Chapter 49

Sensory and Motor Mechanisms
PowerPoint Lectures for Biology, Seventh Edition
Neil Campbell and Jane Reece

Lectures by Chris Romero
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• Overview: Sensing and Acting • Bats use sonar to detect their prey • Moths, a common prey for bats
– Can detect the bat’s sonar and attempt to flee

Figure 49.1
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• Both of these organisms
– Have complex sensory systems that facilitate their survival

• The structures that make up these systems
– Have been transformed by evolution into diverse mechanisms that sense various stimuli and generate the appropriate physical movement

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• Concept 49.1: Sensory receptors transduce stimulus energy and transmit signals to the central 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 the body

• 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
Muscle Dendrites Membrane potential (mV) Weak muscle stretch –50 Receptor potential –70 Action potentials 0 –70 0 1 2 3 4 5 6 7 Time (sec) (a) Crayfish stretch receptors have dendrites embedded in abdominal muscles. When the abdomen bends, muscles and dendrites stretch, producing a receptor potential in the stretch receptor. The receptor potential triggers action potentials in the axon of the stretch 0 –70 01 2 3 4 5 67 Time (sec) receptor. A stronger stretch produces a larger receptor potential and higher requency of action potentials. –50 –70 Strong muscle stretch

Stretch receptor Axon

Figure 49.2a

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– A hair cell found in vertebrates
“Hairs” of hair cell Neurotransmitter at synapse Axon Membrane potential (mV) –50 Membrane potential (mV) –70 Action potentials 0 –70 01 2 3 4 5 6 7 Time (sec) (b) Vertebrate hair cells have specialized cilia or microvilli (“hairs”) that bend when surrounding fluid moves. Each hair cell releases an excitatory neurotransmitter at a synapse No fluid movement More neurotransmitter –50 Receptor potential Membrane potential (mV) –70 Fluid moving in one direction Less neurotransmitter –50 –70 Fluid moving in other direction

0 –70 0 1 2 3 4 5 6 7 Time (sec)

0 –70 01 2 3 4 5 6 7 Time (sec)

with a sensory neuron, which conducts action potentials to the CNS. Bending in one direction depolarizes the hair cell, causing it to release more neurotransmitter and increasing frequency

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

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 with sensory 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 sensory adaptation
– A decrease in responsiveness during continued 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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Mechanoreceptors • Mechanoreceptors sense physical deformation
– Caused by stimuli such as pressure, stretch, motion, and sound

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• The mammalian sense of touch
– Relies on mechanoreceptors that are the dendrites of sensory neurons
Cold Light touch Heat Pain Hair

Epidermis

Dermis

Figure 49.3

Nerve

Connective tissue

Hair movement

Strong pressure

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Chemoreceptors • Chemoreceptors include
– General receptors that transmit information about the total solute concentration of a solution – Specific receptors that respond to individual kinds 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
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0.1 mm

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 enough to 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 detected equally by the two receptors, indicating that the mouse is straight ahead. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings

• Many mammals appear to use the Earth’s magnetic field lines
– To orient themselves as they migrate

Figure 49.5b
(b) Some migrating animals, such as these beluga whales, apparently sense Earth’s magnetic field and use the information, along with other cues, for orientation.

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Thermoreceptors • Thermoreceptors, which respond to heat or cold
– Help regulate body temperature by signaling both surface and body core temperature

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Pain Receptors • In humans, pain receptors, also called nociceptors
– Are a class of naked dendrites in the epidermis – Respond to excess heat, pressure, or specific classes of chemicals released from damaged or inflamed tissues

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• Concept 49.2: The mechanoreceptors involved with hearing and equilibrium detect settling particles 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 called statocysts
– That contain mechanoreceptors and function in their sense of equilibrium

Ciliated receptor cells

Statolith

Cilia

Figure 49.6
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Sensory nerve fibers

• Many arthropods sense sounds with body hairs that vibrate
– Or with localized “ears” consisting of a tympanic membrane and receptor cells

Tympanic membrane

Figure 49.7
1 mm
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Hearing and Equilibrium in Mammals • In most terrestrial vertebrates
– The sensory organs for hearing and equilibrium are closely associated in the ear

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• Exploring the structure of the human ear
1 Overview of ear structure Incus Middle ear Inner ear Stapes Malleus Auditory nerve, to brain 2 The middle ear and inner ear Skull bones Semicircular canals Outer ear

Pinna

Tympanic membrane Eustachian tube Oval window Round window Cochlea

Auditory canal

Hair cells

Tectorial membrane

Tympanic membrane

Eustachian tube

Cochlear duct Bone Vestibular canal Auditory nerve Basilar membrane Axons of sensory neurons To auditory nerve Tympanic canal

Figure 49.8

4 The organ of Corti

3 The cochlea

Organ of Corti

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Hearing • Vibrating objects create percussion waves in the air
– That cause the tympanic membrane to vibrate

• The three bones of the middle ear
– Transmit the vibrations to the oval window on the cochlea

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• These vibrations create pressure waves in the fluid in the cochlea
– That travel through the vestibular canal and ultimately strike the round window

Cochlea Stapes Axons of sensory neurons Vestibular canal Perilymph

Oval window

Apex

Base

Figure 49.9

Round window

Tympanic Basilar canal membrane

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• The pressure waves in the vestibular canal
– Cause the basilar membrane to vibrate up and down causing its hair cells to bend

• The bending of the hair cells depolarizes their membranes
– Sending action potentials that travel via the auditory nerve to the brain

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• The cochlea can distinguish pitch
– Because the basilar membrane is not uniform along its length
Cochlea (uncoiled) Apex (wide and flexible) Basilar membrane

1 kHz 2 kHz 4 kHz 8 kHz 16 kHz (high pitch)

500 Hz (low pitch)

Frequency producing maximum vibration

Figure 49.10

Base (narrow and stiff)

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• Each region of the basilar membrane vibrates most vigorously
– At a particular frequency and leads to excitation of a specific auditory area of the cerebral cortex

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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 in the inner ear
– Function in balance and equilibrium
The semicircular canals, arranged in three spatial planes, detect angular movements of the head. Each canal has at its base a swelling called an ampulla, containing a cluster of hair cells. When the head changes its rate of rotation, inertia prevents endolymph in the semicircular canals from moving with the head, so the endolymph presses against the cupula, bending the hairs.

Flow of endolymph Flow of endolymph Vestibular nerve Cupula Hairs Hair cell Vestibule Utricle Saccule Nerve fibers Body movement

Figure 49.11

The utricle and saccule tell the brain which way is up and inform it of the body’s position or linear acceleration.

The hairs of the hair cells project into a gelatinous cap called the cupula.

Bending of the hairs increases the frequency of action potentials in sensory neurons in direct proportion to the amount of rotational acceleration.

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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 both sides of their body

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• The lateral line system contains mechanoreceptors
– With hair cells that respond to water movement
Lateral line Opening of lateral line canal

Lateral line canal Scale Epidermis Neuromast

Segmental muscles of body wall

Lateral nerve Cupula Sensory hairs

Supporting cell

Hair cell

Figure 49.12

Nerve fiber

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• Concept 49.3: The senses of taste and smell are closely related in most animals • The perceptions of gustation (taste) and olfaction (smell)
– Are both dependent on chemoreceptors that detect specific chemicals in the environment

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• The taste receptors of insects are located within sensory hairs called sensilla
– Which are located on the feet and in mouthparts

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EXPERIMENT Insects taste using gustatory sensilla (hairs) on their feet and mouthparts. Each sensillum contains four chemoreceptors with dendrites that extend to a pore at the tip of the sensillum. To study the sensitivity of each chemoreceptor, researchers immobilized a blowfly (Phormia regina) by attaching it to a rod with wax. They then inserted the tip of a microelectrode into one sensillum to record action potentials in the chemoreceptors, while they used a pipette to touch the pore with various test substances.

To brain

Chemoreceptors

Sensillum

Microelectrode To voltage recorder

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

Pore at tip Pipette containing test substance

Number of action potentials in first second of response

Chemoreceptors 50 30 10 0 0.5 M NaCl Meat 0.5 M Sucrose Stimulus Honey

CONCLUSION Any natural food probably stimulates multiple chemoreceptors. By integrating sensations, the insect’s brain can apparently distinguish a very large number of tastes.

Figure 49.13

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Taste in Humans • The receptor cells for taste in humans
– Are modified epithelial cells organized into taste buds

• Five taste perceptions involve several signal transduction mechanisms
– Sweet, sour, salty, bitter, and umami (elicited by glutamate)

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• Transduction in taste receptors
– Occurs by several mechanisms
Taste pore Taste bud Sugar molecule Sensory receptor cells

Tongue

Sensory neuron

1 Sugar Sugar receptor

A sugar molecule binds to a receptor protein on the sensory receptor cell. G protein Adenylyl cyclase 2 Binding initiates a signal transduction pathway involving cyclic AMP and protein kinase A.

ATP cAMP

Protein kinase A SENSORY K+ RECEPTOR CELL Synaptic vesicle —Ca2+

3 Activated protein kinase A closes K+ channels in the membrane. 4 The decrease in the membrane’s permeability to K+ depolarizes the membrane.

Neurotransmitter

5 Depolarization opens voltage-gated calcium ion (Ca2+) channels, and Ca2+ diffuses into the receptor cell. 6 The increased Ca2+ concentration causes synaptic vesicles to release neurotransmitter. Sensory neuron

Figure 49.14
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Smell in Humans • Olfactory receptor cells
– Are neurons that line the upper portion of the nasal cavity

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• When odorant molecules bind to specific receptors
– A signal transduction pathway is triggered, sending action potentials to the brain
Brain Action potentials Odorant Nasal cavity Bone Epithelial cell Odorant receptors Plasma membrane Olfactory bulb

Chemoreceptor Cilia

Figure 49.15

Odorant

Mucus

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• Concept 49.4: Similar mechanisms underlie vision throughout the animal kingdom • Many types of light detectors
– Have evolved in the animal kingdom and may be homologous

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Vision in Invertebrates • Most invertebrates
– Have some sort of light-detecting organ

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• One of the simplest is the eye cup of planarians
– Which provides information about light intensity and direction but does not form images
Light Light shining from the front is detected

Photoreceptor Visual pigment Ocellus

Nerve to brain Screening pigment Light shining from behind is blocked by the screening pigment

Figure 49.16
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• Two major types of image-forming eyes have evolved in invertebrates
– The compound eye and the single-lens eye

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• Compound eyes are found in insects and crustaceans
– And consist of up to several thousand light detectors called ommatidia
(a) The faceted eyes on the head of a fly, photographed with a stereomicroscope. 2 mm (b) The cornea and crystalline cone of each ommatidium function as a lens that focuses light on the rhabdom, a stack of pigmented plates inside a circle of photoreceptors. The rhabdom traps light and guides it to photoreceptors. The image formed by a compound eye is a mosaic of dots produced by different intensities of light entering the many ommatidia from different angles. Cornea

Crystalline cone

Lens

Rhabdom Photoreceptor Ommatidium

Axons

Figure 49.17a–b
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• Single-lens eyes
– Are found in some jellies, polychaetes, spiders, and many molluscs – Work on a camera-like principle

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The Vertebrate Visual System • The eyes of vertebrates are camera-like
– But they evolved independently and differ from the single-lens eyes of invertebrates

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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 surface of 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
Sclera Choroid Retina Ciliary body Fovea (center of visual field)

Suspensory ligament

Cornea Iris Pupil Optic nerve

Aqueous humor Lens Vitreous humor Central artery and vein of the retina

Figure 49.18
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Optic disk (blind spot)

• Humans and other mammals
– Focus light by changing the shape of the lens
Ciliary muscles contract, pulling border of choroid toward lens Choroid Suspensory ligaments relax Retina Front view of lens and ciliary muscle Lens (rounder)

Ciliary muscle Lens becomes thicker and rounder, focusing on near objects (a) Near vision (accommodation) Suspensory ligaments

Ciliary muscles relax, and border of choroid moves away from lens

Lens (flatter)

Suspensory ligaments pull against lens

Lens becomes flatter, focusing on distant objects

Figure 49.19a–b

(b) Distance vision

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• The human retina contains two types of photoreceptors
– Rods are sensitive to light but do not distinguish colors – Cones distinguish colors but are not as sensitive

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Sensory Transduction in the Eye • Each rod or cone in the vertebrate retina
– Contains visual pigments that consist of a lightabsorbing molecule called retinal bonded to a protein called opsin

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• Rods contain the pigment rhodopsin
– Which changes shape when it absorbs light
Rod Outer segment H H2C Disks H2C CH3 C CH3 C H C C C C C H C H3C H C C H C C H C H O H

CH3 H Cell body Inside of disk Light Synaptic terminal

CH3

cis isomer

Enzymes

H H2C H2C Cytosol Rhodopsin Retinal Opsin

H C

CH3 CH3 C H C C C C

H C C

H C C

H C C O H

C

CH3 H

CH3

CH3

CH3

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

Figure 49.20a, b

(a) Rods contain the visual pigment rhodopsin, which is embedded in a stack of membranous disks in the rod’s outer segment. Rhodopsin consists of the light-absorbing molecule retinal bonded to opsin, a protein. Opsin has seven α helices that span the disk membrane.

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Processing Visual Information • The processing of visual information
– Begins in the retina itself

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• Absorption of light by retinal
– Triggers a signal transduction pathway
Light Active rhodopsin PDE Membrane potential (mV) 0 Dark Light INSIDE OF DISK EXTRACELLULAR FLUID

CYTOSOL cGMP GMP

Plasma membrane

Inactive rhodopsin

Transducin

Disk membrane

– 40 Na+ – 70 – Hyperpolarization Time 5 The Na+ channels close when cGMP detaches. The membrane’s permeability to Na+ decreases, and the rod hyperpolarizes.

1 Light isomerizes retinal, which activates rhodopsin.

2 Active rhodopsin in turn activates a G protein called transducin.

3 Transducin activates the enzyme phosphodiesterae (PDE).

4 Activated PDE detaches cyclic guanosine monophosphate (cGMP) from Na+ channels in the plasma membrane by hydrolyzing cGMP to GMP.

Na+

Figure 49.21

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• In the dark, both rods and cones
– Release the neurotransmitter glutamate into the 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
Dark Responses Rhodopsin inactive Light Responses Rhodopsin active Na+ channels open Na+ channels closed

Rod depolarized

Rod hyperpolarized

Glutamate released Bipolar cell either depolarized or hyperpolarized, depending on glutamate receptors

No glutamate released Bipolar cell either hyperpolarized or depolarized, depending on glutamate receptors

Figure 49.22

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• Three other types of neurons contribute to information processing in the retina
– Ganglion cells, horizontal cells, and amacrine cells
Retina

Optic nerve

To brain

Retina Neurons Photoreceptors Cone Rod

Amacrine cell

Figure 49.23

Optic nerve fibers

Horizontal cell Pigmented epithelium

Ganglion cell

Bipolar cell

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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 optic nerve
– That transmit information to the brain
Left visual field Right visual field

Optic nerve Optic chiasm Lateral geniculate nucleus

Left eye

Right eye

Figure 49.24

Primary visual cortex

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• Most ganglion cell axons lead to the lateral geniculate nuclei of the thalamus
– Which relays information to the primary visual cortex

• Several integrating centers in the cerebral cortex
– Are active in creating visual perceptions

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• Concept 49.5: Animal skeletons function in support, protection, and movement • The various types of animal movements
– All result from muscles working against some type of skeleton

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Types of Skeletons • The three main functions of a skeleton are
– Support, protection, and movement

• The three main types of skeletons are
– Hydrostatic skeletons, exoskeletons, and endoskeletons

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Hydrostatic Skeletons • A hydrostatic skeleton
– Consists of fluid held under pressure in a closed body compartment

• This is the main type of skeleton
– In most cnidarians, flatworms, nematodes, and annelids

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• Annelids use their hydrostatic skeleton for peristalsis
– A type of movement on land produced by rhythmic waves of muscle contractions
(a) Body segments at the head and just in front of the rear are short and thick (longitudinal muscles contracted; circular muscles relaxed) and anchored to the ground by bristles. The other segments are thin and elongated (circular muscles contracted; longitudinal muscles relaxed.) Longitudinal muscle relaxed (extended) Circular muscle contracted Circular muscle relaxed Longitudinal muscle contracted

Bristles

Head

(b) The head has moved forward because circular muscles in the head segments have contracted. Segments behind the head and at the rear are now thick and anchored, thus preventing the worm from slipping backward.

Figure 49.25a–c

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

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Exoskeletons • An exoskeleton is a hard encasement
– Deposited on the surface of an animal

• Exoskeletons
– Are found in most molluscs and arthropods

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Endoskeletons • An endoskeleton consists of hard supporting elements
– 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 more than 200 bones
– Some fused together and others connected at joints by ligaments that allow freedom of movement

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• The human skeleton
key
Axial skeleton Appendicular skeleton Skull

Examples of joints

Head of humerus Scapula

1
Shoulder girdle Sternum Rib Humerus Vertebra Radius Ulna Pelvic girdle Carpals Clavicle Scapula

2 3

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

Phalanges Metacarpals Femur Patella

Ulna

2 Hinge joints, such as between the humerus and the head of the ulna, restrict movement to a single plane.

Tibia Fibula

Ulna

Figure 49.26

Tarsals Metatarsals Phalanges

Radius

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

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Physical Support on Land • In addition to the skeleton
– Muscles and tendons help support large land vertebrates

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• Concept 49.6: Muscles move skeletal parts by contracting • The action of a muscle
– Is always to contract

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• Skeletal muscles are attached to the skeleton in antagonistic pairs
– With each member of the pair working against each other
Human Grasshopper Biceps contracts Extensor muscle relaxes Tibia flexes Flexor muscle contracts

Triceps relaxes

Forearm flexes

Biceps relaxes

Extensor muscle contracts

Tibia extends

Forearm extends

Figure 49.27

Triceps contracts

Flexor muscle relaxes

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Vertebrate Skeletal Muscle • Vertebrate skeletal muscle
– Is characterized by a hierarchy of smaller and smaller units
Muscle Bundle of muscle fibers Nuclei

Single muscle fiber (cell) Plasma membrane Myofibril Light band Z line

Dark band

Sarcomere

TEM I band Thick filaments (myosin) A band M line

0.5 µm I band

Figure 49.28

Thin filaments (actin) Z line H zone Sarcomere Z line

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• A skeletal muscle consists of a bundle of long fibers
– Running parallel to the length of the muscle

• A muscle fiber
– Is itself a bundle of smaller myofibrils arranged longitudinally

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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 myosin molecules

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• Skeletal muscle is also called striated muscle
– Because the regular arrangement of the myofilaments creates a pattern of light and dark 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

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The Sliding-Filament Model of Muscle Contraction • According to the sliding-filament model of muscle contraction
– The filaments slide past each other longitudinally, producing more overlap between the thin and thick filaments

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• As a result of this sliding
– The I band and the H zone shrink
0.5 µm

(a) Relaxed muscle fiber. In a relaxed muscle fiber, the I bands and H zone are relatively wide.

Z

H A Sarcomere

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

Figure 49.29a–c

(c) Fully contracted muscle fiber. In a fully contracted muscle fiber, 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.

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• The sliding of filaments is based on
– The interaction between the actin and myosin molecules 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 muscle fiber contraction
Thick filament Thin filaments
1 Starting here, the myosin head is bound to ATP and is in its lowenergy confinguration.

5 Binding of a new molecule of ATP releases the myosin head from actin, and a new cycle begins.

Thin filament
ATP ATP

Myosin head (lowenergy configuration)

2 The myosin head hydrolyzes ATP to ADP and inorganic phosphate ( P I ) and is in its high-energy configuration.

Thin filament moves toward center of sarcomere. Myosin head (lowenergy configuration)

Thick filament

Actin
ADP Pi

Cross-bridge binding site
Myosin head (highenergy configuration)

ADP

+

Pi

ADP

Pi

Cross-bridge

Figure 49.30

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

1 The myosin head binds to 3 actin, forming a crossbridge.

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The Role of Calcium and Regulatory Proteins • 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 filament are blocked by the regulatory protein tropomyosin

Tropomyosin Actin

Ca2+-binding sites Troponin complex

Figure 49.31a

(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 (Ca2+)
– Bind to another set of regulatory proteins, the troponin complex
Ca2+ Myosinbinding site

Figure 49.31b

(b) Myosin-binding sites exposed

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• The stimulus leading to the contraction of a skeletal muscle fiber
– Is an action potential in a motor neuron that makes a synapse with the muscle fiber
Motor neuron axon Mitochondrion

Synaptic terminal

T tubule

Sarcoplasmic reticulum Myofibril

Ca2+ released from sarcoplasmic reticulum Sarcomere

Figure 49.32

Plasma membrane of muscle fiber

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• The synaptic terminal of the motor neuron
– Releases the neurotransmitter acetylcholine, depolarizing the muscle and causing it to produce an action potential

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• Action potentials travel to the interior of the muscle fiber
– Along infoldings of the plasma membrane called transverse (T) tubules

• The action potential along the T tubules
– Causes the sarcoplasmic reticulum to release Ca2+

• The Ca2+ binds to the troponin-tropomyosin complex on the thin filaments
– Exposing the myosin-binding sites and allowing the cross-bridge cycle to proceed
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• Review of contraction in a skeletal muscle fiber
Synaptic terminal of motor neuron 1 Acetylcholine (ACh) released by synaptic terminal diffuses across synaptic cleft and binds to receptor proteins on muscle fiber’s plasma membrane, triggering an action potential in muscle fiber. Synaptic cleft 2 Action potential is propagated along plasma membrane and down T tubules. T TUBULE PLASMA MEMBRANE

ACh

SR 3 Action potential triggers Ca2+ release from sarcoplasmic reticulum (SR).

Ca2+

7 Tropomyosin blockage of myosinbinding sites is restored; contraction ends, and muscle fiber relaxes. Ca2+ CYTOSOL
2+ 6 Cytosolic Ca is removed by active transport into SR after action potential ends.

4 Calcium ions bind to troponin; troponin changes shape, removing blocking action of tropomyosin; myosin-binding sites exposed.

ADP P2

Figure 49.33
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5 Myosin cross-bridges alternately attach to actin and detach, pulling actin filaments toward center of sarcomere; ATP powers sliding of filaments.

Neural Control of Muscle Tension • Contraction of a whole muscle is graded
– Which means that we can voluntarily alter the extent and strength of its contraction

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• There are two basic mechanisms by which the nervous system produces graded contractions of whole muscles
– By varying the number of fibers that contract – By varying the rate at which muscle fibers are stimulated

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• In a vertebrate skeletal muscle
– Each branched muscle fiber is innervated by only one motor neuron

• Each motor neuron
– May synapse with multiple muscle fibers
Spinal cord Motor unit 1 Motor unit 2 Synaptic terminals Nerve Motor neuron cell body Motor neuron axon

Muscle

Muscle fibers

Figure 49.34
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Tendon

• A motor unit
– Consists of a single motor neuron and all the muscle 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 a motor neuron

• More rapidly delivered action potentials
– Produce a graded contraction by summation
Tetanus Tension

Summation of two twitches Single twitch

Action potential

Time Pair of action potentials Series of action potentials at high frequency

Figure 49.35
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• Tetanus is a state of smooth and sustained contraction
– Produced when motor neurons deliver a volley of action potentials

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Types of Muscle Fibers • Skeletal muscle fibers are classified as slow oxidative, 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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Other Types of Muscle • Cardiac muscle, found only in the heart
– Consists of striated cells that are electrically connected by intercalated discs – Can generate action potentials without neural input

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• In smooth muscle, found mainly in the walls of hollow organs
– The contractions are relatively slow and may be initiated by the muscles themselves

• In addition, contractions may be caused by
– Stimulation from neurons in the autonomic nervous system

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• Concept 49.7: Locomotion requires energy to overcome 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

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Swimming • 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

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Locomotion on Land • Walking, running, hopping, or crawling on land
– Requires an animal to support itself and move against gravity

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• Diverse adaptations for traveling on land
– Have evolved in various vertebrates

Figure 49.36
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Flying • Flight requires that wings develop enough lift
– To overcome the downward force of gravity

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Comparing Costs of Locomotion •The energy cost of locomotion
–Depends on the mode of locomotion and the environment
Physiologists typically determine an animal’s rate of energy use during locomotion by measuring its oxygen consumption or carbon dioxide production while it swims in a water flume, runs on a treadmill, or flies in a wind tunnel. For example, the trained parakeet shown below is wearing a plastic face mask connected to a tube that collects the air the bird exhales as it flies. This graph compares the energy cost, in joules per kilogram of body mass per meter traveled, for animals specialized for running, flying, and swimming (1 J = 0.24 cal). Notice that both axes are plotted on logarithmic scales. For animals of a given body mass, swimming is the most energyCONCLUSION efficient and running the least energyefficient mode of locomotion. In any mode, a small animal expends more energy per kilogram of body mass than a large animal.

EXPERIMENT

RESULTS

CONCLUSION

Flying
Energy cost (J/Kg/m) 10
2

Running

10 1 10–1 10–3

Swimming
1 103 Body mass(g) 106

Figure 49.37
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• Animals that are specialized for swimming
– Expend less energy per meter traveled than equivalently sized animals specialized for flying or running

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