Chapter 34

Vertebrates

PowerPoint Lectures for

Biology, Seventh Edition
Neil Campbell and Jane Reece

Lectures by Chris Romero

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
 • Overview: Half a Billion Years of Backbones
• By the end of the Cambrian period, some 540 million years ago
– An astonishing variety of animals inhabited Earth’s oceans

• One of these types of animals

– Gave rise to vertebrates, one of the most successful groups of
animals

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 • The animals called vertebrates
– Get their name from vertebrae, the series of bones that make up
the backbone

Figure 34.1
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
 • There are approximately 57,000 species of vertebrates
– Which include the largest organisms ever to live on the Earth

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 • Concept 34.1: Chordates have a notochord and a dorsal, hollow
nerve cord
• Vertebrates are a subphylum of the phylum Chordata
• Chordates are bilaterian animals
– That belong to the clade of animals known as Deuterostomia
• Two groups of invertebrate deuterostomes, the urochordates
and cephalochordates
– Are more closely related to vertebrates than to invertebrates

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 • A hypothetical phylogeny of chordates
Chordates
Craniates
Vertebrates
Gnathostomes

(sister group to chordates)

Osteichthyans

(sharks, rays, chimaeras)

Cephalaspidomorphi
Lobe-fins

(frogs, salamanders)
Tetrapods

Cephalochordata

(ray-finned fishes)
Amniotes

Chondrichthyes

crocodiles, birds)
Echinodermata

(turtles, snakes,
Actinopterygii

(coelacanths)
Urochordata

(lampreys)

(lungfishes)
(hagfishes)
(lancelets)
(tunicates)

(mammals)
Mammalia
Amphibia
Actinistia

Reptilia
Myxini

Dipnoi
Milk

Amniotic egg

Legs

Lobed fins

Lungs or lung derivatives

Jaws, mineralized skeleton

Vertebral column

Head

Brain

Notochord

Ancestral deuterostome
Figure 34.2
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
 Derived Characters of Chordates
• All chordates share a set of derived characters
– Although some species possess some of these traits only
during embryonic development
Dorsal,
hollow
nerve cord

Muscle Notochord Brain

segments

Mouth

Anus
Pharyngeal
slits or clefts
Muscular,
post-anal tail
Figure 34.3
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
 Notochord
• The notochord
– Is a longitudinal, flexible rod located between the digestive tube
and the nerve cord
– Provides skeletal support throughout most of the length of a
chordate

• In most vertebrates, a more complex, jointed skeleton develops

– And the adult retains only remnants of the embryonic notochord

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 Dorsal, Hollow Nerve Cord

• The nerve cord of a chordate embryo

– Develops from a plate of ectoderm that rolls into a tube
dorsal to the notochord
– Develops into the central nervous system: the brain and the
spinal cord

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 Pharyngeal Slits or Clefts
• In most chordates, grooves in the pharynx called pharyngeal
clefts
– Develop into slits that open to the outside of the body
• These pharyngeal slits
– Function as suspension-feeding structures in many invertebrate
chordates
– Are modified for gas exchange in aquatic vertebrates
– Develop into parts of the ear, head, and neck in terrestrial
vertebrates

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 Muscular, Post-Anal Tail
• Chordates have a tail extending posterior to the anus
– Although in many species it is lost during embryonic
development

• The chordate tail contains skeletal elements and muscles

– And it provides much of the propelling force in many aquatic
species

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 Tunicates
• Tunicates, subphylum Urochordata
– Belong to the deepest-branching lineage of chordates
– Are marine suspension feeders commonly called sea squirts

• Tunicates most resemble chordates during their larval stage

– Which may be as brief as a few minutes

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 Notochord

Dorsal, hollow
nerve cord

Tail

Excurrent
siphon
Muscle
segments
Incurrent
siphon
Intestine
Stomach
Atrium

Pharynx with slits

(a) A tunicate larva is a free-swimming but

Figure 34.5 nonfeeding “tadpole” in which all four
chief characters of chordates are evident.

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 • As an adult
– A tunicate draws in water through an incurrent siphon, filtering
food particles
Incurrent
siphon
to mouth

Excurrent
siphon
Excurrent
siphon

Atrium

Pharynx
with
numerous
slits Anus
Intestine
Tunic
Esophagus
Stomach

(c) An adult tunicate, or (b) In the adult, prominent

sea squirt, is a sessile pharyngeal slits function
animal (photo is in suspension feeding,
approximately life-sized). but other chordate
Figure 34.5 (b) characters are not obvious.

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 Lancelets
• Lancelets, subphylum Cephalochordata
– Are named for their bladelike shape
Tentacle 2 cm

Mouth

Pharyngeal slits
Atrium

Notochord
Digestive tract

Dorsal, hollow Atriopore

nerve cord Segmental
muscles

Anus

Tail
Figure 34.4
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
 • Lancelets are marine suspension feeders
– That retain the characteristics of the chordate body
plan as adults

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 Early Chordate Evolution
• The current life history of tunicates
– Probably does not reflect that of the ancestral
chordate

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 • Gene expression in lancelets
– Holds clues to the evolution of the vertebrate form
BF1

Otx Hox3

Nerve cord of lancelet

embryo

BF1
Otx Hox3

Brain of vertebrate embryo

(shown straightened)
Midbrain
Figure 34.6 Forebrain Hindbrain
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
 • Concept 34.2: Craniates are chordates that have a head
• The origin of a head
– Opened up a completely new way of feeding for chordates:
active predation

• Craniates share some common characteristics

– A skull, brain, eyes, and other sensory organs

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 Derived Characters of Craniates
• One feature unique to craniates
– Is the neural crest, a collection of cells that appears near the
dorsal margins of the closing neural tube in an embryo
Neural Neural
Dorsal edges
crest tube
of neural plate Ectoderm
Ectoderm

Migrating neural
Notochord crest cells

(a) The neural crest consists of (b) Neural crest cells migrate to
bilateral bands of cells near distant sites in the embryo.
the margins of the embryonic
Figure 34.7a, b folds that form the neural tube.
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
 • Neural crest cells
– Give rise to a variety of structures, including some of the bones
and cartilage of the skull

(c) The cells give rise to some

of the anatomical structures
unique to vertebrates, including
some of the bones and cartilage
of the skull.

Figure 34.7c

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 The Origin of Craniates
• Craniates evolved at least 530 million years ago
– During the Cambrian explosion

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 • The most primitive of the fossils
– Are those of the 3-cm-long Haikouella

(a) Haikouella. Discovered in 1999 in

southern China, Haikouella had eyes
and a brain but lacked a skull, a
Figure 34.8a derived trait of craniates.
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
 • In other Cambrian rocks
– Paleontologists have found fossils of even more advanced
chordates, such as Haikouichthys
5 mm

(b) Haikouichthys. Haikouichthys had a

Figure 34.8b skull and thus is considered a true craniate.
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
 Hagfishes
• The least derived craniate lineage that still survives
– Is class Myxini, the hagfishes

Slime glands

Figure 34.9

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 • Hagfishes are jawless marine craniates
– That have a cartilaginous skull and axial rod of cartilage derived
from the notochord
– That lack vertebrae

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 • Concept 34.3: Vertebrates are craniates that have a backbone
• During the Cambrian period
– A lineage of craniates evolved into vertebrates

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 Derived Characters of Vertebrates
• Vertebrates have
– Vertebrae enclosing a spinal cord
– An elaborate skull
– Fin rays, in aquatic forms

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 Lampreys
• Lampreys, class Cephalaspidomorphi
– Represent the oldest living lineage of vertebrates
– Have cartilaginous segments surrounding the notochord and
arching partly over the nerve cord

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 • Lampreys are jawless vertebrates
– Inhabiting various marine and freshwater habitats

Figure 34.10

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 Fossils of Early Vertebrates
• Conodonts were the first vertebrates
– With mineralized skeletal elements in their mouth and
pharynx

Dorsal view
of head

Dental
Figure 34.11 elements

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

 • Armored, jawless vertebrates called ostracoderms
– Had defensive plates of bone on their skin

Pteraspis

Pharyngolepis

Figure 34.12
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings
 Origins of Bone and Teeth
• Mineralization
– Appears to have originated with vertebrate mouthparts

• The vertebrate endoskeleton

– Became fully mineralized much later

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings