PROTOCHORDATES

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PROTOCHORDATES

A. Phylum: Hemichordata

Characteristics:
1. Pharyngeal clefts usually present
2. Dorsal nerve cord

Similarities with invertebrates exclusive of chordates include:


1. Ventral nerve cord, solid.
2. No notochord; stomochord present.
3. Free-swimming larva--tornaria.

Major groups of hemichordates:

1. Class: Pterobranchia.
• Mostly sessile, stalked, colonial
• Gather food my muco-ciliary arms (lophophores).
• Few or no pharyngeal slits.

2. Class: Enteropneusta (Acorn Worms).

• Mostly free-swimming, sedentary, burrowing.


• Proboscis, mouth at anterior end of collar leads into pharynx, pharyngeal slits present.
• Gather food by muco-ciliary proboscis.
• Dorsal hollow nerve cord only in collar.

Figure from Kent & Miller, courtesy of WCB/McGraw-Hill

B. Urochordata (Tunicata)

• About 2000 species.


• Usually have notochord during early development (confined to tail--strengthens tail for locomotion); not retained in ascidians.
• Heart pumps blood in one direction for a time and then reverses the pump in the other direction.
• No head, tail, or metamerism in adults that metamorphose.
• Cellulose-like tunic.

Major groups (3):

1. Ascidiacea (sea squirts).


a. Larvae.
• Have notochord and dorsally located nerve cord.
• Food and respiratory water enters through incurrent siphon and then passes on to pharynx.
• Pharynx and atrium begin to develop in the larva.
b. Adults.
• At metamorphosis the larva attaches to substrate by adhesive suckers located at its anterior end and tail is resorbed.
• Notochord dissappears.
• Rearrangement of internal organs.
• No special sense organs.

Figure from Kent & Miller, courtesy of WCB/McGraw-Hill

2. Thaliacea.

• Pelagic tunicates; barrel-shaped.


• Move by expelling water from their excurrent siphon.

3.Larvacea.
• Also pelagic and free-swimming throughout life.

• Resemble larval ascidians (example of paedogenesis).

• Retain notochord throughout life--aids in stiffening tail for locomotio

igure from Kent & Miller, courtesy of WCB/McGraw-Hill

C. Cephalochordata (lancelets).
• About 45 species.
• Branchiostoma
• Eel-like, free-swimming, sedentary, burrowing.
• Persistent notochord.
• Epidermis single cell layer.
• Caudal, dorsal, and ventral fins.
• Musculature--myomeres and myosepta (metamerism).
•No highly organized sense organs.

Figure from Kent & Miller, courtesy of WCB/McGraw-Hill

D. Differences between cephalochordates and vertebrates.

• Almost no cephalization.
• No paired sense organs.
• No vertebral column.
• High number of gill slits.
• Segmented musculature extends to anterior tip of head.
• No paired appendages.
• Outer layer of skin (epidermis) one-cell thick.
• No muscular heart.
• Excretory protonephridia resemble those on non-chordates.

E. Protochordate and vertebrate relationships.

1. Evidence for relationship between chordates and certain other invertebrates.

• Protochordates and vertebrates resemble echinoderms and hemichordates because they are deuterostomes. Characteristics of this group
include:

Relationship of the Vertebrata

with other dueterostomes.


2. Other evidence for relationships between protochordates and vertebrates

• The most primitive living vertebrates are jawless fishes (cyclostomes).


• The free-swimming, filter-feeding larval stage of lampreys closely resembles protochordate larvae--this provides us with further evidence for a
close relationship between vertebrates and protochordates.
• Ammocoete larvae (filter feeders) have:
a. 7 gill slits (respiratory in function).
b. Notochord extending from head region into tail (no vertebral column).
c. Dorsal, hollow nerve tube; typical embryonic vertebrate brain with three primary vesicles.
d. A stratified epidermis.
e. Typical embryonic vertebrate kidney.
f. Typical closed, embryonic vertebrate circulatory system with two-chambered heart and red blood cells.
g. Tadpole-like morphology typical of vertebrate larvae (such as frogs).
h. Sense organs more numerous than lancelets--median naris that leads to an olfactory sac, two middorsal eyes on head, otic vesicles that develop
into inner ears.

Figure from Kent & Miller, courtesy of WCB/McGraw-Hill

F. Garstang theory of vertebrate evolution; possible steps in the evolution of vertebrates.

1. Pterobranchs and primitive echinoderms--derived from a hypothesized common ancestor--a sessile arm-feeder.
2. Shift from lophophores to gill-filtering to obtain food.
3. Development of a tadpole-like larva.
4. Neoteny--larval locomotor structures retained throughout life.

Protochordates and Other Interesting Critters

1. The exact origin of vertebrates remains unresolved. Members of the group known as protochordates appear to
be close relatives and so are included in the phylum Chordata

2. Distinguishing characteristics of the phylum Chordata:

Gill Slits: in wall of pharynx (at some point during the life of the animal).
Notochord: also present sometime during life.
Dorsal Hollow Nerve Cord
Postanal Tail

3. Protochordates make up 2 of the 3 chordate subphyla: the Urochordata and Cephalochordata (the remaining one is,
as you might guess, the Craniata).

These animals all share the distinguishing characteristics listed above but lack vertebrae.

4. Urochordata: (meaning the notochord is in the tail) includes the ascidians (sea squirts,tunicates), larvaceans,
and thaliaceans. Larvae are free swimming. Larval ascidians do not feed and swim for only a few days (or less).
Water entering the mouth passes over the gills and out through the atrium. Adhesive papillae attach larvae to
substrate (pilings, rocks, etc.). Following metamorphosis, adults lose notochord, mouth becomes the incurrent
siphon, atriopore becomes the excurrent siphon. Rearrangement of viscera. Food is trapped by mucus in the
endostyle and directed to esophagus. Respiratory water passes over gills and into atrium.

5. Cephalochordata: (meaning the notochord is in the head) includes Amphioxus, a very common marine organism.
Adults are nearly all trunk and tail. Muscles metameric and separated by myosepta. Pharyngeal gill slits open into
the atrium. Filter feeders, but have the ability to swim. Notochord persists throughout life as the primary
supporting structure.

6. Craniata: animals with a vertebral column of bone or cartilage. Nine major extinct or extant classes:
Agnatha - Cyclostomes, 45 species, jawless, poorly developed or absent fins.
Acanthodii - extinct jawed fish
Placodermi - also extinct; bony armor
Chondrichthyes - cartilaginous fish. 575 species. Sharks, rays, and chimeras
Osteichthyes - over 17,000 species. Bony fish
Tetrapods
Amphibia - amphibians, 2,400 species
Reptilia - reptiles. 6,000 living species
Aves - birds, 8,700
Mammalia - mammals. 4,500 species

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