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Phylum Chordata
The chordates are a group of particularinterest to us as we belong to it, beingmembers of the subphylum Vertebrata.
The chordates include all of thevertebrates (fish, amphibians, reptiles,
mammals and birds), but also two non-vertebrate subphyla: the Urochordata andthe Cephalochordata.
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Phylum Chordata
The chordates were in the 19thcenturyconsidered to have been derived fromprotostome ancestors (the annelid, mollusc,arthropod group).
However, a better understanding of embryologyshows that chordates are deuterostomes and
the invertebrates they are most closely related toare the Echinodermata (sea stars, sand dollars,sea urchins) and the Hemichordata (acornworms).
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Important differences in development
between protostomes and deuterostomes
The differences in development that distinguish
the protostomes and deuterostomes include:
Whether cleavage of cells in the early zygote is spiral
or radial. Whether or not, if the early blastomere is separated,
each cell can develop into a normal larva or not.
Whether the blastopore ultimately forms the mouth or
anus of the organism. Whether or not the organism possesses a coelom
and how that coelom is formed.
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Protostomes and Deuterostomes
Protostomes include the annelids,
mollusks, and arthropods.
Deuterostomes include the echinoderms
and Chordates, which includes the
vertebrates.
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Figure 23.02
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Characteristics of the Chordata
Chordates are:
bilaterally symmetrical
triploblastic
have a well developed coelom
have a complete digestive system
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Germ layers of a triploblastic
organism Endoderm: innermost germ layer of an embryo. Forms
the gut, liver, pancreas.
Ectoderm: Outer layer of cells in early embryo.
Surrounds the blastocoel. Forms outer epithelium ofbody and nervous system.
Mesoderm: Third germ layer formed in gastrula between
ectoderm and endoderm. The coelomforms in themesoderm. Mesoderm gives rise to connective tissue,muscle, urogenital and vascular systems andperitoneum.
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Coeloms
The coelom is a cavity entirely surroundedby mesoderm.
A coelom provides a tube-within-a-tube
arrangement which has many advantages:
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Advantages of a coelom
In mammals the pericardial, peritoneal, andpleural cavities are formed from the coelom.
Allows flexibility in arranging visceral organs
Permits greater size and complexity ofinternal organs as there is space for them togrow
Fluid-filled coelom can act as a hydrostatic
skeleton Cushions organs against impact
Allows organs to grow and moveindependently of your outer body wall.
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Five distinctive characteristics of
the chordates Five distinctive characteristics separate the
chordates from all other phyla: Notochord
Single, dorsal, tubular nerve cord
Pharyngeal pouches or slits
Endostyle
Postanal tail
Not all of these characteristics are apparent inadult organisms and may appear only in theembryonic or larval stages.
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Notochord
Notochord: the notochord is a flexible,
rodlike structure. It extends the length of
the body and is an anchor point for
muscles.
The notochord bends without shortening
so it permits the animal to undulate.
Figure 23 01
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Figure 23.01
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Notochord
In nonvertebrates and the jawless
vertebrates the notochord is present
throughout life.
However, in the jawed vertebrates it is
replaced by the vertebral column; the
remnants of the notochord being found in
the intervertebral disks.
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Single, dorsal, tubular nerve
cord In most invertebrates the nerve cord, if present,
is ventral to the gut.
In chordates, in contrast, the nerve cord is dorsalto the gut and notochord. The nerve cordpasses through the neural arches of thevertebrae, which protect it.
The nerve cord is enlarged in vertebrates into abrain, which is surrounded by a bony orcartilaginous cranium.
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Pharyngeal pouches and slits
Pharyngeal slits occur in aquatic chordates andlead from the pharyngeal cavity to the outside.
The pharyngeal slits are used as a filter feedingdevice in protochordates (i.e., Urochordata(Tunicates)) and Cephalochordata (lanceletse.g. Amphioxus).
Water containing food is drawn in through themouth by cilia and exits via the pharyngeal slitswhere the particles are trapped in mucus.
Figure 23 09b
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Figure 23.09b
Amphioxus
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Pharyngeal pouches and slits
In vertebrates the pharyngeal arches have
been modified into gills by the addition of a
rich blood supply and thin gas permeable
walls.
The contraction of muscles in the pharynx
drive water through the gills.
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Pharyngeal pouches and slits
In amniotes an opening may not form and
rather than slits only grooves called
pharyngeal pouches develop.
In tetrapods these pouches give rise
during development to a variety of
structures including the middle ear cavity,
eustachian tube, and tonsils.
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Endostyle or thyroid gland
The endostyle is found in protochordates
and in lamprey larvae. It is located on the
floor of the pharynx and secretes mucus,
which is used to trap particles.
The endostyle works with the pharyngeal
slits in filter feeding.
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Endostyle or thyroid gland
Some cells in the endostyle secrete
iodinated proteins and are homologous
with the iodinated-hormone secreting
thyroid gland, which is found in adultlampreys and vertebrates.
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Postanal tail
The postanal tail, some musculataure and thenotochord enable larval tunicates andamphioxus to swim.
The postanal tail evolved to allow organisms toswim and its efficiency has been enhanced bythe addition of fins. The postanal tail is present
only in vestigial form in humans (the coccyx)although tails as a whole are widespread amongvertebrates.
Figure 23.09b
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Amphioxus
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Classification of the Chordata
There are three subphyla in the Chordata:
Subphylum Urochordata: tunicates
Subphylum Cephalochordata: lancelets
Subphylum Vertebrata: fish, amphibians,
reptiles, birds, mammals, etc.
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Subphylum Urochordata
The Urochordata (tunicates named for the
tough tunic that surrounds the adult) look like
most unpromising candidates to be chordates
and relatives of the vertebrates.
The largest group, the ascidians or sea squirts
(Class Ascidiacea) as adults are marine, sessile,
filter feeding organisms that live either solitarily
or in colonies.
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Ciona intestinalis
(a solitary sea squirt)
0147.jpg
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Synoicum pulmonaria a colonial sea squirt
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Ascidians
Adult ascidians lack a notochord and there
is only a single ganglion in place of the
dorsal nerve cord.
Of the five characteristics of chordates
adults possess only two: pharyngeal gill
slits and an endostyle, both of which theyuse in filter feeding.
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Ascidians
The adult sea squirt draws water in
through an incurrent siphon and pushes it
back out an excurrent one.
Food particles are filtered out in the
pharyngeal slits with mucus from the
endostyle used to trap particles.
Figure 23.04
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15.4
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http://seapeachbio.blogspot.com/2012/06/ascidian-filter-feeding-mechanism.html
Ascidian filter feeding process. Water is drawn
through the animal by the beating of cilia.
Cilia lining the slits in the branchial sac or pharynx
Branchial sac orpharynx
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Larval Ascidian
Even though the adult ascidian hardly
resembles a chordate its larva does.
Larval ascidians are very small and
tadpole-like and possess all five chordate
characteristics previously outlined.
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Young larval ascidian
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Larval Ascidian
The larval ascidians role is to disperse andto achieve this it is free swimming.However, it has only a short larval life
(minutes to a couple of days) and does notfeed during this time.
Instead it searches for a place to settleand then attaches and metamorphosesinto an adult.
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Ascidian metamorphosis
During metamorphosis the notochord
disappears, the nerve cord is reduced to a
single nerve ganglion and a couple of
nerves.
Figure 23.06
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15.5
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Other Urochordate classes
Besides the ascidians there are two otherclasses of the Urochordata: the Larvaceaand Thaliacea.
Both are small, transparent planktonicforms. Thaliaceans are cylindrical or
spindle shaped whereas larvaceans aretadpolelike and resemble an ascidianlarva.
G f
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Garstangs hypothesis of chordate
larval evolution
In the 1920s it was proposed that the
vertebrates were derived from an
ancestral ascidian that retained its
characteristics into adulthood (the processby which juvenile characteristics are
retained into adulthood is referred to as
paedomorphosis).
Figure 23.12
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G t h th i f h d t
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Garstangs hypothesis of chordate
larval evolution
Garstangs hypothesis is supported by
embryological evidence, but more recently
molecular analyses have suggested that
sessile ascidians are a derived form andthat the free-living larvaceans are more
likely to be the closest relatives of the
chordates.
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Subphylum Cephalochordata
The cephalochordates are the lancelets, whichare small (3-7 cm long) laterally compressedfishlike animals that inhabit sandy sediments ofcoastal waters. They lack a distinct head and
have no cranium.
They are commonly referred to as Amphioxus asthis was the original genus name. There are 29species, five of which occur in North Americancoastal waters.
Figure 23.09b
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Amphioxus
Amphioxus is a filter feeder.
Water enters the mouth and then passes
through the pharyngeal slits, where food is
trapped in mucus. Cilia then move the
food to the gut.
Figure 23.09a
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Amphioxus
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Amphioxus
Amphioxus is interesting because it displays the
basic chordate characteristics in a simple and
obvious form because of its transparency.
Amphioxus is considered to be the closest living
relative of the vertebrates because it shares
several characteristics with vertebrates that
Urochordates do not possess.
Amphioxus characteristics
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Amphioxus characteristics
shared with vertebratesAs well as possessing the chordate
characteristics, amphioxus shares a
number of traits with vertebrates including:
Segmented myomeres (muscle blocks)
Dorsal and ventral aortas
Branchial (gill) arches (blood vessels running
over the gills).
Characteristics of vertebrates
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Characteristics of vertebrates
notpossessed by AmphioxusAmphioxus however lacks several
characteristics that biologists think the
ancestor of vertebrates possessed. These
include: Tripartite brain(with forebrain, midbrain and
hindbrain) protected by a cranium
Chambered heart Muscular gut and pharynx
List continues on next slide
Characteristics of vertebrates
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Characteristics of vertebrates
notpossessed by Amphioxus Various special sensory organs(eyes,
chemical and pressure receptors)
Neural crest(ectodermal cells that are found
on the embryonic neural tube and areengaged in the formation of the cranium, tooth
dentine, some endocrine glands and
Schwann cells, which provide myelin
insulation to nerve cells).