Cordate info: last page

"Protochordates" Hemichordates: Some biologists believe they're classes within a single phylum;
a second interpretation places them as phyla on either side of the echinoderms

living hemichordates may provide some clues to how chordates evolved.

Whether this is true is being challenged by molecular data suggesting that these
animals may be more closely related to the echinoderms than to the chordates.
(Regardless of which taxonomic position is correct for the hemichordates,
it doesn't change that these animals have two distinct body plans with some
shared characteristics and others unique to each).

Hemichordates and
Protochordates
In chordates: notochord forms along dorsal surface of entire gut, seperates into unique
solid skeletal structure

In hemichordates: have a STOMOCHORD, dervived from primitive gut but DIFFERENT

analogous to notochord

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tripartate coelom: protocoel, mesocoel, metacoel

Porifera
Placozoa Hemichordates are:
Cnidaria
Ctenophora - bilateral, unsegmented with 2 body plans/"homes";
Platyhelminthes
Gastrotricha
Gnathostomulida solitary ENTEROPNEUSTA: U-shaped burrows
Cycliophora colonial PTEROBRANCHIA: Tubes they secrete
Rotifera
Annelida
Mollusca they share these traits:
Sipuncula
Nemertea -Tripartate (eucoelomate) deuterostomes
Bryozoa -ciliated epidermis with mucus producing gland cells (trap food, line burrows, build
Brachiopoda
Phoronida tubes)
autapomorphies Arthropoda
Onychophora
-sessile
!! Hemichordata Tardigrada -dioecious, gametes released in water
• Stomochord form a complex
Nematomorpha -hydrostatic skeleton (coelomic fluid + muscles) which
Nematoda
• Proboscis Priapulida in some parts of body includes circular & longitudinal muscles, probiscus*, collar of
• Glomerulus Kinorhyncha the enteropneusts or only longitudinal muscles in the enteropneust trunk, and all
Loricifera
Echinodermata parts of the pterobranchs.
!! Hemichordata
Chordata

NOT "probiscis"
proboscis: Tubular feeding structure that extends from an animal. In many invertebrates, it is eversible and extends from the body to permit feeding

Enteropneusta ("acorn worm")

Sessile, 3 distinct parts derived from tripartate regions

Hemichordate
ProbOscis = protocoel, the first compartment
Collar Collar = mesocoel, the second
(mesocoel) Trunk = metacoel (includes branchiogenital trunk, hepatic, intestinal
Proboscis trunk)
(protocoel) Branchiogenital
trunk (metacoel) Gill slits: exit for water that's filtered from the pharynx, gas exchange
across walls.
Gill slits
Anus Probiscus collects food, allows limited locomotion

Intestinal trunk (metacoel) Hepatic (metacoel)

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 Nervous system: a net connected to a dorsal nerve cord visible in the enteropneust
as a dorsal neurochord and in pterobranchs as only a dorsal thickening in the nerve net that covers the body

"Protochordates" Neurocord: Dorsal nerve cord or concentration of nerves in hemichordates.

It is not clear if the neurochord and notochord are homologous or dissimilar structures

Substrate feeding strategy in burrowers, suspension feeding in tube dwellers

Acorn worm burrow

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analogous: do not have the same evolutionary origin but have the same functions. Eg insect and bat wings, notocord and stomocord

food trapped in mucus covering probiscus, cilia pass to mouth on collar edge
Feeding
water carrying mucus passes through pharynx wall openings (EXITS VIA GILL PORES)
Rejected particles
NOTE:
Gill pores
(Gill pores are important in water movement, not in food selection,
Proboscis which is not the case in the chordates).
Gut
CILIA, not muscles, propel along digestive tract
Collar

Preoral ciliary organ Esophagus pharyngeal gill slits shared with chordates
Food entering mouth

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Open circulatory system with Heart vesicle located in proboscis. ->Posterior to anterior = DORSAL vessel
-Blood from dorsal vessel combines with blood from proboscis
-> flows over heart vesicle, GLOMERULUS
Proboscis complex -> then VENTRAL vessel -> posterior
supported by stomochord
formed from protocoel GLOMERULUS: removal of metabolic wastes
Mesocoel
Heart vesicle STOMOCHORD: forms from mesocoel, extends as hallow
collar, mesocoel
pocket into proboscis
Glomerulus with an opening INTO gut and ACTS AS A HYDROSTATIC
SKELETON
Protocoel
supporting the proboscis

It is ANALOGOUS to the chordate notocord (NOT homologous)

Mouth Stomocord

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GLOMERULUS IN:
VERTEBRATES: Part of kidney consisting of capillaries surrounded by the renal capsule;

HEMICHORDATES: network of blood sinuses forming a spongy tissue, assumed to have an excretory function
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 MORE on GLOMERULUS: when blood is forced into the central sinus by the heart vesicle,
there is sufficient pressure to force fluid from the blood across the glomerulus and into the protocoel.
Essential nutrients and ions are recovered from the coelomic fluid of the proboscis by cells lining the cavity,
"Protochordates" and urine is released through the proboscis pore.

Locomotion limited using proboscis to drag them along

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not all are vertebrates 4 characteristics: a notochord (gives the phylum its name);
a dorsal hollow nerve cord above the notochord;
pharyngeal gill slits;
and post-anal segmentation--a tail.
Chordates?

Segmentation via myomere muscle block arrangement (myotomes (Embryological

block of mesoderm in vertebrates that forms a myomere. In some cases
myomere and myotome are used interchangeably)) down body length.

2 major cranium based groups: acraniate and craniate

The notochord, which extends the length of the body, is a cartilaginous, flexible rod
covered with a fibrous sheath. Its original function was skeletal, and it maintained
the shape of the body when myomeres contracted.
In higher chordates it is replaced with vertebrae formed from cartilage and bone.
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8 gills slits important ancestral characteristic, replaced by gills and lungs in advanced
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cranium: cartilage or bone casing that contains

the enlarged anterior portion of nerve cord; the brain chordates: serious adaptative radiation (Evolution of a variety of different species from a single common ancestor.
Each is adapted for a particular niche, and the appearance of the descendants may vary considerably from
each other and the ancestor)
Porifera
Placozoa
Cnidaria
Ctenophora
Platyhelminthes
Gastrotricha
Gnathostomulida
Cycliophora
Rotifera
Annelida
Mollusca
Sipuncula
Nemertea
Bryozoa
Brachiopoda
Phoronida
Arthropoda
autapomorphies? Onychophora
!! Chordata Tardigrada
Nematomorpha
• Notochord
Nematoda
• Post anal segmentation Priapulida
• Pharynx with endostyle Kinorhyncha
• Ventral heart Loricifera
Echinodermata
Hemichordata
!! Chordata

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 "Protochordates"

there are 4 of them. There's a paraphyletic group.

Major chordate subphyla

O Urochordata
these 2 are lower
O Cephalochordata
O Vertebrata

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common characters of Chordata: notochord, pharyngeal gill slits,

post-anal segmentation (a tail), and a dorsal nerve cord; all found in larval urochordates
What’s
What’s new in Chordata
What’

O Notochord rod, new way of swimming

O Dorsal tubular nerve cord hallow
linked
O Pharyngeal gill slits respiration byproduct of role in getting water away from food
O Post anal segmentation
O Endostyle consolidating food particles. Ancestor filter feeder.
O (Ventral pumping heart)

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Originally used in filter feeding.

Pharyngeal gill slits unique filter feeding
tactic Pharynx perforated with gill slits, allow water to pass from inside to outside.

ENDOSTYLE produces mucus net lining inside of pharynx, traps food.

Food passed back to digestive system when net wound up by the cilia
lining the epipharyngeal groove along the dorsal surface of the pharynx.

GILL BARS (support for slits, formed from tissue btwn slits) vascularized and
gas exchange occurs as water flows over them.

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12 Side effect of food filtering is respiration.
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pharyngeal gill slit: Lateral opening in the wall of the pharynx that allows water
to enter into the mouth and exit through the pharynx.
This is one of the ancestral characteristics of the phylum Chordata and are
also found in Hemichordata and Enteropneusta. It allowed water to be removed
from ingested food before it was passed back into the digestive system
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 "Protochordates"
ENDOSTYLE: ciliated groove lying along the midventral line (and dorsal?) of the pharynx of chordates. Secretes a mucus net that rises from the groove
across the inner surface of the pharynx to the hyperpharyngeal groove on the opposite side of the pharynx. The net is porous and allows water through
while trapping particulate food on its inner surface (passed down dig syst to be wound into a cord by the hyperpharyngeal groove). Allows this to occur inside, not outside
like most filter feeders. The endostyle also incorporates iodine into the net, suggesting that the cells of the endostyle are the evolutionary precursor of the
thyroid gland found in other chordates.
Dorsal nerve chord
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Notochord
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fluid-filled rod of tightly packed cells surrounded by a sheath of elastic connective tissue.
uncompressible from anterior to posterior. Fixed length! Can bend
Notochord + myomeres = unique locomotion
Elastic sheath
Fibrous sheath
Notochord
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During embryological development, the presence of the notochord
signals the ectoderm above it to form a neural plate. As neurulation
proceeds, the two sides of the neural plate fold up and meet along the
midline, creating the hollow neural tube with a central neurocoel.
While this is happening, the whole structure sinks below the
ectoderm. The result is the hollow nerve chord that is characteristic of
the hemichordates and the chordates.
eventually dissapears, replaced by bony skeletal structure.
Cartilage between vertebrae remnants.
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nerve chord sits on top of notochord
ancestral characteristic of the Chordata, consists of a stiff
cartilaginous rod near the dorsal surface of the animal.
It is skeletal and helps support to the body
in ancestor, no protective casing around nervechord. notochord
strictly structural
remains in primitive/lower chordates
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defines chordata, BUT:
remains only in primitive subphyla: urochordates and cephalochordates.
In vertebrates, the notochord's role as a skeletal structure has been
replaced by the vertebral column.
Myomeres and movement:
Muscles are arranged segmentally in blocks (MYOMERES) along length of noto,
(myomeres connected to neighbour with mysosepta).
Fibres are aligned anterior->posterior with no fluid filled coelomic space as hydrostatic.
Result: NO notochord: myomere contracts, length shrinks and adjacent myo stretched
-> accordian like ripple
WITH notochord: contraction bends notochord.
Relax & elastic sheath returns to straight position --> restretches myomere to original.
Rinse and repeat on alternating sides (coordinated contractions down length)
->typical aquatic swimming motion, trunk & tail undulating side to side propelling
foreward
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 "Protochordates"

See above for myomeres+notochord = undulating swimming motion

Segmentation
muscles segmented along notocord as myomeres

extended notochord and muscle block segmentation

beyond the anus: post anal segments -> TAIL

creates more undulation, more power, more speed

Tail is a modification for a new form of locomotion unique to chordates.

Post anal segments (Tail)

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pharyngeal basket: In ascidian adults the pharynx is enlarged to form the pharyngeal basket.
Pharyngeal openings in the wall of the basket allow water to pass through with food being
trapped on the inner surface before being passed into the gut

ciliated Pharyngeal basket = main body;

Sessile ancestor surrounded by atrium which opens to outside via ATRIAL SIPHON
Urochordates
solitary and colonial food trapped by mucus sheet produced by endostyle which is gathered into
a cord by cilia and passed by the ciliated endostyle to the U shaped digestive
system.
Mouth SIMPLE DIFFUSION accounts for lack of excretory/respiratory systems.
Atrium
Endostyle open circulatory w/ tubular heart, small ganglion anterior to pahrynx
Pharyngeal gill slits
Gonad Missing most chordate traits!
Stomach

ADULT, specialized suspension METAMORPHOSIS TO LARVA

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common name: tunicate, from the living TUNIC covering adult body.
Tunic: composed of cellulose like protein called TUNICIN (gelatenous to leathery)
it is LIVING (unlike cuticle) with wandering AMOEBOCYTES

ALL 4 CHORDATE TRAITS main organs in PRECAUDAL (before the tail) region
Urochordate larva notocord, myomeres and dorsal tubular nerve cord extend length of body

notocord sole skeletal structure

Incurrent Excurrent siphon
siphon and anus Dorsal tubular branchial basket (pharygeal?) is present and still surrounded by atrium, same feeding
nerve cord

Endostyle
Notochord
Pharyngeal gill slits Post anal segments (Tail)

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18 take adult basket, but one end add a motile tail.
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tail has segmentation, post anal, nerve cord, notochord.

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 "Protochordates"

anchors with sucker/papillae.

Urochordate metamorphosis
Loses tail, pharyngeal basket enlarges and rotates 180 degrees so opening
faces up.

see all chordate traits in larvae, not adult. Very distinct!

monoecious

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all 4 characteristics: notocord, pharygeal gill slit, dorsal hallow nerve cord
above it, tail(segmentation/myomeres)
Cephalochordates
lancelets

notocord only skeletal element, myomeres chevron shaped

filter feeders, burrow into substrate

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ATRIOPORES!

tentacles/cilia create food current drawing in, passes through pharynx

Cephalochordata and pharyngeal slits (gills) which open into internal space (atrium).

Water exits body through ATRIOPORE in the Atrium.

Nerve cord gills open to interior space. Gill bars ciliated to trap.
Notochord
Cecum Intestine
Tail Endostyle is ventral

BOTH intra and extracellular digestion in digestion!

Digestive CECUM may be homologous to vertebrate liver
Mouth Endostyle Pharyngeal Atriopore Anus
gill slits closed circulatory system; contractile ventral vessel and bulbs at gill arches
instead of heart.
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Cecum: A blind-ended pouch that extends from the main digestive tract.
Digestive ceca may be the sites for final digestion of ingested food or may
be regions of the gut with specialized enzymes or conditions required for digestion

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 "Protochordates"

The dorsal hollow nerve cord runs the length of the body but is not
Cross section enlarged at the anterior end to form a brain.
(Cephalochordata)
(Cephalochordata)

Neural tube

Notochord

Myomeres muscles beginning to overlap.Smoother surface with contractions

efficient swimmer.
mucus Atrium

Gill Bars

Endostyle
Pharynx region Intestinal region
monoecious
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protonephridia:Osmoregulatory-excretory structure found in some invertebrates.

Also called a flame-cell, this tubule is closed at its distal end.
The beating of internal cilia pull water across the cell membrane and then propels it down the tubule.

filter coelomic space at top of gills, excrete urine into atrium

atriopore: The external opening to the atrium. In cephalochordates water passes

across the pharyngeal slits into the atrium and from there leaves through the
atriopore. (L. atrium, entrance hall

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