an early offshoot of the lineage who are leading to [[Chordata]].
 
{{wikispecies|Hemichordata}}
 
==References==
 
* [http://cluster3.biosci.utexas.edu/faculty/cameronc/Hemichordate.Species.html Cameron, C. B. ''Classification of the extant hemichordata'']
 
* [http://tolweb.org/tree?group=Hemichordata&contgroup=Deuterostomia At the Tree of Life]
 
  
 
==Enteropneusta (Acorn worms)==
 
==Enteropneusta (Acorn worms)==
The '''Acorn worms''' or '''Enteropneusta''' are a [[Hemichordata|hemichordate]] class of [[invertebrate]]s. Acorn worms are classified in the phylum [[Hemichordata]], closely related to the [[chordate]]s. There are about 70 species of acorn worm in the world.
+
The '''Acorn worms''' or '''Enteropneusta''' include about 70 known species. As with all hemichordates, acorn worms are benthic and may be deposit feeders (substrate feeders) or suspension feeders (filter feeders). Substrate feeders, like ''Balanoglossus clavigerus'', are generally larger than filter feeders (Ramel 2005).
 
 
All species are [[Fauna (animals)#Infauna|infaunal]] [[benthos]] that either may be [[deposit feeder]]s or [[suspension feeder]]s. Some of these worms may grow to be very long; one particular species may reach a length of 2.5 meters (almost eight feet), although most acorn worms are much, much smaller.  
 
  
One genus, ''[[Balanoglossus]]'', is also known as the tongue [[worm]].  
+
The genus ''Balanoglossus'' is also known as the tongue worm.  
  
 
===Anatomy===
 
===Anatomy===
The Acorn worm's body is [[cylinder (geometry)|cylindrical]]. The body is made up of three main parts: the acorn-shaped proboscis, a short fleshy collar that lies behind it, and the long trunk, which is the rest of the body. The creature's [[mouth]] is located at the collar behind the proboscis. One theory is that this 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.
+
The Acorn worm's body is cylindrical and made up of three main parts: the acorn-shaped proboscis, a short fleshy collar that lies behind it, and the long trunk, which is the rest of the body. The creature's mouth is located at the collar behind the proboscis. One theory is that this 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.
  
The skin is covered with [[cilium|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.
+
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 by cilia movements and body contractions.
  
Acorn worms move by cilia movements and body contractions.
+
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]].
  
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.
+
====Similarities to Chordates====
  
====Similarities to Chordates====
+
Acorn worms are considered more highly specialized and advanced than other similarly shaped worm-like creatures. They have a circulatory system with a [[heart]] that also functions as a [[kidney]]. Acorn worms have gill-like structures that they use for breathing, similar to the gills of primitive fish. Hence, acorn worms are sometimes said to be a link between classical [[invertebrate]]s and [[vertebrate]]s.
  
Acorn worms are considered more highly specialised and advanced than other similarly shaped worm-like creatures. They have a circulatory system with a heart that also functions as a kidney. Acorn worms have  gill-like structures that they use for breathing, similar to the gills of primitive fish. Hence, acorn worms are sometimes said to be a link between classical invertebrates and vertebrates. Some also have a postanal tail which sometimes show weak signs of segmentation. An interesting trait is that its three-section body plan is no longer present in the vertebrates, except from 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.
+
Some also have a post-anal tail that sometimes show weak signs of segmentation. An interesting trait is that its three-section body plan is not present in the vertebrates, except from the anatomy of the frontal neural tube, later developed into a [[brain]] that is divided into three main parts. This means some of the original anatomy of the early chordate ancestors may still present even if it is not always visible.
  
 
===Lifestyle===
 
===Lifestyle===
  
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.
+
Acorn worms are rarely seen by [[human]]s 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).  
+
To obtain [[food]], many acorn worms swallow sand or mud that contains organic matter and microorganisms in the manner of [[earthworm]]s (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). They rarely leave their burrows, which may have several openings (Ramel 2005).
  
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.
+
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. Organic material adheres to mucus on the proboscis and is moved by cilia to the mouth. The mouth can be covered by the collar to avoid eating inorganic or other undesirable items (Ramel 2005).  
  
 
===Reproduction===
 
===Reproduction===
  
Acorn worms have separate genders that release eggs and sperm into the water for external fertilization. In some, eggs develop into free-swimming larvae that look very similar to echinoderm larvae (this suggests vertebrates and echinoderms are linked phylogenically). The larvae eventually settle down and change into tiny acorn worms on the surface and take on the burrowing lifestyle. Others don't have a larval stage, but develops directly into small juveniles.
+
Acorn worms have separate genders that release eggs and [[sperm]] into the water for external fertilization.  
 +
 
 +
In some, eggs develop into free-swimming larvae that look very similar to echinoderm larvae. After a number of weeks, the larvae change into tiny acorn worms and settle on the surface and take on the burrowing lifestyle. Others do not have a larval stage, but develop directly into small juveniles.
  
 
==Pterobranchia==
 
==Pterobranchia==
'''Pterobranchia''' is a [[clade]] of small, worm-shaped [[animal|animals]]. They belong to the [[hemichordata]], and live in [[secretion|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.
+
'''Pterobranchia''' is a class of Hemichordata that live in secreted tubes on the ocean floor, and feed by filtering [[plankton]]  out of the water with the help of cilia attached to tentacles. Unlike the enterpneusts, the pterobranchs possess only one, or even no, pharylgeal slits and each animal has only a single gonad, while enterpneuts have numerous gonads (Ramel 2005). The collar has between 1 and 9 pairs of tentacles, each of which has a double row of smaller ciliated tentacles (Ramel 2005).
 +
 
 +
There are about 30 known living species in the group. These are small, and range from 1 millimeter to 12 millimeters (Ramel 2005).  
  
Pterobranchia were established by [[Ray Lankester]] in [[1877]]. It contained, at that time, the single [[genus]] ''[[Rhabdopleura]]''. ''Rhabdopleura'' was at first regarded as an aberrant [[Polyzoon]], but with the publication of the Challenger Report (''[[Cephalodiscus]]'') in [[1887]], it became clear that ''Cephalodiscus'', the second genus now included in the [[Order (biology)|order]], had affinities in the direction of the [[Enteropneusta]].
+
Pterobranchia was established by Ray Lankester in 1877. It contained, at that time, the single genus ''Rhabdopleura''. ''Rhabdopleura'' was at first regarded as an aberrant Polyzoon, but with the publication of the Challenger Report (''Cephalodiscus'') in 1887, it became clear that ''Cephalodiscus'', the second genus now included in the order, had affinities in the direction of the Enteropneusta.
  
Recent advances in [[electron microscope|electron microscopy]] have suggested that pterobranchs belong to the same [[clade]] as the extinct [[graptolite]]s.
+
Recent advances in electron microscopy have suggested that pterobranchs belong to the same clade as the extinct graptolites.
  
 
==References==
 
==References==
 
* {{1911}}
 
* {{1911}}
* [http://faculty.washington.edu/bjswalla/Hemichordata/Pterobranchia/class%20pterobranchia.html At Billie Swalla's site]
+
* Ramel, G. J. L. 2005. [http://www.earthlife.net/inverts/hemichordate.html The Phylum Hemichordata]. Earthlife.net website. (accessed December 8, 2006).
 +
 
 +
 
 +
[http://faculty.washington.edu/bjswalla/Hemichordata/Pterobranchia/class%20pterobranchia.html At Billie Swalla's site]
 +
* [http://cluster3.biosci.utexas.edu/faculty/cameronc/Hemichordate.Species.html Cameron, C. B. ''Classification of the extant hemichordata'']
 +
 
 
* [http://cluster3.biosci.utexas.edu/faculty/cameronc/Hemichordate.Species.html Cameron, C. B. ''Classification of the extant hemichordata'']
 
* [http://cluster3.biosci.utexas.edu/faculty/cameronc/Hemichordate.Species.html Cameron, C. B. ''Classification of the extant hemichordata'']
 +
* [http://tolweb.org/tree?group=Hemichordata&contgroup=Deuterostomia At the Tree of Life]
  
  

Revision as of 23:57, 8 December 2006

Hemichordata
Acorn worm
Generalized hemichordate. Colors used here to emphasize the different body sections.
Scientific classification
Kingdom: Animalia
Subkingdom: Eumetazoa
Branch: Bilateria
Superphylum: Deuterostomia
Phylum: Hemichordata
Bateson, 1885
Classes

Hemichordata is a small phylum of worm-shaped, marine invertebrates. Hemichordates are bilaterially symmetrical and their bodies are divided into three sections (Protosome or proboscis, collar, and trunk). The musculature in their gut is very poorly developed, and food is mostly transported through it by using the cilia that covers the inside.

Hemichordates are considered a sister group of the chordates and echinoderms. They were once considered part of Chordata, but lack a true notochord. A hollow nerve cord, or notochord, is found in all chordates, even tunicates, and some hemichordates also seem to have a primitive form of notochord (at least in early life). In the embryonic stage, this tubular nerve cord looks like the hollow nerve cord of chordates. Both Hemichordata and Chordata have gill slits, and primitive fossil echinoderms also show signs of gill slits.

As with the Chordata and Echinodermata phyla, Hemichordata are deuterostomes. As deuterostomes, they have true coeloms (body cavities), with the coelom forming through enterocoely (the mesoderm forms as evaginations of the developed gut that pinch off, forming the coelom); the first opening becomes the anus rather than the mouth as in protostomes; and the early divisions of the zygote occurs parallel or perpendicular to the polar axis (radial cleavage).

Hemichordata includes about 100 living species. These are widely distributed, being found both in shallow costal waters and in the deeper sea, and are benthic (live on the sea floor), at least in their adult form (Ramel 2005). Hemichordates range from only a few millimeters long to the Balanoglossus gigas, which reaches 1.5 meters (4.7 feet) (Ramel 2005).

Hemichordata classes

Hemichordates date back to the Lower or Middle Cambrian and include an important class of fossils called graptolites, most of which became extinct in the Carboniferous.

Hemichordata is divided into two classes: the Enteropneusta, commonly called acorn worms, and the Pterobranchia, which may include the graptolites.

A third class, Planctosphaeroidea, is proposed based on a single species known only from larvae.

One of the suggestions are that the pterobranchs are more basal deuterostomes, while the enteropneusts are an early offshoot of the lineage that lead to Chordata.

Enteropneusta (Acorn worms)

The Acorn worms or Enteropneusta include about 70 known species. As with all hemichordates, acorn worms are benthic and may be deposit feeders (substrate feeders) or suspension feeders (filter feeders). Substrate feeders, like Balanoglossus clavigerus, are generally larger than filter feeders (Ramel 2005).

The genus Balanoglossus is also known as the tongue worm.

Anatomy

The Acorn worm's body is cylindrical and made up of three main parts: the acorn-shaped proboscis, a short fleshy collar that lies behind it, and the long trunk, which is the rest of the body. The creature's mouth is located at the collar behind the proboscis. One theory is that this 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.

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 by cilia movements and body contractions.

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.

Similarities to Chordates

Acorn worms are considered more highly specialized and advanced than other similarly shaped worm-like creatures. They have a circulatory system with a heart that also functions as a kidney. Acorn worms have gill-like structures that they use for breathing, similar to the gills of primitive fish. Hence, acorn worms are sometimes said to be a link between classical invertebrates and vertebrates.

Some also have a post-anal tail that sometimes show weak signs of segmentation. An interesting trait is that its three-section body plan is not present in the vertebrates, except from the anatomy of the frontal neural tube, later developed into a brain that is divided into three main parts. This means some of the original anatomy of the early chordate ancestors may still present even if it is not always visible.

Lifestyle

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). They rarely leave their burrows, which may have several openings (Ramel 2005).

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. Organic material adheres to mucus on the proboscis and is moved by cilia to the mouth. The mouth can be covered by the collar to avoid eating inorganic or other undesirable items (Ramel 2005).

Reproduction

Acorn worms have separate genders that release eggs and sperm into the water for external fertilization.

In some, eggs develop into free-swimming larvae that look very similar to echinoderm larvae. After a number of weeks, the larvae change into tiny acorn worms and settle on the surface and take on the burrowing lifestyle. Others do not have a larval stage, but develop directly into small juveniles.

Pterobranchia

Pterobranchia is a class of Hemichordata that live in secreted tubes on the ocean floor, and feed by filtering plankton out of the water with the help of cilia attached to tentacles. Unlike the enterpneusts, the pterobranchs possess only one, or even no, pharylgeal slits and each animal has only a single gonad, while enterpneuts have numerous gonads (Ramel 2005). The collar has between 1 and 9 pairs of tentacles, each of which has a double row of smaller ciliated tentacles (Ramel 2005).

There are about 30 known living species in the group. These are small, and range from 1 millimeter to 12 millimeters (Ramel 2005).

Pterobranchia was established by Ray Lankester in 1877. It contained, at that time, the single genus Rhabdopleura. Rhabdopleura was at first regarded as an aberrant Polyzoon, but with the publication of the Challenger Report (Cephalodiscus) in 1887, it became clear that Cephalodiscus, the second genus now included in the order, had affinities in the direction of the Enteropneusta.

Recent advances in electron microscopy have suggested that pterobranchs belong to the same clade as the extinct graptolites.

References
ISBN links support NWE through referral fees

  • This article incorporates text from the Encyclopædia Britannica Eleventh Edition, a publication now in the public domain.
  • Ramel, G. J. L. 2005. The Phylum Hemichordata. Earthlife.net website. (accessed December 8, 2006).


At Billie Swalla's site


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