CHAPTER 29

Support, Protection,
and Movement
Powerpoints revised by Franklyn Tan Te

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Copyright McGraw-Hill Education. All rights reserved. No reproduction or distribution without the prior written consent of
McGraw-Hill Education.

An ant carries with ease a flower petal that is heavier than the
ants body weight.
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Grasshoppers and Superman

Grasshoppers can jump to a height of

50 times the length of their body
But their muscles are no more powerful
than human muscles
Muscles of small and large animals exert
the same force per cross-sectional area
Grasshoppers leap high in proportion to
their size because they are small and not
because they have extraordinary muscles

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Integument

Integument is a protective outer

covering that includes skin, hair, setae,
scales, feathers, and horns
Tough and pliable to provide mechanical
protection against abrasion and puncture
Provide moisture proofing against water
loss or gain
Effective barrier against bacterial invasion
Protect underlying layers from the
damaging effects of ultraviolet light
Have other important regulatory functions

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Integument
The integument of endothermic animals
Contributes to temperature regulation
Contains sensory receptors to provide essential
information about the environment
Has excretory and respiratory functions
Assists in camouflage and signaling or display
(pigmentation)
Secretes molecules that may play role in mate
attraction, predator repulsion, and detection of
pheromonal cues that influence behavioral
interactions between animals

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Integument

Invertebrate Integument
Unicellular eukaryotes have either delicate
plasma membranes or a protective pellicle
Most invertebrates have complex tissue
coverings and some secrete a noncellular
cuticle over epidermis
Parasitic platyhelminths have syncytial
tegument that is resistant to immune
response of host and to digestive enzymes
Molluscs have soft epidermis, which
contains mucous glands that secrete
calcium carbonate shell

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Integument
Cephalopods have more a complex
integument made of cuticle; simple
epidermis, connective tissue, and
reflective cells called iridocytes
Arthropods have the most complex
invertebrate integuments that provide
protection and skeletal support

Developed a firm exoskeleton and jointed

appendages that allow for muscle attachment
Led to the extraordinary diversification of this
phylum to become the largest and most varied
of all animal groups

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Integument

Arthropods have single-layered epidermis

called hypodermis that secretes a complex
cuticle in two zones on the body
Thicker, inner zone called procuticle made of
protein and chitin that is deposited in layers
called lamellae
Thinner, outer zone called epicuticle is
nonchitinous, made of proteins and lipids, and
provides moisture-proofing barrier

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Arthropod cuticle is one of the toughest

animal materials that is resistant to
pressure, boiling, and tearing but is very
light and pliable

Integument

Cuticle may remain soft and flexible, like in

many small crustaceans and insect larvae,
but can also be found hardened in two
general ways
Decapod cuticle is hardened by deposition of
calcium carbonate in the outer layer of the
procuticle called calcification
Insect cuticle hardens by sclerotization, which
is the formation of stabilizing cross-linkages
between the protein molecules of the
procuticle lamellae

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This results in sclerotin, which is very resistant to

damage and insoluble in water

Figure 29.1
Integumentary
systems of animals,
showing
the major layers.
A, Arthropod body
wall B, Amphibian
integument. C,
Human integument.
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Integument

Since arthropod cuticle is so tough,

molting must occur to allow for growth
Molting starts with epidermal cells dividing by
mitosis
Enzymes secreted by epidermis digest most of
the procuticle
Digested materials are absorbed and reused
Space beneath the old cuticle grows new
epicuticle and procuticle are formed
After old cuticle is shed, new cuticle is
thickened and calcified or sclerotized

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Integument

Basic vertebrate integument and its

derivatives are best exemplified by frog
and human skin
Thin, outer stratified epithelial layer called
epidermis is derived from ectoderm

Inner, thicker layer called dermis is derived

from mesoderm

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Gives rise to hair, feathers, claws, and hooves

Composed of dense connective tissue that

contains blood vessels, collagenous fibers,
nerves, pigment cells, fat cells, and unique
connective tissue cells called fibroblasts

Integument

Epidermis is made of several layers of

stratified squamous epithelium
Basal layer of cells undergo frequent mitosis to
renew cell layers lying above so new cells are
made and old cells are displaced upward
Exceedingly tough fibrous protein called keratin
accumulates in the interior of cells in the
process called keratinization
As old cells die, keratin accumulates in all the
cytoplasm of the cells and become cornified
Stratum corneum is formed when highly
cornified cells become thick and highly resistant
to abrasion and water diffusion that eventually
forms calluses and footpads of mammals

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Figure 29.2 Integument of

bony fishes and lizards.

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Integument
Dermis supports, cushions, and nourishes
the epidermis
May contain true bony structure of dermal
origin much like scales of modern fishes,
which evolved from bony armor of early
fishes

Most amphibians lack dermal bones in their

skin but have vestiges of dermal scales
In reptiles, dermal bones form the armor of
crocodilians, the beaded skin of lizards, and
the shell of turtles
For some mammals, dermal bone gives rise to
antlers and the bony cores of horns

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Integument

Claws, beaks, nails, and horns are made

up of combinations of epidermal and
dermal components
Most mammalian horns, nails, and claws have
a bony core covered by vascularized nutritive
layer of dermis
The outer epithelia layer has germinative role
for continued growth and keratinization
Overgrowth is prevented by constant use that
leads to wear and abrasion

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29-17

Figure 29.3 Similarity of structure of integumentary

derivatives. Claws, beaks, and horns are all built of
similar combinations of epidermal (keratinized) and
dermal components.

Integument

Animal coloration tends to be vivid and

dramatic when used for warning coloration or
as recognition markers
Some colors are subdued or cryptic for
camouflage and disguise
Integumentary colors can be from pigments
or from surface structures
Structural colors are reflected by surface tissues
that make bright iridescences and metallic hues
as seen in insects and fish
In birds, small air-filled spaces or pores in
feathers can reflect a portion of color spectrum
while other animals use thin films to angle light

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Integument
Pigments (biochromes) are molecules that
reflect specific light rays and are more
commonly used in the animal kingdom
Crustaceans and ectothermic vertebrates have
pigments that are in large cells with branching
processes called chromatophores
Pigments may concentrate in center of cell or
be dispersed throughout cell
Cephalopod chromatophores are very different
in that small sac-like cells with pigment
granules are surrounded by muscle cells that
can stretch the cell into a pigmented sheet
When muscles relax, the elastic cell shrinks
quickly and allow rapid color changes

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Figure 29.4 Chromatophores. A, The crustacean chromatophore

dispersed ( left ) and concentrated ( right ). B, The cephalopod
chromatophore muscles contracted ( left ) to expose the pigment
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Integument
Melanin pigments are the most widespread and
are usually black or brown polymers
responsible for earth-colored shades found in
melanocytes or melanophores
Carotenoid pigments impart yellow and red
colors contained inside xanthophores
Ommochromes and pteridines are for yellow
pigments in molluscs and arthropods, while
green coloration is produced by yellow
pigment overlying blue structural color
Iridophores contain crystals of guanine or
other purines, not pigments, that produce
silvery or metallic colors when these reflect
light

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Integument
Mammals are relatively somber-colored
(uncolorful) and most species are colorblind
However, some primate species of baboons
and mandrills have brightly colored skin
patches since these have color-vision
Dermal melanophores deposit melanin in
growing hair of mammals and give the general
dull colors of most mammalian species

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Integument

Injurious effects of sunlight like human

sunburn demonstrates the damaging
effect of ultraviolet radiation on cells
Flatworms exposed to sun in shallow water
are damaged or killed
Arthropod cuticle, scales of reptiles,
feathers of birds, and fur of mammals
provide screening action against the sun
Humans are naked apes and lack furry
protection

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Depend on thickening of the stratum corneum

and epidermal pigmentation

Integument
The epidermis absorbs most of the ultraviolet
radiation but about 10% penetrate the dermis
Sunburn is from blood-vessel enlargement due
to release of histamine and other vasodilator
substances
A sun tan results from increased melanin
secretion in the dermis and from pigment
darkening in the epidermis
Ultraviolet radiation causes about one million
new cases of skin cancer annually in the U.S.
High doses of ultraviolet radiation in childhood
may result in genetic mutations that cause skin
cancer later in life

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Skeletal Systems

Skeletons provide support, rigidity,

surface for muscle attachment, and
protection of delicate body organs

Hydrostatic skeletons of invertebrates are

not rigid and use their body fluids
Muscles in the body wall of earthworms
contract against the incompressible coelomic
fluids enclosed in a limited space
Alternative contraction of muscles enable worm
to be thin or thick and produce backwardmoving waves of motion to propel the animal
forward using tiny bristles (setae) as anchors

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Figure 29.5 Earthworm movement.

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Skeletal Systems

The elephant trunk, tongues of mammals

and reptiles, and tentacles of cephalopods
are examples of structures that lack any
obvious skeletal support but is capable of
bending, twisting, and lifting heavy objects

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Septa separate the worm body into

compartments that allow each part to develop
pressure even when other compartments are
punctured or cut into pieces

These structure are called muscular hydrostats

because of incompressible tissues that remain
at constant volume and arranged in complex
patterns

Figure 29.6
Muscular trunk
of an elephant,
an example of a
muscular
hydrostat.

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Skeletal Systems

Rigid skeletons provide firm elements

to which muscles can attach and serve
as anchor points for the extension and
flexion of muscle movement

Two types of rigid skeletons are the outer

exoskeleton and inner endoskeleton
Exoskeleton is typical of mollusks, arthropods,
and other invertebrates that will have shells,
spicules, and calcareous, proteinaceous or
chitinous plates
Can be for protection and locomotion but has
to be periodically molted since it does not grow
with the animal in many cases

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Skeletal Systems
Endoskeleton is found in echinoderms, some
cnidarians and vertebrates
Vertebrate endoskeleton is formed inside the
body and composed of specialized connective
tissue like bone and cartilage
Functions as protection and support but also
as reservoir for calcium and phosphorus
In amniotic vertebrates, the bone marrow
makes red blood cells, white blood cells, and
platelets

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Skeletal Systems

Notochord is a semi-rigid supportive axial rod

of protochordates and all vertebrate larvae

Made of large vacuolated cells surrounded by

layers of elastic fibrous collagen sheaths
Act as stiffening structure to maintain body
shape during locomotion

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Notochord surrounded or replaced by vertebrae

during development except in jawless
vertebrates like lampreys and hagfishes

Cartilage is a major skeletal element of jawless

fishes and elasmobranchs

Skeletal Systems

Most vertebrates have bony skeletons with

some interspersed cartilage

Cartilage is soft, pliable tissue that resists

compression with a basic form called hyaline
cartilage that is clear and glassy in form

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This type of cartilage is made of cartilage cells called

chondrocytes that are surrounded by a firm complex
protein-carbohydrate gel interlaced with a meshwork
of collagen fibers
Forms most of the cartilaginous skeletons of
vertebrate embryos, the articulating surfaces of
joints and the supportive structures for tracheal,
laryngeal and bronchial rings; most have no blood
vessels so heal slowly

Skeletal Systems

Other types of cartilage include elastic and

fibrous tissues that form bundles and are
arranged in herringbone design
Cartilage structure similar to hyaline cartilage
are found in the radula of gastropods and
lophophore feeding structures of brachiopods
Cephalopods have specialized cartilage with
long, branching processes that resemble cells
of vertebrate bone

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Skeletal Systems

Bone is living tissue with significant deposits

of inorganic calcium salts in an extracellular
matrix of collagen fibers in proteincarbohydrate gel

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Highly vascular and capable of rapid healing,

unlike cartilage
Nearly as strong as cast iron, yet only one-third as
heavy due to its structural organization
Bone never forms in vacant space but is laid down
by the replacement of connective tissue

Skeletal Systems

Most bone develops by replacing hyaline

cartilage called endochondral bone
replacement (within cartilage)
Embryonic cartilage is eroded and becomes
honeycombed; it is then filled with boneforming cells
Extracellular bone matrix gets deposited and
later calcified around strand-like remnants of
the cartilage

Intramembranous bone comes directly

from sheets of embryonic cells

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Forms face, cranium, and clavicle while the rest

of the skeleton is endochondral bone

Skeletal Systems

Fully formed bone varies in density and

comes in two types- spongy (cancellous)
and compact (lamellar) bones
Spongy bone usually has open interlacing
framework of bony tissue that is oriented to
give maximum strength under normal stress
and strains
Compact bones are denser bones formed from
spongy bones through further deposition of
bone matrix arranged in concentric rings
Both spongy and compact bones form the long
bones of tetrapods

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29-37

Figure 29.7 Structure of compact bone. A, Adult long bone with a

cut into the medullary cavity. B, Enlarged section showing osteons
C, Enlarged view of an osteon D, An osteocyte within a lacuna.

Skeletal Systems

Microscopic structure of bone consists

of bundles of osteons cemented
together with interconnected blood
vessels and nerves

Osteons (Haversian system) are elongated

cylindrical ring structures containing
organized lacunae and canaliculi

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Lacunae are cavities between the rings that

contain bone cells called osteocytes, which are
connected by minute passages called
canaliculi that allow nutrients and growth
factors to be distributed throughout the bone

Skeletal Systems

Blood vessels and nerves interconnect

within the bones and allow rapid healing
Bone is a dynamic tissue such that bone
growth and remodeling are complex
restructuring processes
Osteoclasts slowly resorb bone (cell
destruction) while osteoblasts deposit new
additional bone (cell building)
These simultaneous processes allow growth of
the bone without any weakening
Bone marrow cavity grows larger by bone
resorption of the inner surfaces of surrounding
bone with new bone being laid down on the
outer surface by bone deposition

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Skeletal Systems

Bone growth responds to several hormones

Bone growth responds to usage much like

muscles

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Parathyroid hormone stimulates bone resorption

Calcitonin is a hormone from thyroid gland that inhibits
bone resorption
Both hormones and vitamin D3 (1,25-dihydroxyvitamin
D3), maintain constant blood calcium levels

When astronauts have been living without gravity for

some time, they can suffer bone loss and weakness

Skeletal Systems

Plan of the vertebrate skeleton has

both the axial and appendicular
skeletal divisions
Axial skeleton includes the skull, vertebral
column, sternum, and ribs
Appendicular skeleton are bones of limbs
along with pectoral and pelvic girdles
Movement from water to land forced
dramatic changes in body form
Increased cephalization made the skull the
most intricate part of the skeleton

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Figure 29.8 Skeleton of a perch.

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Skeletal Systems

Vertebrate skulls have increased

concentration of brain, sense organs, and
food gathering apparatus

Some early fishes had 180 skull bones but over

time, many skull bones were lost or fused and
now are greatly reduced

Amphibians have from 50 to 95, mammals have 35 or

fewer while humans have 29

Vertebral column is the main stiffening axis

and serves as points for muscle
attachment while preserving body shape
Movement from water to land implies that
body is no longer supported by water
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Skeletal Systems

Vertebral column became structurally

adapted to withstand new regional
stresses given by the two pairs of limb
appendages
Amniotic tetrapod vertebrae are separated into
cervical (neck), thoracic (chest), lumbar (back),
sacral (pelvic), and caudal (tail) vertebrae
In frogs, birds, and humans, the caudal
vertebrae reduced in size and number while the
sacral vertebrae are fused
The python has over 400 vertebrae while the
human child has 33 vertebrae and the adult has
5 fused to form the sacrum and 4 form the
coccyx or tail bone

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Figure 29.9 Human

skeleton. A, Ventral
view. B, Dorsal view.
29-45

Skeletal Systems
Humans also have 7 cervical vertebrae, 12
thoracic vertebrae, and 5 lumbar vertebrae
Nearly all mammals have 7 cervical
vertebraefrom the short necks of
dolphins to the long necks of giraffes
The first two cervical vertebrae are present
in all vertebrates

Atlas is the 1st cervical vertebra and supports

the skull while allowing it to pivot
Axis is the 2nd cervical vertebra and allows the
head to turn side-to-side

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Skeletal Systems

Ribs are either long or short skeletal

structures that articulate medially with the
vertebrae and extend into the body wall
Fishes have single or paired of ribs for every
vertebra that serve as stiffening rods for
improved effectiveness of muscle contractions
Many fishes have both dorsal and ventral ribs,
some with numerous rib-like intermuscular bones
Some vertebrates have reduced ribs

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The Leopard frog that has no ribs at all

Mammalian ribs form a thoracic basket to prorect

the heart, lungs, and other soft body parts

Skeletal Systems
Sloths have 24 pairs of ribs, while horses have
18 pairs
Primates other than humans have 13 pairs of
ribs
Humans generally have 12 pairs; some have a
rare 13th pair

Most vertebrates and fishes have paired

appendages
Most fishes, except agnathans, have pectoral
and pelvic girdles supporting thin pectoral and
pelvic fins
Some eels lack pectoral or pelvic fins while the
Moray eels are lacking in both

29-48

Skeletal Systems
Tetrapods, unless they are limbless, have
two pairs of pentadactyl limbs (five-toed)
that are supported by respective girdles
The pentadactyl limb is similar in all
tetrapods , alive or extinct, even when
these are highly modified for various
modes of life
Modifications due to different living
environments often involve bone loss or
fusion rather than addition of new bones

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The ends of the appendages like fingers and

toes are more likely to change

Skeletal Systems
Horses and their relatives have gained speed
by elongation of a longer third toe with the
horse standing on its third finger nail (hoof)
Bird embryos demonstrate distal modification
where 13 distinct wrist and hand bones
(carpals and metacarpals) and finger bones
(phalanges) are reduced to only 4 bones in 3
digits as found in the adults
In tetrapods, the pelvic girdle is firmly attached
to the axial skeleton and absorb the greatest
locomotory force transmitted by hind limbs
The pectoral girdle is more loosely attached to
provide forelimbs with greater freedom for
manipulation

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Skeletal Systems

Effect of body size on bone stress

follows what Galileo realized in 1638:
The ability of limbs to support a load
decreases as the animals increase size
Consider one animal twice as long, wide,
and tall as a second animal:

The larger animal has eight times the volume

and eight times the weight
The strength of the legs, however, is based on
the cross-sectional area of bones, tendons and
muscles, which is only four times greater
Therefore, eight times the weight is to be carried
by four times the strength

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Skeletal Systems

Mammalian bone is uniform per crosssectional area, which places an upper limit
on overall size
Bone shape in different sized animals does not
change much, so many mammals adapted by
shifting limb posture to align with body axis
Bones and muscles are capable of carrying
more weight when aligned closely with the
ground reaction force, as in the horses leg
Peak bone stresses during strenuous activity is
no greater for a galloping horse than for a
running chipmunk
Elephants and large dinosaurs had thick and
robust bones but this decreases running speed

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Figure 29.10 Comparison of postures in small and

large mammals, showing the effect of scale.

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Animal Movement

Movement is an important characteristic of

animals and includes streaming of cytoplasm
and massive muscular movements

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Animal movement relies on a fundamental

mechanism called contractile proteins that
facilitate relaxation and contraction
Contractile machinery is composed of ultrafine
fibrils that are arranged to relax or contract when
powered by ATP
The most important protein contractile system is
composed of actin and myosin

Animal Movement

The actomyosin system is almost a universal

biomechanical system

Found from protozoa to vertebrates with a diverse

set of functional roles

However, cilia and flagella are composed of

proteins other than actin and myosin
The three principal types of animal movement
are

Ameboid
Ciliary and flagellar
Muscular

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Animal Movement
Ameboid movement:
Occurs in wandering cells of metazoans
Is characteristic of amebas and other
unicellular forms
Also occurs in macrophages, white blood
cells, embryonic mesenchyme, and other
mobile cells that move in tissue spaces
Ameboid cells change shape by extending
and withdrawing pseudopodia (false feet)

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Non-granular gel-like ectoplasm that encloses a

more liquid endoplasm

Animal Movement

Movement depends on actin, actin-binding

proteins, and other regulatory proteins
Under one model, as the pseudopod extends,
hydrostatic pressure forces actin subunits into
the pseudopod, the subunits dissociate from
actin-binding proteins and reassemble into a
network to form a gel-like ectoplasm
At trailing edge of the gel, the network
disassembles; actin filaments interact with
myosin to create a contractile force that pulls
the cell along behind the extending pseudopod
Locomotion is assisted by membrane-adhesion
proteins that attach temporarily to substrate,
providing traction that enables the cell to crawl

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Animal Movement
Ciliary and flagellar movement
Are

particularly distinctive for ciliated

unicellular eukaryotes
Occur in all animal groups except nematodes
and arthropods

Cilia are minute, hairlike, motile processes that

extend from surfaces of many animal cells

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Function to move whole unicellular organisms and

ctenophores in their aquatic environments

Also used to propel fluids and materials across

epithelial surfaces in larger animals

Animal Movement

All cilia have a uniform diameter of 0.2 to

0.5 micrometers and a basal body
(kinetosome) similar to centrioles
Basal body gives rise to peripheral circle of 9+2
arrangement of microtubules, forming the
structural support and machinery of ciliary
movement
Microtubules are composed of a spiral array of
tubulin protein subunits
Microtubule doublets around the periphery are
connected to each other and the central pair by
microtubule-associated proteins (MAPs)
Extending from each doublet is a pair of arms
composed of the MAP called dynein

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Animal Movement
Dynein arms act as cross bridges between
doublets and produce a sliding force between
microtubules
During ciliary movement, microtubules behave
like sliding filaments that move past one
another much like vertebrate muscular action
During ciliary flexion, dynein arms link to
adjacent microtubules then swivel and release
in repeated actions, causing the microtubules
on one side to slide outwards past the
microtubules on the other side
During the recovery stroke, the microtubules of
the opposite side slide outward to bring the
cilia back into its starting position

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Figure 29.11 A, Longitudinal and cross

section of a cilium showing the
microtubules and microtubuleassociated proteins (MAPs)
B, Electron micrograph of section
through several cilia. (x133,000)
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Animal Movement

Flagella occur in flagellated protozoans, animal

spermatozoa, and in sponges
The flagellum is whiplike, longer than a cilium, and
occurs singularly or in small numbers at one end of the
cell
Flagella have the same basic internal structure as cilia
although several exceptions to this 9+2 arrangement
are in the sperm tails of flatworms (9+1) and mayflies
(9+0)
Cilia and flagella differ more in their beating patterns
than in structure
A flagellum beats symmetrically with snakelike
undulations to propel water parallel to the long axis of
the flagellum

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Figure 29.12 A, Flagellum beats in wavelike undulations, propelling

water parallel to its main axis B, Movement of cilia in comb plates
of a ctenophore.
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Animal Movement

A cilium beats asymmetrically with a fast power

stroke in one direction followed by a slow
recovery, wherein water is propelled parallel to
the ciliated surface

Muscular movement occurs via

contractile tissue called muscle fibers
and are the most highly developed form
of muscle cells
Muscle fibers generally work only by
contraction and cannot actively lengthen
Can be arranged in a variety of ways to make
every movement possible

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Animal Movement

The types of vertebrate muscles are

characterized by appearance: skeletal,
cardiac, and smooth muscles

Skeletal (striated) muscles are transversely

striped with alternating dark and light
bands (striations) and are multinucleated
Organized into sturdy, compact bundles that are
mostly attached to skeletal elements that move
the trunk, appendages, eyes, respiratory organs,
mouthparts, and other body structures
Muscle fibers are long cylindrical cells that are
packed together in bundles called fascicles and
enclosed by tough connective tissue

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Figure 29.13 Photomicrographs of

types of vertebrate muscle. A, Skeletal
muscle (human) showing several
striated fibers B, Cardiac muscle
(monkey) is striated, similar to skeletal
muscle, C, Smooth muscle (human)
showing absence of striations.

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Animal Movement
Fascicles are grouped into a discrete muscle
enclosed by another layer of thick connective
tissue
Some muscles taper at ends as they connect to
bones via tendons, while others are flattened
sheets as in abdominal muscles
Skeletal muscles can contract powerfully and
quickly, but fatigues more rapidly than smooth
muscle

29-67

Generally called voluntary muscle as these are

stimulated by motor fibers under conscious control

Figure 29.14 Organization of skeletal muscle from gross to

molecular level.
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Animal Movement

Cardiac muscle is the tireless muscle of

the vertebrate heart; it shares some
characteristics of skeletal muscle
Made of closely opposed, separated
uninucleated cells joined by junctional
complexes within vertical bars called
intercalated discs
Fast acting, striated, and with similar
contraction mechanism as skeletal muscle but
has involuntary autonomic and hormonal
control like smooth muscle
Heart beat originates within a specialized
cardiac muscle and can continue to beat
outside of the body

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Animal Movement

Smooth (visceral) muscle lacks striations

and the cells are much smaller, tapering at
both ends with a single, central nucleus
Cells interdigitate with each other such that the
tapered ends of one cell is next to the central
nuclear region of the next cell
Smooth muscle cells form sheets of muscle
circling the walls of the alimentary canal, blood
vessels, respiratory passages, and urinary and
reproductive ducts and cavities
Usually slow acting and can maintain prolonged
contractions with little energy use
Normally controlled by the autonomic nervous
system, hormones and local regulatory systems

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Animal Movement
Smooth muscle contractions are involuntary
and unconscious; function by sustained
contractions and relaxations
Most smooth muscles push material through a
tube like the intestines or regulate tube diameter
during blood flow or air flow

Types of invertebrate muscles include

variations of smooth, striated, and
oblique striated muscles

Striated muscles are found in a variety of

cnidarians and arthropods

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Giant barnacles and Alaskan king crabs have

giant muscle fibers that 3 mm wide by 6 cm long

Animal Movement

Bivalve molluscan muscle fibers are of two

types- striated and smooth muscles
Scallops use fast striated muscle fibers to
close valves during swimming actions as these
can contract rapidly
A slower smooth muscle is able to sustain
long-lasting contraction for hours or days
Slow adductors use very little energy and
require very little neural stimulation to remain
contracted
The contracted state resembles a catch
mechanism with low rate of cross-bridge
cycling between contractile proteins within the
muscle fibers

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Animal Movement

Insect flight muscles can beat their wings

more than 1,000 beats per second
This fibrillar muscle contracts at frequencies
much faster than any vertebrate muscle but has
limited extensibility
The wing leverage system is arranged so the
muscles shorten very little during each
downbeat; both muscles and wings operate as a
rapidly oscillating system in an elastic thorax
Muscles recoil elastically, are activated by
stretch during flight, and require excitatory
neural signals periodically with one signal per
20 to 30 contractions

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Animal Movement

Structure of striated muscle is from the

periodic bands visible under the light
microscope

Each cell or fiber is a multinucleated tube

with many myofibrils packed together and
surrounded by a cell membrane called the
sarcolemma
Myofibrils contains two filaments of proteins
called myosin and actin
Actin extends in parallel filaments from a dense
protein complex called the Z line
Sarcomere is the functional unit of myofibrils,
extending between successive Z lines

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Animal Movement

Myosin filament is composed of myosin

molecules packed together in a bundle
Each myosin molecule consists of two
polypeptide chains, each with a club-shaped
head region
Two myosin bundles are held end-to-end at the
center of each sarcomere
The double heads of each myosin molecule
face outward from the center of the filament
and point towards the Z line
The heads act as binding sites for ATP and
form molecular cross bridges that interact with
the actin filaments during muscle contraction

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Figure 29.15 Molecular structure of actin and myosin filaments of

skeletal muscle A, The myosin molecule. B, The myosin filament
extended towards actin filaments. C, The actin filament.
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Animal Movement

Actin filaments are made of a backbone of

double stranded actin twisted into a
double helix and two actin-binding
proteins called tropomyosin and troponin
Each tropomyosin is a double helix that lies
near the grooves between the actin strands
Troponin is a complex of three globular
proteins located at intervals along the actin
filament that act as a calcium-dependent
switch that control contraction
Actin filament complexes extend outward from
both sides of the Z line and overlap myosin
bundles towards the sarcomere

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Animal Movement

Sliding filament model of muscle

contraction was independently
proposed in the 1950s by A.F. Huxley
and H.E Huxley to explain contraction

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Nebulin regulates actin length and the elastic

protein titin support and anchor the myosin to
the middle of the sarcomere at the M line

The actin and myosin filaments link

together by molecular cross bridges and
then act as levers to pull the filaments past
each other

Animal Movement
The club shaped heads of the myosin filaments
form cross bridges that snap rapidly back and
forth, attaching and releasing from special
receptor sites on the actin filaments
This ratcheting action draws the actin past the
myosin and pulls the Z lines together
All sarcomere units shorten together as the
muscle contracts
Relaxation is passive: when the cross bridges
between the filaments release, the sarcomeres
are free to lengthen
This requires some force, usually supplied by
recoil of elastic fibers within the muscle, and
by antagonistic muscles or by gravity

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Figure 29.16 Sliding-filament hypothesis, showing how actin

and myosin filaments interact during contraction. A, Muscle
relaxed. B, Muscle contracted.
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Animal Movement

Muscles contract in response to nerve

control and stimulation

If a nerve to a muscle is severed, the muscle

atrophies or wastes away

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Skeletal muscles are innervated by motor neurons

whose cell bodies are located in the central nervous
system (brain and spinal cord)
Each cell body leads to a motor axon that leaves the
brain and spinal cord, travels through the peripheral
nerve trunk, and branches to many terminal points on
a muscle
Each terminal branch innervates a single muscle fiber,
while a single motor axon may innervate a few fibers
for precise control like the eye

Animal Movement
Large muscle groups like the leg can have
single motor axons controlling up to 2,000
muscle fibers
A motor neuron and all of the muscle fibers it
innervates are called a motor unit
When a motor neuron fires, the action potential
passes simultaneously to all motor units and
each is contracted simultaneously

Precise control of movement requires varying

number of motor units activated at one time

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The total force of muscle contraction depends on the

number of motor units activated

Motor unit recruitment is a steady increase in muscle

tension by increasing motor units

Animal Movement

The neuromuscular (myoneural) junction

is the place where the motor axon
terminates on a muscle fiber

At this junction, there is a synaptic cleft,

which separates the nerve terminal from the
muscle fiber
Neuron stores acetylcholine in small synaptic
vesicles near the synaptic cleft
When the nerve signal or action potential reaches
the synapse, the acetylcholine is released and acts
as chemical neurotransmitter
Acetylcholine causes depolarization of the muscle
fiber membrane by binding to receptor sites and
induces muscle contraction

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Animal Movement
Synapse provides a chemical bridge that
couples the nerve impulse and muscle fibers
Vertebrate skeletal muscles have elaborate
conduction system that carries the
depolarization from the neuromuscular junction
to the densely packed filaments of the muscle
fiber
Numerous invaginations on the sarcolemma
surface project into muscle fibers called Ttubules

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T-tubule system is continuous with the sarcoplasmic

reticulum (specialized endoplasmic reticulum) that
runs parallel to the actin and myosin filaments, where
it releases calcium to enable muscle contraction

Figure 29.17 Section of vertebrate skeletal muscle showing a

nerve-muscle synapse (neuromuscular or myoneural junction)
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Animal Movement

Excitation-contraction coupling occurs

when electrical depolarization of the
sarcolemma and t-tubules activate the
contractile system

In resting muscle, contraction does not

occur since the tropomyosin strands
around the actin filaments block the myosin
heads from attaching to actin
When muscle is stimulated, the action potential
is transmitted down the t-tubules
Electrical depolarization causes Ca+2 to be
released from sarcoplasmic reticulum; it then
binds with troponin

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Animal Movement
Troponin changes shape, shifts the
tropomyosin out of its blocking position, and
allows active sites on actin filaments to be
exposed
Myosin heads binds to active site and form
cross bridges between the adjacent myosin
and actin filaments

Muscle action follows a series of attachpull-release cycle or cross-bridge cycling

Release of bond energy from ATP activates the
myosin head that swings 45 degrees and
releases a molecule of ADP
This power stroke pulls the actin filament
about 10 nanometers in distance

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Animal Movement
Movement stops when another ATP molecule
binds to the myosin head and thus freeing it
from the active site
Shortening continues as long as nerve action
potential arrive at the neuromuscular junction
and that free calcium remains available
The cross-bridge cycling can repeat 50100
times per second in pulling actin and myosin
filaments past each other
Each sarcomere shortens a very small distance

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This distance is multiplied by the thousands of

sarcomeres lying end-to-end in muscle fiber, such that
strongly contracting muscle may shorten by one-third
of its length

Animal Movement
When stimulation stops, calcium is pumped
back into sarcoplasmic reticulum
Troponin resumes its original configuration
and tropomyosin moves back to its blocking
position on the actin, leading to muscle
relaxation

Large amounts of energy is needed for

muscle contraction

ATP is the immediate source of energy and

is normally present at constant levels

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Aerobic metabolism catabolizes glucose that is

transported through the blood to produce ATP

Figure 29.18 Excitationcontraction coupling in

vertebrate skeletal
muscle. Step 1: An
action potential spreads
along the sarcolemma
Step 2: Myosin forms
cross bridges Step 3:
Using ATP, the myosin
head swings toward the
center of the sarcomere
Step 4: The myosin
head binds another ATP
Step 5: The myosin
head splits ATP,
retaining the energy.
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Animal Movement

Glycogen is stored within the muscles to

supply glucose for ATP production
It is a polysaccharide chain of glucose
molecules that is stored in the liver and muscles
Abundant as more than three-fourths of all
glycogen is stored in muscles
Can be mobilized quickly to provide energy in
both aerobic and anaerobic conditions

Muscles also have an energy reserve called

creatine phosphate that is a high-energy
phosphate compound stored during rest
Creatine phosphate releases stored bond
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energy to convert ADP to ATP

Animal Movement

Different muscle types, like slow and fast

oxidative fibers, rely heavily on glucose
and oxygen transported by blood
If muscle contraction is not too vigorous or
prolonged, glucose is completely oxidized to
CO2 and water by aerobic respiration
During prolonged heavy exercise, blood flow
cannot provide enough oxygen for complete
oxidation of glucose

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Muscles must then rely on energy from anaerobic

glycolysis

Anaerobic glycolysis is not as efficient as

aerobic respiration but is needed for all forms
of heavy muscular exertion

Animal Movement

Fast glycolytic fibers rely exclusively on

anaerobic glycolysis for energy
Anaerobic glycolysis degrades glucose to
lactic acid to release energy
Lactic acid accumulates in the muscle and
diffuses rapidly into general circulation
Continued muscular exertion causes a buildup
of lactic acid that leads to muscle fatigue that
was initially thought to be due to decreased pH
and enzyme inhibition
Recent evidence suggests that muscle fatigue
in muscles relying on creatine phosphate may
be due to accumulation of inorganic phosphate

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Animal Movement
Anaerobic pathway is self-limiting; continued
heavy exertion leads to exhaustion
Muscles incur an oxygen debt because
accumulated lactic acid must be converted to
pyruvic acid, which can be fed into the Krebs
Cycle via conversion to Acetyl-CoA
Lactic acid is then oxidized by extra oxygen as
the oxygen debt gets repaid via increased
oxygen consumption, even after muscle
exertion has ended
Oxygen replenishment continues until all lactic
acid has been oxidized in the body and
glycogen is resynthesized

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Animal Movement

Muscle performance depends on the

type of muscle fiber

Slow oxidative fibers (red muscle fibers)

are specialized for slow, sustained
contractions without fatigue
These fibers function in postural muscles
Contain extensive blood supply (red color), a
high density of mitochondria for constant ATP
supply, and abundant stored myoglobin that
supplies additional stored oxygen

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Two kinds of fast fibers capable of fast,

powerful contractions but with different
energy production pathways

Animal Movement

Fast glycolytic fibers (white muscles) lack

efficient blood supply, have low density of
mitochondria and myoglobin, and thus
fatigue easily when used
Usually pale in color and function anaerobically,
as exemplified by the white meat of chicken
breast and running muscles of cats
During a chase, such muscles rapidly develop an
oxygen debt in less than a minute

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Cheetahs must rest 3040 minutes after a chase

Human weight lifters favor these muscles and

can not sustain lifting heavy objects for long
periods of time

Animal Movement
Fast

oxidative fibers have extensive

blood supply and high density of

mitochondria and myoglobin
Functions aerobically and is used for rapid,
sustained activities since the muscles are
fatigue resistant
Migratory birds, like geese and swans, dogs,
and ungulates have limb muscles with a high
percentage of fast oxidative fibers capable of
active locomotion for long periods
Most muscles possess a mixture of the three
different types of muscle fibers to permit a
wide range of activities

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Animal Movement

Importance of tendons in energy

storage is due to stored elastic strain
energy during walking and running

The Achilles tendon is stretched by the

combination of downward force of the body
and the contraction of the calf muscles
As tendon recoils, this extends the foot while
the muscle is still contracted and propels the
leg forward
Kangaroo uses the recoil energy in tendons to
bounce along called the bouncing ball
principle

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Animal Movement

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This bouncing ball type of movement uses

far less energy than would be required if every
step relied solely on alternate muscle
contraction and relaxation
Elastic storage of energy also occurs in the
legs of grasshoppers and fleas, wing hinges
of flying insects, hinge ligaments of bivalve
molluscs, and the dorsal ligament
(ligamentum nuchae) that supports the head
of hoofed mammals during running

29-100

Figure 29.19 Energy storage in the Achilles tendon of human

and kangaroo legs.