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Labs
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Phylum Chordata
Chordata (L., chorde =
cord)
Important features of the group:
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All members of the phylum
Chordata exhibit the following four major morphological features at some
stage of their life cycle.
- Notochord
- Dorsal Hollow Nerve Cord (DHNC)
- Pharyngeal Slits
- Post-anal Tail
The notochord, the feature for which the subphylum is named, is an elastic,
rod-like structure that runs parallel to and between the digestive tract
and central
nervous
system.
It is
found
in
all chordate
embryos and serves as a support structure, resisting "telescoping" of
the body
when longitudinal muscles shorten. The notochord is composed of fluid-filled
cells wrapped in a fibrous sheath. Although functionally replaced by
the vertebral column in many chordate groups (principally the "higher"
vertebrates), the notochord is still an important embryonic structure
in all chordates. In adult humans, and other mammals, the notochord
is preserved as the nucleus pulposus, a small gel-like region at the
core
of each
of
the
intervertebral discs between adjacent vertebrae.
The dorsal, hollow nerve cord forms the central nervous
system (brain and spinal cord) of all chordates. It is distinct from
the central nervous systems of most non-chordate invertebrates in its
position,
form,
and
development.
The chordate nerve cord is positioned dorsal to the gut tube, while the
nerve cord of most non-chordates is found ventral to the digestive tract.
The chordate nerve cord is also hollow and fluid-filled, while
that of non-chordates is typically solid. Lastly, the chordate nerve
cord forms by a invagination, an embryonic process in which neural tissue
gathers dorsally on the outer surface of the embryo, folds into a tube,
and then sinks inward (invaginates) to take up its internal position.
The nerve cords of non-chordates do not form by invagination, instead
cells typically move inward to their internal positions individually.
The pharyngeal slits are paired longitudinal series
of perforations through the wall of the pharynx (the region of the digestive
tract between
the
mouth
and
esophagus). These slits appear in in the embryo stage of all chordates
and may persist into adulthood or may close and disappear before birth
or hatching (as is the case in mammals, for example). Pharyngeal slits
allow water that is brought in via the mouth to exit without passing
through the entire digestive tract. In ancestral chordates and modern
urochordates and cephalochordates (as well as hemichordates) pharyngeal
slits function as suspensio-feeding structures. Vertebrates have modified
the slits and their support structures for other functions including
gas exchange, jaw support, and hearing.
The post-anal tail is a muscular region of the body
that extends beyond the anus. In non-chordates the digestive tract usually
extends the entire length of the body and the anus is typically positioned
at the posterior end of the body (an earthworm is a good example of this
type of body plan). The chordate post-anal tail includes
skeletal support and musculature that improves the locomotion of many
aquatic
chordate
species.
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Chordate characteristics All
chordates possess the four trademarks of the phylum: a notochord;
a dorsal, hollow nerve cord; pharyngeal slits; and a muscular,
postanal tail. (From Campbell
6th edition's Fig
34-2.)

Notochord function The
notochord resists shortening in length, but is flexible laterally.
Without a notochord, lateral muscle
contraction collapses the body along its length. With a notochord
muscle contractions on alternating sides efficiently flex the
body
in simple
swimming strokes. (From Kardong
3rd edition's Fig
2.5.)

Neurulation - development of the
dorsal hollow nerve cord. The neural tube originates as
a plate of dorsal ectoderm just above the developing notochord.
This "neural plate" soon folds inward, rolling itself
into the neural tube, which will become the central nervous system
- the brain and spinal cord. These organs are hollow in most chordates
because of this mechanism of development. (From Campbell
6th edition's Fig
47-11.)

Post-anal tail. Recall
that the digestive tract of the shark terminates at the
cloaca, between the pelvic fins. The remainder of the body
then represents a post-anal tail and plays a huge role in
propulsion during aquatic locomotion. The paired fins play
little or no propulsive role in most fish. Seen in this
photo is a Blue Shark (Prionace glauca) swimming
underwater in the Pacific Ocean, thirty miles off of San
Diego, California. Blue sharks have been recorded swimming
at speeds in excess of 35 mph!
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Examples:
The phylum Chordata includes three subphyla:
Examples of each subphylum are shown at right. For more detailed information
on each subphylum follow the links above.
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A larval tunicate - note the chordate
characteristics in the diagrammatic view.

Metamorphosis of a larval urochordate. Note
the loss of all chordate characteristics except the pharyngeal slits.
Click image to enlarge.

Adult urochordates. Note the
inhalant (larger) and exhalant siphons and the striking convergence with
sea sponges -click here for an example.
Subphylum Urochordata: a tunicate. (a)
An adult tunicate, or sea squirt, is a sessile animal commonly arranged
in a U shape (photo
is approximately life-sized). (b) In the adult, prominent pharyngeal
slits function in suspension feeding, but the other chordate characteristics
are not obvious. (c) In the tunicate larva, which is a free-swimming
but nonfeeding "tadpole," the chordate characteristics
are evident: It has a notochord, dorsal nerve tube, and a tail with
muscle
segments. (From Campbell 6th edition's Fig
34-3.)

Subphylum
Cephalochordata: the lancelet Branchiostoma
(also called amphioxus). This
small invertebrate displays all four chordate characteristics. The
pharyngeal
slits function
in suspension feeding. Water passes into the pharynx and through
slits into the atrium, a chamber that vents to the outside via the
atriopore.
Food particles trapped by mucus are swept by cilia into the digestive
tract. The muscle segments you can see in this photo of a lancelet
produce the sinusoidal swimming of these animals. (From Campbell
6th edition's Fig
34-4a.)

Subphylum Cephalochordata: the
lancelet Branchiostoma. The
muscle segments you can see in this photo of a lancelet produce the
sinusoidal swimming of these animals. (From Campbell
6th edition's Fig
34-4b.)

Subphylum Vertebrata (Class Mammalia): The Vertebrates
are characterized by a well-developed brain, cranium and sense
organs. As their name implies, they are also characterized
by vertebrae which protect their dorsal hollow nerve cord. Bone
and a muscular phayngeal pump are also characteristics of vertebrates. The
mammal seen here (Canis familiaris) is representative
of the class Mammalia, displaying both of the class' defining characteristics
- mammary glands and hair.
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ECHINODERMS, HEMICHORDATES, AND CHORDATES SHARE CERTAIN CHARACTERISTICS
The
hemichordates have apparent affinities to both the echinoderms and the chordates,
showing an evolutionary relationship between those two large and important
phyla. The ciliated larvae of hemichordates are so similar to those of some
echinoderms that they were mistaken for echinoderm larvae when first discovered.
The dipleurula type of larva is found only in the echinoderms and hemichordates.
It has a band of cilia encircling the mouth, whereas the trochophore type
of larva found in many protostomes (including some flatworms, molluscs, and
annelids)
has a band of cilia encircling the body anterior to the mouth (see Figure
below). The similarity of the larvae of hemichordates and echinoderms, as well
as the
similarities in their early embryology, indicate that these two groups probably
stem from a common ancestor.
The most obvious resemblance of hemichordates to chordates is
their possession of pharyngeal slits (and perhaps a short dorsal nerve cord)
which are found
in all chordates but nowhere else in the animal kingdom. While the hemichordates
are closer to the echinoderms than to the chordates, recognition of their
ties with both Chordata and Echinodermata has helped clarify the evolutionary
relationship
between these two major groups. Note that there is no suggestion here
that chordates evolved from echinoderms, but simply that the two groups diverged
from a common ancestor at some remote time.

Chordate Origins via Paedomorphosis

Summary of Garstang's hypothesis
regarding the origin of chordates and vertebrates. Beginning
with an echinoderm larva, Garstang proposed a series of literal
evolutionary steps through the larval stages that involved paedomorphosis
and eventually produced chordates. (From Kardong
3rd edition's Fig
2.30.)
The hypothesis described in the figure
above is one possible explanation of how chordates originated from an echinoderm
ancestor. There is strong evidence for some version of Garstang's hypothesis
though. Echindoerms and chordates are clearly both deuterostomes, sharing
similar embryology. Bilateral symmetry is also evident in both groups, especially,
and notably, in the larvae of echinoderms. These facts solidify the union
of echinoderms and chordates as a monophyletic group (Deuterostomes) and
are suggestive of an evolutionary origin for both groups from something resembling
an larval echinoderm.
Add to this the fact that the larvae of
echinoderms and hemichordates (the group in which some chordate characteristics
are first evident) and the next step is explained. Shared characteristics,
in particular those seen in the larvae, link echinoderms to hemichordates.
Shared characteristics also link adult hemichordates to chordates - the pharyngeal
slits, and dorsal nerve cord for example.
This pattern of paedomorphosis may have
been repeated again in the transition from urochordates to cephalochordates.
See the Cephalochordate page in this demo
for more information on this.
It is important to emphasize that scientists
do not agree on a single explanation for chordate origins. The hypothesis presented
above is one somewhat popular and well-supported theory.
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