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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 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Both bats and moths 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 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Concept: Sensory receptors transduce stimulus energy and transmit signals to the central nervous system • Sensations and perceptions begin with sensory reception, the detection of stimuli by sensory receptors • Sensations are action potentials that reach the brain via sensory neurons • Exteroreceptors • Once the brain is aware of sensations it interprets them, giving the perception of stimuli. – Detect stimuli coming from the outside of the body • Interoreceptors – Detect internal stimuli Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 1 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 • Sensory receptors perform four functions in this process • 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 • Amplification is the strengthening of stimulus energy by cells in sensory pathways • 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 • Integration is the evaluation and coordination of stimuli to produce a response. It occurs at all levels of the nervous system. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Hair cell found in vertebrates Fluid moving in one direction No fluid movement “Hairs” of hair cell Neurotransmitter at synapse Types of Sensory Receptors More neurotransmitter Less neurotransmitter –70 Action potentials 0 Membrane potential (mV) Membrane potential (mV) –50 –70 0 hair cells have specialized cilia or microvilli (“hairs”) that bend when sur-rounding fluid moves. Each hair cell releases an excitatory neurotransmitter at a synapse – Chemoreceptors –70 – Electromagnetic receptors 0 –70 –70 0 1 2 3 4 5 6 7 Time (sec) (b) Vertebrate –70 0 1 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 • Based on the energy they transduce, sensory receptors fall into five categories – Mechanoreceptors: –50 Receptor potential Membrane potential (mV) –50 Axon Fluid moving in other direction 0 1 2 3 4 5 6 7 Time (sec) 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 – Thermoreceptors – Pain receptors Figure 49.2b Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 2 Mechanoreceptors Mechanoreceptors • Mechanoreceptors sense physical deformation • The mammalian sense of touch relies on mechanoreceptors that are the dendrites of sensory neurons – Caused by stimuli such as pressure, stretch, motion, and sound Cold Light touch Pain Hair Heat Epidermis Dermis Figure 49.3 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Nerve Connective tissueHair movementStrong pressure Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Chemoreceptors • Chemoreceptors include – General receptors that transmit information about the total solute concentration of a solution • Two of the most sensitive and specific chemoreceptors known are present in the antennae of the male silkworm moth. The males use their antennae to detect two components in pheromones released by females. Figure 49.4 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 0.1 mm – Specific receptors that respond to individual kinds of molecules Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 3 Electromagnetic Receptors • Electromagnetic receptors detect various forms of electromagnetic energy such as visible light, heat, electricity, and magnetism • Some snakes have very sensitive infrared receptors that detect body heat of prey against a colder background (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 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Thermoreceptors • Many mammals and birds appear to use the Earth’s magnetic field lines to orient themselves as they migrate • Thermoreceptors, which respond to heat or cold help regulate body temperature by signaling both surface and body core temperature 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. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 4 Pain Receptors Hearing, balance and the ear. • In humans, pain receptors, also called nociceptors are a class of naked dendrites in the epidermis • The mechanoreceptors involved with hearing and equilibrium detect settling particles or moving fluid • They respond to excess heat, pressure, or specific classes of chemicals released from damaged or inflamed tissues • Hearing and the perception of body equilibrium are related in most animals Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Hearing and Equilibrium in Mammals Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Structure of the human ear • In most terrestrial vertebrates the sensory organs for hearing and equilibrium are closely associated in the ear. 1 Overview of ear structure 2 The middle ear and inner ear Incus Semicircular Skull bones canals for balance Stapes Middle ear Inner ear Malleus Outer ear Auditory nerve, to brain • The ear is divided into three areas: Pinna – The outer ear which contains the auditory canal. Tympanic membrane – The middle ear which includes the tympanic membrane and the three ear bones that transmit vibrations to the inner ear Hair cells Cochlea Eustachian tube Auditory canal Tectorial membrane Tympanic membrane Oval window Cochlear duct – The inner ear which contains the cochlea where vibrations are converted to nerve impulses and sent to the brain as well as the organs of balance: utricle, saccule and semicircular canals. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Round window Eustachian tube Bone Vestibular canal Basilar membrane Figure 49.8 Axons of sensory neurons Auditory nerve To auditory nerve 4 The organ of Corti Tympanic canal 3 The cochlea Organ of Corti Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 5 Hearing Hearing • Vibrating objects create percussion waves in the air that cause the tympanic membrane to vibrate • These vibrations create pressure waves in the fluid in the cochlea that travel through the vestibular canal and ultimately strike the round window • The three bones of the middle ear transmit the vibrations to the oval window on the cochlea Cochlea Stapes Oval window Axons of sensory neurons Vestibular Perilymph canal Apex Base Figure 49.9 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Round window Tympanic Basilar canal membrane Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Hearing Hearing • The pressure waves in the vestibular canal cause the basilar membrane in the organ of Corti within the cochlea to vibrate up and down causing its hair cells to bend • The cochlea can distinguish pitch because the basilar membrane is not uniform along its length. 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. Cochlea (uncoiled) • The bending of the hair cells depolarizes their membranes sending action potentials that travel via the auditory nerve to the brain where they are interpreted as sounds. Apex (wide and flexible) Basilar membrane 1 kHz 500 Hz (low pitch) 2 kHz 4 kHz 8 kHz 16 kHz (high pitch) Figure 49.10 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Frequency producing maximum vibration Base (narrow and stiff) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 6 Equilibrium • Several of the organs of the inner ear detect body position and balance The semicircular canals, arranged in three spatial planes, detect angular movements of the head. • The utricle and saccule tell the brain which way is up. • Semicircular canals in the inner ear detect angular movements of the head and function in balance and equilibrium Each canal has at its base a swelling called an ampulla, containing a cluster of hair cells. Flow of endolymph 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 Vestibular nerve Cupula Hairs Hair cell Nerve fibers Vestibule Utricle Body movement Saccule The utricle and saccule tell the brain which way is up and inform it of the body’s position or linear acceleration. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings The hairs of the hair cells project into a gelatinous cap called the cupula. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Bending of the hairs increases the frequency of action potentials in sensory neurons in direct proportion to the amount of rotational acceleration. Odor detection in humans Odor detection in humans • Olfactory receptor cells • When odorant molecules bind to specific receptors a signal transduction pathway is triggered, sending action potentials to the brain – Are neurons that line the upper portion of the nasal cavity Brain Action potentials Odorant Olfactory bulb Nasal cavity Bone Epithelial cell Odorant receptors Chemoreceptor Plasma membrane Figure 49.15 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Odorant Cilia Mucus Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 7 Vision Structure of the Vertebrate Eye • Many types of light detectors have evolved in the animal kingdom ranging from simple eye cups of planarians (flatworms) to the compound eyes of insects and crustaceans to the camera-like eyes of vertebrates and cephalopods (octopuses and relatives). • The main parts of the vertebrate eye are – The sclera, a layer of tough connective tissue, which includes the transparent cornea which allows light into the eye and acts as a fixed lens. – The choroid, a pigmented layer inside the sclera. – The conjunctiva, that covers the outer surface of the sclera (except the cornea) and keeps it moist. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Structure of the Vertebrate Eye Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings The structure of the vertebrate eye – The iris, which regulates the size of the pupil the hole in the middle of the iris that lets light in. Sclera Choroid Retina Ciliary body Fovea (center of visual field) Suspensory ligament – The retina, which contains photoreceptors that detect light and convert it to nervous impulses. Cornea Iris – The lens, which focuses light on the retina Optic nerve Pupil Aqueous humor Lens Vitreous humor Central artery and vein of the retina Figure 49.18 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Optic disk (blind spot) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 8 Humans and other mammals focus light by changing the shape of the lens Ciliary muscles contract, pulling border of choroid toward lens Suspensory ligaments relax Choroid Front view of lens and ciliary muscle Lens (rounder) Retina Ciliary muscle Lens becomes thicker and rounder, focusing on near objects (a) Near vision (accommodation) Ciliary muscles relax, and border of choroid moves away from lens Suspensory ligaments • 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 to light. Lens (flatter) Suspensory ligaments pull against lens Lens becomes flatter, focusing on distant objects Figure 49.19a–b (b) Distance vision Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Sensory Transduction in the Eye • Each rod or cone in the vertebrate retina contains visual pigments that consist of a lightabsorbing pigment called retinal bonded to a protein called opsin. • Opsins vary in structure from one type of photoreceptor to another • Rods contain the visual pigment rhodopsin, which changes shape when it absorbs light. • Cones contain one or other of three different photopsins and are either red, green or blue cones. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Rods are used under low light conditions and cones in bright light for color vision. • Absorption spectra of the different types of cones overlap and the differential stimulation of the three types of cones allows the brain to produce the perception of color. • For example, if both red and green cones are stimulated we may see orange or yellow depending on which cones are more strongly stimulated. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 9 Processing Visual Information • The processing of visual information begins in the retina itself Light PDE • When a cone or rod is struck it absorbs light and its retinal changes shape. • This change in shape triggers a signal transduction pathway that generates an action potential and nerve impulses that travel to the brain. CYTOSOL Inactive rhodopsin Transducin Na+ 2 Active rhodopsin in turn activates a G protein called transducin. 3 Transducin activates the enzyme phosphodiestera e(PDE). 0 Dark Light – 40 GMP 1 Light isomerizes retinal, which activates rhodopsin. Membrane potential (mV) 4 Activated PDE detaches cyclic guanosine monophosphate (cGMP) from Na+ channels in the plasma membrane by hydrolyzing cGMP to GMP. – 70 – Hyperpolarization Time Na+ 5 The Na+ channels close when cGMP detaches. The membrane’s permeability to Na+ decreases, and the rod hyperpolarizes. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • 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 Left Right brain visual visual field Optic nerve Plasma membrane cGMP Disk membrane Figure 49.21 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings EXTRACELLULAR FLUID INSIDE OF DISK Active rhodopsin Left eye field • Several integrating centers in the cerebral cortex are active in creating visual perceptions. • You should be aware that the “reality” you perceive is a a simulation generated by your brain that builds a mental model of the world. Right eye Optic chiasm Lateral geniculate nucleus Primary visual cortex Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 10 Animal Skeletons Functions of the skeleton • Animal skeletons function in support, protection, and movement • The three main functions of a skeleton are – Support, • The various types of animal movements all result from muscles working against some type of skeleton Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings – Protection, – and Movement Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Human skeleton • The mammalian skeleton is an endoskeleton of bones buried within the soft tissue. It is built from more than 200 bones • Some bones are fused together and others are connected at joints by ligaments that allow freedom of movement 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 Phalanges Metacarpals Femur Patella 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 Ulna 2 Hinge joints, such as between the humerus and the head of the ulna, restrict movement to a single plane. Tibia Fibula Ulna Tarsals Metatarsals Phalanges Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Radius 3 Pivot joints allow us to rotate our forearm at the elbow and to move our head from side to side. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 11 Human • Muscles move skeletal parts by contracting • The action of a muscle is always to contract. • Skeletal muscles are attached to the skeleton in antagonistic pairs with each member of the pair working against each other e.g. triceps and biceps in the upper arm. Grasshopper Extensor muscle relaxes Biceps contracts Tibia flexes Flexor muscle contracts Triceps relaxes Forearm flexes Extensor muscle contracts Biceps relaxes Tibia extends Forearm extends Triceps contracts Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Flexor muscle relaxes Figure 49.27 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Muscle • Vertebrate skeletal muscle is characterized by a hierarchy of smaller and smaller units. Bundle of muscle fibers • 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. Z line Sarcomere TEM I band • Myofibrils are composed of myofilaments. Figure 49.28 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Nuclei Single muscle fiber (cell) Plasma membrane Myofibril Light band Dark band Thick filaments (myosin) Thin filaments (actin) Z line A band M line H zone Sarcomere 0.5 µm I band Z line Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 12 • The myofibrils are composed of 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 • Skeletal muscle is also called striated muscle because the regular arrangement of the myofilaments creates a pattern of light and dark bands Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Each repeating unit is a sarcomere bordered by Z lines. The Z lines are aligned vertically and are attached to the thin filaments (made of actin). • The thin filaments project into the sarcomere. • The thick filaments are aligned in the center of the sarcomere and align with and partially overlap with the thin filaments. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Muscle • At rest the thick and thin fibers only partially overlap and this produces a pattern of bands in the sarcomere that are identified by letters. Bundle of muscle fibers • The A-band is the area in the middle of the sarcomere where the thick filaments are located and appears dark. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Z line Sarcomere • The I-band is the area at either end of the sarcomere where only thin filaments are found. This appears light. • The H-zone in the middle is where only thick filaments occur. It appears less dark than the rest of the A-band. Nuclei Single muscle fiber (cell) Plasma membrane Myofibril Light band Dark band TEM I band Figure 49.28 Thick filaments (myosin) Thin filaments (actin) Z line A band M line H zone Sarcomere 0.5 µm I band Z line Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 13 I band 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. (a) Relaxed muscle fiber. In a relaxed muscle fiber, the I bands and H zone are relatively wide. • As a result of this sliding the I band and the H zone shrink. 0.5 µm 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. (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. Figure 49.29a–c Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Myosin-actin interactions underlying muscle fiber contraction • 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 Thick filament 1 Starting here, the myosin head is bound to ATP and is in its low-energy confinguration. Thin filaments Thin filament 5 Binding of a new molecule of ATP releases the myosin head from actin, and a new cycle begins. ATP ATP Thin filament moves toward center of sarcomere. Thick filament Cross-bridge binding site Actin Pi ADP Pi 4 Releasing ADP and ( Pi), myosin relaxes to its low-energy configuration, sliding the thin filament. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings myosin head hydrolyzes ATP to ADP and inorganic phosphate ( PI ) and is in its high-energy configuration. ADP Myosin head (lowenergy configuration) ADP + P i 2 The Myosin head (lowenergy configuration) Cross-bridge Myosin head (highenergy configuration) 13 The myosin head binds to actin, forming a crossbridge. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 14 The Role of Calcium and Regulatory Proteins • A skeletal muscle fiber contracts – Only when stimulated by a motor neuron • When a muscle is at rest the myosin-binding sites on the thin filament are blocked by the regulatory protein tropomyosin Tropomyosin Actin Figure 49.31a Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Ca2+-binding sites Troponin complex (a) Myosin-binding sites blocked Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • For a muscle fiber to contract the myosinbinding sites must be uncovered • This occurs when calcium ions (Ca2+) bind to another set of regulatory proteins, the troponin complex • 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 Ca2+ T tubule Myosinbinding site Sarcoplasmic reticulum Figure 49.31b (b) Myosin-binding sites exposed Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 49.32 Myofibril Plasma membrane of muscle fiber Ca2+ released from sarcoplasmic reticulum Sarcomere Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 15 Neural Control of Muscle Tension • The synaptic terminal of the motor neuron releases the neurotransmitter acetylcholine, depolarizing the muscle and causing it to produce an action potential • Contraction of a whole muscle is graded – Which means that we can voluntarily alter the extent and strength of its contraction. • 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 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • 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 • 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 Motor Motor unit 1 unit 2 Synaptic terminals Spinal cord Nerve Motor neuron cell body Motor neuron axon Muscle Muscle fibers Figure 49.34 Tendon Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 16 • A twitch results from a single action potential in a motor neuron • More rapidly delivered action potentials produce a graded contraction by summation • Tetanus is a state of smooth and sustained contraction produced when motor neurons deliver a volley of action potentials. Tension Tetanus Summation of two twitches Single twitch Action potential Time Pair of action potentials Series of action potentials at high frequency Figure 49.35 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Cardiac muscle Smooth muscle • Cardiac muscle, found only in the heart • In smooth muscle, found mainly in the walls of hollow organs (e.g. the digestive tract) the contractions are relatively slow and may be initiated by the muscles themselves – Consists of striated cells that are electrically connected together. As a result an action potential in one part of the heart spreads to all cardiac cells and the heat contracts. – Cardiac muscle can generate action potentials without neural input because the heart has its own pacemaker cells that cause rhythmic depolarizarions. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • In addition, contractions may be caused by stimulation from neurons in the autonomic nervous system Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 17