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    Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

    PowerPoint Lectures for Biology, Seventh Edition

    Neil Campbell and Jane Reece

    Lectures by Chris Romero

    Chapter 49

    Sensory and Motor

    Mechanisms

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    Overview: Sensing and Acting

    Bats use sonar to detect their prey

    Moths, a common prey for bats

    Can detect the bats sonar and attempt to flee

    Figure 49.1

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    Both of these organisms

    Have complex sensory systems that facilitatetheir survival

    The structures that make up these systems

    Have been transformed by evolution intodiverse mechanisms that sense various stimuliand generate the appropriate physicalmovement

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    Concept 49.1: Sensory receptors transduce

    stimulus energy and transmit signals to thecentral nervous system

    Sensations are action potentials

    That reach the brain via sensory neurons

    Once the brain is aware of sensations

    It interprets them, giving the perception of stimuli

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    Sensations and perceptions

    Begin with sensory reception, the detection of stimuli by sensory receptors

    Exteroreceptors

    Detect stimuli coming from the outside of thebody

    Interoreceptors Detect internal stimuli

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    Functions Performed by Sensory Receptors

    All stimuli represent forms of energy

    Sensation involves converting this energy

    Into a change in the membrane potential of sensory receptors

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    Sensory receptors perform four functions in this

    process Sensory transduction, amplification,

    transmission, and integration

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    Two types of sensory receptors exhibit these

    functions A stretch receptor in a crayfish

    Figure 49.2a

    (a)Crayfish stretch receptors have dendritesembedded in abdominal muscles. When theabdomen bends, muscles and dendrites

    stretch, producing a receptor potential in thestretch receptor. The receptor potential triggersaction potentials in the axon of the stretch

    receptor. A stronger stretch producesa larger receptor potential and higher requency of action potentials.

    Muscle

    Dendrites

    Stretchreceptor

    Axon M e m

    b r a n e

    p o

    t e n

    t i a l ( m

    V )

    50

    70

    0

    70

    0 1 2 3 4 5 6 7Time (sec)

    Action potentials

    Receptor potential

    Weak

    muscle stretch

    50

    70

    0

    70

    0 1 2 3 4 5 6 7Time (sec)

    Strong

    muscle stretch

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    A hair cell found in vertebrates

    of action potentials in the sensory neuron.Bending in the other direction has the oppositeeffects. Thus, hair cells respond to the directionof motion as well as to its strength and speed.s

    (b) Vertebrate hair cells have specialized ciliaor microvilli (hairs) that bend when sur-rounding fluid moves. Each hair cell releasesan excitatory neurotransmitter at a synapse

    with a sensory neuron, which conducts actionpotentials to the CNS. Bending in one directiondepolarizes the hair cell, causing it to releasemore neurotransmitter and increasing frequency

    50 70

    0

    70

    0 1 2 3 4 5 6 7Time (sec)

    Action potentials

    No fluidmovement

    50 70

    0

    70

    0 1 2 3 4 5 6 7Time (sec)

    Receptor potential

    Fluid moving inone direction

    50 70

    0

    70

    0 1 2 3 4 5 6 7Time (sec)

    Fluid moving inother direction

    M e m

    b r a n e

    p o

    t e n

    t i a l ( m

    V )

    M e m

    b r a n e

    p o

    t e n

    t i a l ( m

    V )

    M e m

    b r a n e

    p o

    t e n

    t i a l ( m

    V )

    Hairs of hair cell

    Neuro-trans-mitter atsynapse

    Axon

    Lessneuro-trans-mitter

    Moreneuro-trans-mitter

    Figure 49.2b

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    Sensory Transduction

    Sensory transduction is the conversion of

    stimulus energy Into a change in the membrane potential of a

    sensory receptor

    This change in the membrane potential

    Is known as a receptor potential

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    Many sensory receptors are extremely

    sensitive With the ability to detect the smallest physical

    unit of stimulus possible

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    Amplification

    Amplification is the strengthening of stimulus

    energy By cells in sensory pathways

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    Transmission

    After energy in a stimulus has been transduced

    into a receptor potential Some sensory cells generate action potentials,

    which are transmitted to the CNS

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    Sensory cells without axons

    Release neurotransmitters at synapses withsensory neurons

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    Integration

    The integration of sensory information

    Begins as soon as the information is received

    Occurs at all levels of the nervous system

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    Some receptor potentials

    Are integrated through summation

    Another type of integration is sensoryadaptation

    A decrease in responsiveness duringcontinued stimulation

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    Types of Sensory Receptors

    Based on the energy they transduce, sensory

    receptors fall into five categories Mechanoreceptors

    Chemoreceptors

    Electromagnetic receptors

    Thermoreceptors

    Pain receptors

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    The mammalian sense of touch

    Relies on mechanoreceptors that are thedendrites of sensory neurons

    Figure 49.3

    Heat

    Light touch Pain

    Cold

    Hair

    Nerve Connective tissue Hair movement Strong pressure

    Dermis

    Epidermis

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    Chemoreceptors

    Chemoreceptors include

    General receptors that transmit informationabout the total solute concentration of asolution

    Specific receptors that respond to individualkinds of molecules

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    Two of the most sensitive and specific

    chemoreceptors known Are present in the antennae of the male

    silkworm moth

    Figure 49.4 0 . 1

    m m

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    Electromagnetic Receptors

    Electromagnetic receptors detect various forms

    of electromagnetic energy Such as visible light, electricity, and

    magnetism

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    Some snakes have very sensitive infrared

    receptors That detect body heat of prey against a colder

    background

    Figure 49.5a(a) This rattlesnake and other pit vipers have a pair of infrared receptors,

    one between each eye and nostril. The organs are sensitive enoughto detect the infrared radiation emitted by a warm mouse a meter away.The snake moves its head from side to side until the radiation is detectedequally by the two receptors, indicating that the mouse is straight ahead.

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    Many mammals appear to use the Earths

    magnetic field lines To orient themselves as they migrate

    Figure 49.5b

    (b) Some migrating animals, such as these beluga whales, apparentlysense Earths magnetic field and use the information, along withother cues, for orientation.

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    Thermoreceptors

    Thermoreceptors, which respond to heat or cold

    Help regulate body temperature by signalingboth surface and body core temperature

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    Pain Receptors

    In humans, pain receptors, also callednociceptors

    Are a class of naked dendrites in the epidermis

    Respond to excess heat, pressure, or specificclasses of chemicals released from damagedor inflamed tissues

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    Concept 49.2: The mechanoreceptors involvedwith hearing and equilibrium detect settlingparticles or moving fluid

    Hearing and the perception of body equilibrium

    Are related in most animals

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    Sensing Gravity and Sound in Invertebrates

    Most invertebrates have sensory organs calledstatocysts

    That contain mechanoreceptors and function intheir sense of equilibrium

    Figure 49.6

    Ciliatedreceptor cells

    CiliaStatolith

    Sensory nerve fibers

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    Many arthropods sense sounds with body hairsthat vibrate

    Or with localized ears consisting of atympanic membrane and receptor cells

    Figure 49.7

    1 mm

    Tympanicmembrane

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    Hearing and Equilibrium in Mammals

    In most terrestrial vertebrates

    The sensory organs for hearing andequilibrium are closely associated in the ear

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    Exploring the structure of the human ear

    Figure 49.8

    Pinna

    Auditorycanal

    Eustachiantube

    Tympanicmembrane

    Stapes

    Incus

    Malleus

    Skullbones

    Semicircular canals

    Auditory nerve,to brain

    Cochlea

    Tympanicmembrane

    Ovalwindow

    Eustachiantube

    Roundwindow

    Vestibular canal

    Tympanic canal

    Auditory nerve

    BoneCochlear duct

    Hair cells Tectorialmembrane

    Basilar membrane

    To auditorynerve

    Axons of sensory neurons

    1 Overview of ear structure 2 The middle ear and inner ear

    4 The organ of Corti 3 The cochleaOrgan of Corti

    Outer ear Middle

    ear Inner ear

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    Hearing

    Vibrating objects create percussion waves inthe air

    That cause the tympanic membrane to vibrate

    The three bones of the middle ear

    Transmit the vibrations to the oval window onthe cochlea

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    The cochlea can distinguish pitch

    Because the basilar membrane is not uniformalong its length

    Cochlea(uncoiled)

    Basilar membrane

    Apex(wide and flexible)

    Base(narrow and stiff)

    500 Hz(low pitch)1 kHz

    2 kHz

    4 kHz

    8 kHz

    16 kHz(high pitch)

    Frequency producing maximumvibration

    Figure 49.10

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    Each region of the basilar membrane vibratesmost vigorously

    At a particular frequency and leads toexcitation of a specific auditory area of thecerebral cortex

    E ilib i

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    Equilibrium

    Several of the organs of the inner ear

    Detect body position and balance

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    The utricle, saccule, and semicircular canals inthe inner ear

    Function in balance and equilibrium

    Figure 49.11

    The semicircular canals, arranged in threespatial planes, detect angular movementsof the head.

    Body movement

    Nervefibers

    Each canal has at its base aswelling called an ampulla,containing a cluster of hair cells.

    When the head changes its rateof rotation, inertia preventsendolymph in the semicircular canals from moving with the head,so the endolymph presses against

    the cupula, bending the hairs.

    The utricle and saccule tell the brain whichway is up and inform it of the bodysposition or linear acceleration.

    The hairs of the hair cellsproject into a gelatinous capcalled the cupula.

    Bending of the hairs increases thefrequency of action potentials insensory neurons in directproportion to the amount of

    rotational acceleration.

    Vestibule

    Utricle

    Saccule

    Vestibular nerve

    Flowof endolymph

    Flowof endolymph

    CupulaHairs

    Hair cell

    H i g d E ilib i i Oth V t b t

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    Hearing and Equilibrium in Other Vertebrates

    Like other vertebrates, fishes and amphibians

    Also have inner ears located near the brain

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    Most fishes and aquatic amphibians

    Also have a lateral line system along bothsides of their body

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    The lateral line system containsmechanoreceptors

    With hair cells that respond to water movement

    Figure 49.12 Nerve fiber

    Supporting cell

    Cupula

    Sensoryhairs

    Hair cell

    Segmental muscles of body wall Lateral nerve

    Scale EpidermisLateral line canal

    Neuromast

    Opening of lateralline canal

    Lateralline

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    Concept 49.3: The senses of taste and smellare closely related in most animals

    The perceptions of gustation (taste) andolfaction (smell)

    Are both dependent on chemoreceptors thatdetect specific chemicals in the environment

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    The taste receptors of insects are locatedwithin sensory hairs called sensilla

    Which are located on the feet and inmouthparts

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    Figure 49.13

    EXPERIMENT Insects taste using gustatory sensilla (hairs) on their feet andmouthparts. Each sensillum contains four chemoreceptors with dendrites thatextend to a pore at the tip of the sensillum. To study the sensitivity of eachchemoreceptor, researchers immobilized a blowfly ( Phormia regina ) by attachingit to a rod with wax. They then inserted the tip of a microelectrode into onesensillum to record action potentials in the chemoreceptors, while they used apipette to touch the pore with various test substances.

    N u m

    b e r o

    f a c t

    i o n p o

    t e n

    t i a l s

    i n f i r s t s e c o n d

    o f r e s p o n s e

    CONCLUSION Any natural food probably stimulates multiple chemoreceptors. Byintegrating sensations, the insects brain can apparently distinguish a very largenumber of tastes.

    To brain

    Chemo-receptors

    Pore at tip

    Pipette containingtest substance

    To voltagerecorder

    Sensillum

    Microelectrode

    50

    30

    10

    00.5 M NaCl

    Meat 0.5 M Sucrose

    Honey

    Stimulus

    Chemoreceptors

    RESULTS Each chemoreceptor is especially sensitive to a particular class of substance, but this specificity is relative; each cell can respond tosome extent to a broad range of different chemical stimuli.

    Taste in Humans

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    Taste in Humans

    The receptor cells for taste in humans

    Are modified epithelial cells organized intotaste buds

    Five taste perceptions involve several signaltransduction mechanisms

    Sweet, sour, salty, bitter, and umami (elicitedby glutamate)

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    Taste pore Sugar molecule

    Sensoryreceptor cells

    Sensoryneuron

    Taste bud

    Tongue

    G protein Adenylyl cyclase

    Ca 2+

    ATP

    cAMP

    Proteinkinase A

    Sugar

    Sugar receptor

    SENSORYRECEPTORCELL Synaptic

    vesicle

    K+

    Neurotransmitter

    Sensory neuron

    Transduction in taste receptors

    Occurs by several mechanisms

    Figure 49.14

    4 The decrease in the membranes permeability toK+ depolarizes the membrane.

    5 Depolarization opens voltage-gated calcium ion(Ca 2+) channels, and Ca 2+ diffuses into the receptor cell.

    6 The increased Ca 2+ concentration causessynaptic vesicles to release neurotransmitter.

    3 Activated protein kinase A closes K + channels inthe membrane.

    2 Binding initiates a signal transduction pathwayinvolving cyclic AMP and protein kinase A.

    1 A sugar molecule binds

    to a receptor protein onthe sensory receptor cell.

    Smell in Humans

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    Smell in Humans

    Olfactory receptor cells

    Are neurons that line the upper portion of thenasal cavity

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    When odorant molecules bind to specificreceptors

    A signal transduction pathway is triggered,sending action potentials to the brain

    Brain

    Nasal cavity

    Odorant

    Odorantreceptors

    Plasmamembrane

    Odorant

    Cilia

    Chemoreceptor

    Epithelial cell

    Bone

    Olfactory bulb

    Action potentials

    MucusFigure 49.15

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    Concept 49.4: Similar mechanisms underlievision throughout the animal kingdom

    Many types of light detectors

    Have evolved in the animal kingdom and maybe homologous

    Vision in Invertebrates

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    Vision in Invertebrates

    Most invertebrates

    Have some sort of light-detecting organ

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    Light

    Light shining fromthe front is detected

    Photoreceptor

    Visual pigment

    Ocellus

    Nerve tobrain

    Screeningpigment

    Light shining frombehind is blockedby the screening pigment

    One of the simplest is the eye cup of planarians

    Which provides information about lightintensity and direction but does not formimages

    Figure 49.16

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    Two major types of image-forming eyes haveevolved in invertebrates

    The compound eye and the single-lens eye

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    Compound eyes are found in insects andcrustaceans

    And consist of up to several thousand lightdetectors called ommatidia

    Figure 49.17ab

    Cornea

    Crystallinecone

    Rhabdom

    Photoreceptor Axons

    Ommatidium

    Lens

    2 m m

    (a) The faceted eyes on thehead of a fly,

    photographed witha stereomicroscope.

    (b) The cornea and crystalline cone of each ommatidium function asa lens that focuses light on therhabdom, a stack of pigmented

    plates inside a circle of photoreceptors. The rhabdomtraps light and guides it tophotoreceptors. The imageformed by a compound eye is amosaic of dots produced by differentintensities of light entering themany ommatidia from different angles.

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    Structure of the Eye

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    Structure of the Eye

    The main parts of the vertebrate eye are

    The sclera, which includes the cornea

    The choroid, a pigmented layer

    The conjunctiva, that covers the outer surfaceof the sclera

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    The iris, which regulates the pupil

    The retina, which contains photoreceptors The lens, which focuses light on the retina

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    The structure of the vertebrate eye

    Figure 49.18

    Ciliary body

    Iris

    Suspensoryligament

    Cornea

    Pupil

    Aqueoushumor

    Lens

    Vitreous humor

    Optic disk(blind spot)

    Central artery andvein of the retina

    Opticnerve

    Fovea (center of visual field)

    Retina

    ChoroidSclera

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    The human retina contains two types of photoreceptors

    Rods are sensitive to light but do notdistinguish colors

    Cones distinguish colors but are not assensitive

    Sensory Transduction in the Eye

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    y y

    Each rod or cone in the vertebrate retina

    Contains visual pigments that consist of a light-absorbing molecule called retinal bonded to aprotein called opsin

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    Rods contain the pigment rhodopsin

    Which changes shape when it absorbs light

    Figure 49.20a, b

    Rod

    Outer segment

    Cell body

    Synapticterminal

    Disks

    Insideof disk

    (a) Rods contain the visual pigment rhodopsin, which is embedded ina stack of membranous disks in the rods outer segment.Rhodopsin consists of the light-absorbing molecule retinalbonded to opsin, a protein. Opsin has seven helices that spanthe disk membrane.

    (b) Retinal exists as two isomers. Absorption of light convertsthe cis isomer to the trans isomer, whichcauses opsin to change its conformation (shape).After a few minutes, retinal detaches from opsin.In the dark, enzymes convert retinal back to its cisform, which recombines with opsin to form rhodopsin.

    Retinal

    OpsinRhodopsin

    Cytosol

    HC

    CH2C

    CH2C C

    HCH 3

    CH 3H

    CC

    CH 3 H CH 3

    CC

    CC

    CC

    C

    H

    H

    H

    H

    OH

    H3C

    HC

    CH2C

    CH

    2C C

    HCH 3

    CH 3H

    CC

    CH 3 H CH 3

    C C C C

    HH

    CH 3

    H

    C C CHO

    CH 3

    trans isomer

    cis isomer

    EnzymesLight

    Processing Visual Information

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    g

    The processing of visual information

    Begins in the retina itself

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    In the dark, both rods and cones

    Release the neurotransmitter glutamate intothe synapses with neurons called bipolar cells,which are either hyperpolarized or depolarized

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    In the light, rods and cones hyperpolarize

    Shutting off their release of glutamate

    The bipolar cells

    Are then either depolarized or hyperpolarized

    Figure 49.22

    Dark Responses

    Rhodopsin inactive

    Na + channels open

    Rod depolarized

    Glutamatereleased

    Bipolar cell either depolarized or hyperpolarized,depending onglutamate receptors

    Light Responses

    Rhodopsin active

    Na + channels closed

    Rod hyperpolarized

    No glutamatereleased

    Bipolar cell either hyperpolarized or depolarized,depending onglutamate receptors

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    Three other types of neurons contribute toinformation processing in the retina

    Ganglion cells, horizontal cells, and amacrinecells

    Figure 49.23Opticnervefibers

    Ganglioncell

    Bipolar cell

    Horizontalcell

    Amacrinecell

    Pigmentedepithelium

    NeuronsCone Rod

    Photoreceptors

    Retina

    Retina

    Optic nerve

    Tobrain

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    Signals from rods and cones

    Travel from bipolar cells to ganglion cells

    The axons of ganglion cells are part of the opticnerve

    That transmit information to the brain

    Figure 49.24

    Leftvisualfield

    Rightvisualfield

    Lefteye

    Righteye

    Optic nerve

    Optic chiasm

    Lateralgeniculatenucleus

    Primaryvisual cortex

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    Most ganglion cell axons lead to the lateralgeniculate nuclei of the thalamus

    Which relays information to the primary visualcortex

    Several integrating centers in the cerebralcortex

    Are active in creating visual perceptions

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    Concept 49.5: Animal skeletons function insupport, protection, and movement

    The various types of animal movements

    All result from muscles working against some

    type of skeleton

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    Hydrostatic Skeletons

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    A hydrostatic skeleton

    Consists of fluid held under pressure in aclosed body compartment

    This is the main type of skeleton

    In most cnidarians, flatworms, nematodes, andannelids

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    Annelids use their hydrostatic skeleton for peristalsis

    A type of movement on land produced byrhythmic waves of muscle contractions

    Figure 49.25ac

    (a) Body segments at the head and just in frontof the rear are short and thick (longitudinalmuscles contracted; circular musclesrelaxed) and anchored to the ground bybristles. The other segments are thin andelongated (circular muscles contracted;longitudinal muscles relaxed.)

    (b) The head has moved forward because

    circular muscles in the head segments havecontracted. Segments behind the head andat the rear are now thick and anchored, thuspreventing the worm from slipping backward.

    (c) The head segments are thick again andanchored in their new positions. The rear segments have released their hold on theground and have been pulled forward.

    Longitudinalmuscle relaxed(extended)

    Circular musclecontracted

    Circular musclerelaxed

    Longitudinalmusclecontracted

    HeadBristles

    Exoskeletons

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    An exoskeleton is a hard encasement

    Deposited on the surface of an animal

    Exoskeletons

    Are found in most molluscs and arthropods

    Endoskeletons

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    An endoskeleton consists of hard supportingelements

    Such as bones, buried within the soft tissue of an animal

    Endoskeletons Are found in sponges, echinoderms, and

    chordates

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    The mammalian skeleton is built from morethan 200 bones

    Some fused together and others connected at joints by ligaments that allow freedom of movement

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    The human skeleton

    Figure 49.26

    1 Ball-and-socket joints, where the humerus contactsthe shoulder girdle and where the femur contacts thepelvic girdle, enable us to rotate our arms andlegs and move them in several planes.

    2 Hinge joints, such as between the humerus andthe head of the ulna, restrict movement to a singleplane.

    3 Pivot joints allow us to rotate our forearm at theelbow and to move our head from side to side.

    keyAxial skeletonAppendicular skeleton

    Skull

    Shoulder girdle

    Clavicle

    Scapula

    Sternum

    RibHumerus

    Vertebra

    Radius

    UlnaPelvicgirdle

    Carpals

    Phalanges

    Metacarpals

    Femur Patella

    Tibia

    Fibula

    TarsalsMetatarsalsPhalanges

    1

    Examplesof joints

    2

    3

    Head of humerus

    Scapula

    Humerus

    Ulna

    UlnaRadius

    Physical Support on Land

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    In addition to the skeleton

    Muscles and tendons help support large landvertebrates

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    Skeletal muscles are attached to the skeletonin antagonistic pairs

    With each member of the pair working againsteach other

    Figure 49.27

    Human Grasshopper

    Bicepscontracts

    Tricepsrelaxes

    Forearmflexes

    Bicepsrelaxes

    Tricepscontracts

    Forearmextends

    Extensor musclerelaxes

    Flexor musclecontracts

    Tibiaflexes

    Extensor musclecontracts

    Flexor musclerelaxes

    Tibiaextends

    Vertebrate Skeletal Muscle

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    Vertebrate skeletal muscle

    Is characterized by a hierarchy of smaller andsmaller units

    Figure 49.28

    Muscle

    Bundle of muscle fibers

    Single muscle fiber (cell)

    Plasma membrane

    Myofibril

    Lightband Dark band

    Z line

    Sarcomere

    TEM 0.5 mI band A band I band

    M line

    Thickfilaments(myosin)

    Thinfilaments(actin)

    H zoneSarcomere

    Z lineZ line

    Nuclei

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    A skeletal muscle consists of a bundle of longfibers

    Running parallel to the length of the muscle

    A muscle fiber

    Is itself a bundle of smaller myofibrils arrangedlongitudinally

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    The myofibrils are composed to two kinds of myofilaments

    Thin filaments, consisting of two strands of actin and one strand of regulatory protein

    Thick filaments, staggered arrays of myosinmolecules

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    Skeletal muscle is also called striated muscle

    Because the regular arrangement of themyofilaments creates a pattern of light anddark bands

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    Each repeating unit is a sarcomere

    Bordered by Z lines

    The areas that contain the myofilments

    Are the I band, A band, and H zone

    The Sliding-Filament Model of Muscle Contraction

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    According to the sliding-filament model of muscle contraction

    The filaments slide past each other longitudinally, producing more overlapbetween the thin and thick filaments

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    As a result of this sliding

    The I band and the H zone shrink

    Figure 49.29ac

    (a) Relaxed muscle fiber. In a relaxed muscle fiber, the I bands

    and H zone are relatively wide.

    (b) Contracting muscle fiber. During contraction, the thick andthin filaments slide past each other, reducing the width of theI bands and H zone and shortening the sarcomere.

    (c) Fully contracted muscle fiber. In a fully contracted musclefiber, the sarcomere is shorter still. The thin filaments overlap,eliminating the H zone. The I bands disappear as the ends of the thick filaments contact the Z lines.

    0.5 m

    Z HA

    Sarcomere

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    The sliding of filaments is based on

    The interaction between the actin and myosinmolecules of the thick and thin filaments

    The head of a myosin molecule binds to an

    actin filament Forming a cross-bridge and pulling the thin

    filament toward the center of the sarcomere

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    Myosin-actin interactions underlying musclefiber contraction

    Figure 49.30

    Thick filament

    Thin filaments

    Thin filament

    ATPATP

    ADPADP

    ADP

    P i P i

    P i

    Cross-bridge

    Myosin head (low-energy configuration)

    Myosin head (high-energy configuration)

    +

    Myosin head (low-

    energy configuration)

    Thin filament movestoward center of sarcomere.

    Thickfilament

    ActinCross-bridgebinding site

    1 Starting here, the myosin head isbound to ATP and is in its low-energy confinguration.

    2 The myosin head hydrolyzesATP to ADP and inorganicphosphate ( I ) and is in itshigh-energy configuration.

    P

    1 The myosin head binds toactin, forming a cross-bridge.

    3

    4 Releasing ADP and ( i), myosinrelaxes to its low-energy configuration,sliding the thin filament.

    P

    5 Binding of a new mole-cule of ATP releases themyosin head from actin,

    and a new cycle begins.

    The Role of Calcium and Regulatory Proteins

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    A skeletal muscle fiber contracts

    Only when stimulated by a motor neuron

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    When a muscle is at rest

    The myosin-binding sites on the thin filamentare blocked by the regulatory proteintropomyosin

    Figure 49.31a

    ActinTropomyosin Ca 2+-binding sites

    Troponin complex

    (a) Myosin-binding sites blocked

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    For a muscle fiber to contract

    The myosin-binding sites must be uncovered This occurs when calcium ions (Ca 2+)

    Bind to another set of regulatory proteins, thetroponin complex

    Figure 49.31b

    Ca 2+

    Myosin-binding site

    (b) Myosin-binding sites exposed

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    The stimulus leading to the contraction of askeletal muscle fiber

    Is an action potential in a motor neuron thatmakes a synapse with the muscle fiber

    Figure 49.32

    Motor neuron axon

    Mitochondrion

    Synapticterminal

    T tubule

    Sarcoplasmicreticulum

    Myofibril

    Plasma membraneof muscle fiber

    Sarcomere

    Ca 2+ releasedfrom sarcoplasmicreticulum

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    The synaptic terminal of the motor neuron

    Releases the neurotransmitter acetylcholine,depolarizing the muscle and causing it toproduce an action potential

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    ACh

    Synapticterminalof motor

    neuronSynaptic cleft T TUBULE

    PLASMA MEMBRANE

    SR

    ADP

    CYTOSOL

    Ca 2+

    Ca 2+

    P 2

    Cytosolic Ca 2+ isremoved by activetransport intoSR after actionpotential ends.

    6

    Review of contraction in a skeletal muscle fiber

    Figure 49.33

    Acetylcholine (ACh) released by synaptic terminal diffuses across synapticcleft and binds to receptor proteins on muscle fibers plasma membrane,triggering an action potential in muscle fiber.

    1

    Action potential is propa-gated along plasmamembrane and downT tubules.

    2

    Action potentialtriggers Ca 2+release from sarco-plasmic reticulum(SR).

    3

    Myosin cross-bridges alternately attachto actin and detach, pulling actinfilaments toward center of sarcomere;ATP powers sliding of filaments.

    5

    Calcium ions bind to troponin;troponin changes shape,removing blocking actionof tropomyosin; myosin-bindingsites exposed.

    4

    Tropomyosin blockage of myosin-binding sites is restored; contractionends, and muscle fiber relaxes.

    7

    Neural Control of Muscle Tension

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    Contraction of a whole muscle is graded

    Which means that we can voluntarily alter theextent and strength of its contraction

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    There are two basic mechanisms by which thenervous system produces graded contractionsof whole muscles

    By varying the number of fibers that contract

    By varying the rate at which muscle fibers arestimulated

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    In a vertebrate skeletal muscle

    Each branched muscle fiber is innervated byonly one motor neuron

    Each motor neuron

    May synapse with multiple muscle fibers

    Figure 49.34

    Spinal cord

    Nerve

    Motor neuroncell body

    Motor unit 1

    Motor unit 2

    Motor neuronaxon

    Muscle

    Tendon

    Synaptic terminals

    Muscle fibers

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    A motor unit

    Consists of a single motor neuron and all themuscle fibers it controls

    Recruitment of multiple motor neurons

    Results in stronger contractions

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    A twitch

    Results from a single action potential in amotor neuron

    More rapidly delivered action potentials

    Produce a graded contraction by summation

    Figure 49.35

    Actionpotential Pair of

    actionpotentials

    Series of actionpotentials at

    high frequency

    Time

    T e n s i o n

    Singletwitch

    Summation of two twitches

    Tetanus

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    Tetanus is a state of smooth and sustainedcontraction

    Produced when motor neurons deliver a volleyof action potentials

    Types of Muscle Fibers

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    Skeletal muscle fibers are classified as slowoxidative, fast oxidative, and fast glycolytic

    Based on their contraction speed and major pathway for producing ATP

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    Types of skeletal muscles

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    C 49 7 L i i

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    Concept 49.7: Locomotion requires energy toovercome friction and gravity

    Movement is a hallmark of all animals

    And usually necessary for finding food or

    evading predators

    Locomotion

    Is active travel from place to place

    Swimming

    O i f i i

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    Overcoming friction

    Is a major problem for swimmers Overcoming gravity is less of a problem for

    swimmers

    Than for animals that move on land or fly

    Locomotion on Land

    W lki i h i li l d

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    Walking, running, hopping, or crawling on land

    Requires an animal to support itself and moveagainst gravity

    Di d i f li l d

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    Diverse adaptations for traveling on land

    Have evolved in various vertebrates

    Figure 49.36

    Flying

    Fli ht i th t i d l h lift

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    Flight requires that wings develop enough lift

    To overcome the downward force of gravity

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    A i l th t i li d f i i g

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    Animals that are specialized for swimming

    Expend less energy per meter traveled thanequivalently sized animals specialized for flying or running