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 Chapter 8
Muscle Physiology

CHAPTER OUTLINE

Skeletal muscle contractions can play a part in homeostatic activities, such as heat generation, and in non-homeostatic
activities, such as movement of the body through space. Smooth muscle is found in the walls of hollow organs and
tubes. It functions to regulate movement of substances through the specific structures. Cardiac muscle is found only in
the walls of the heart and functions to move blood throughout the body.

STRUCTURE OF SKELETAL MUSCLE

Movement of specialized intracellular components allows skeletal muscle to generate tension and contract. Controlled
contractions of muscle allows: (1) purposeful movement of the body and parts of the body, (2) manipulation of external
objects, (3) propulsion of contents through hollow internal organs, and (4) emptying of certain organs to the external
environment.
Muscle is the largest group of tissues in the body, constituting approximately half the body’s weight with skeletal
muscle accounting for 40 percent of the body weight in men and 32 percent of the body weight in women.
Skeletal muscles are composed of a number of muscle cells (fibers). Skeletal muscle fibers are elongated cylinders
measuring from 10 to 100 m in diameter and up to 750,000 m in length. These cells have multiple nuclei, as they
formed from the fusion of myoblasts during embryonic development. The muscle fibers lie parallel to one another and
bundled together by connective tissue. Skeletal muscle fibers appear striated due to a highly organized internal
arrangement of thick and thin filaments (sarcomeres).
Thick filaments are composed of polymers of the protein myosin. Thin filaments are composed of polymers of actin,
and include the regulatory proteins troponin and tropomyosin. Sarcomeres also contain the giant protein titan. This
elastic protein extends the entire width of the sarcomere (~2 m) and functions as: (1) scaffolding, (2) elastic “spring”
to help the muscle recoil after having been stretched, and (3) some form of signal transduction within the muscle fiber
especially during hypertrophy induced by weight lifting.

MOLECULAR BASIS OF SKELETAL MUSCLE CONTRACTION

The sliding filament theory describes the molecular mechanisms responsible for contraction. During contractions, the
entire sarcomere shortens. Various components in the sarcomere shorten as well; this includes the H zone and the
I band, however the A band stays a constant length. Skeletal muscle contraction is voluntary and controlled by the
somatic nervous system. A neural signal results in Ca++ being released from the lateral sacs of the sarcoplasmic
reticulum. This Ca++ binds with troponin, which alters the shape of the troponin/tropomyosin complex unmasking the
myosin binding sites on the actin monomers. This results in the formation of the crossbridge where myosin binds with
actin. Once the crossbridge has formed, the powerstroke takes place with the displacement of ATP and P i from the
myosin head. The powerstroke progresses to form the rigor complex, a very strong bond between the actin and myosin
molecules. The powerstroke causes the thin filaments to be pulled over the thick filaments, shortening the sarcomere.
In order to release the bond between actin and myosin, ATP must bind to the myosin head and be hydrolyzed to ADP
and Pi. This sequence of crossbridge and powerstroke continues until the sarcomere is fully contracted.

Calcium is the link between excitation and contraction. When the neuron stimulates the muscle cell, a graded potential
is formed at the end plate (end-plate potential). This stimulates action potentials to propagate across the skeletal muscle
membrane. These action potentials plunge into the transverse tubules (invaginations of the cell membrane running
perpendicular to the long axis of the cell). Transverse tubules (T tubule) are in contact with the membranes of the
lateral sacs of the sarcoplasmic reticulum. In the membrane of the T tubules are voltage-sensitive proteins known as
dihydropyridine receptors (DHP). The DHP is activated by the action potentials. Activated DHP interact with Ca 2++
release channels (ryanodine receptors) in the lateral sacs of the sarcoplasmic reticulum. The ryanodine receptors open
to allow Ca2++ to leak into the cytosol where it can bind with the troponin, initiating the contractile cycle. Relaxation
occurs when there are no more action potentials. The deactivated DHP/ryanodine results in no more Ca2++ leaking from
the sarcoplasmic reticulum. ATP-dependent pumps remove the excess Ca2++ from the cytosol and return it to the lateral
sacs. This allows the sarcomere to passively return to its relaxed state.

Muscle Physiology 57
Contractile activity far outlasts the electrical activity that initiated it. A complete contraction/relaxation cycle (a twitch)
takes approximately 100 ms.

SKELETAL MUSCLE MECHANICS

Whole muscles are groups of muscle fibers bundled together and attached to bones. The connective-tissue covering of
the whole muscle extends beyond the ends of the muscle to form the tendon, which is attached to the bone at the origin
(stationary part of the joint) and the insertion (movable component of the joint). Tendons are considered to be series-
elastic components of the muscle and behave much like a stiff spring. Shortening of the sarcomere generates tension,
which is transmitted to the bone and in turn is the force applied to move the external load. Interactive units of skeletal
muscles, bones, and joints form lever systems.
There are three primary types of contractions: in isotonic contraction, the load remains constant as the muscle changes
length; in isokinetic contraction, the velocity of contraction remains constant as the muscle changes length; and in
isometric contraction, the tension develops at constant muscle length. All three contractions are generated by the same
interactions of thick and thin filaments.
Contractions can be described in two other ways. Concentric contraction occurs when the muscle shortens as the force
is applied, e.g., lifting a weight. Eccentric contractions occur when the muscle is lengthening as the force is applied,
e.g., lowering a weight. The velocity of contraction is related to the load being moved. In concentric contractions,
velocity of shortening is inversely related to load (the heavier the load, the slower the contraction), while in eccentric
contractions velocity of lengthening is directly related to load (the heavier the load the faster the lengthening).
Contractions of a whole muscle can be of varying strength. A single action potential will generate a weak short-
duration contraction called a twitch, which rarely occurs because it is too brief and weak to be useful. Because muscle
fibers are arranged into whole muscle, they can function cooperatively to produce purposeful movements. Two primary
factors generate whole-muscle tension: (1) the number of muscle fibers contracting, and (2) the tension developed by
each fiber.
The number of fibers contracting within a muscle depends on the extent of motor unit recruitment. The motor unit is
the total number of fibers innervated by a single motor neuron. Small motor units (12–50 muscle fibers/neuron) are
useful for delicate movements where precision is required (hand manipulations). Large motor units (1500–2000 muscle
fibers/neuron) are useful when powerful, coarsely controlled movements are required (postural movements).
The frequency of stimulation can influence the tension developed by each muscle fiber. If a muscle fiber is allowed to
completely relax prior to being stimulated again, the second twitch will be of the same magnitude as the first, providing
the same strength stimulus that was used. If the muscle fiber is stimulated again before it has completely relaxed, the
second contraction will piggyback on the first. This summation of events will lead to the production of greater force
than the single twitch, alone. This is called twitch summation. If the muscle fiber is stimulated so rapidly that it does
not have a chance to relax at all, a smooth, sustained contraction of maximal strength called tetanus is generated.
Twitch summation results from a sustained elevation in cytosolic calcium.
There is an optimal muscle length at which maximal tension can be developed. If a muscle is configured so that the
sarcomere is much shorter than optimal muscle length, or is stretched so much that the interaction of the actin and
myosin is interrupted, the tension will be less than optimal. This is called the length-tension relationship.
Although muscles can accomplish work, much of the energy is converted to heat. Approximately 25 percent of the
energy consumed is used for work, while 75 percent of the energy is converted to heat.

SKELETAL MUSCLE METABOLISM AND FIBER TYPES

Muscle fibers have alternate pathways for forming ATP. Creatine phosphate is the first energy storehouse used at the
onset of contractile activity. Creatine kinase is the enzyme responsible for liberating the energy needed for synthesizing
ATP. Aerobic or endurance exercise can be supported by the processes of oxidative phosphorylation. These processes
require adequate supply of oxygen so these muscles are replete with myoglobin and rich vasculature. These muscles do
not fatigue easily. Glycolysis is utilized during high-intensity exercise, when oxygen may be limited. These muscles
produce powerful contractions but fatigue easily. At times of intense exercise, the glycolytic process may overwhelm
oxidative phosphorylation, and the glycolytic end products (pyruvate) may build up in the muscle. The muscle will
convert the pyruvate to lactate by the process of fermentation.
Muscle fatigue occurs when the muscle can no longer respond to stimulation with equivalent contractile activity. This
mechanism prevents the muscle from being worked to the point where it can no longer generate ATP. The primary
factors involved include a local increase in inorganic phosphate, which reduces the strength of the myosin/actin
interaction and decreases the sensitivity of the troponin/tropomyosin complex to Ca ++. Additionally, the depletion of
glycogen stores plays a role in fatigue. Central fatigue occurs when the CNS no longer adequately activates the motor

58 Chapter Eight
neuron supplying the muscles. This is often psychologically based, stemming from the discomfort (pain) associated
with exercise. Boredom and monotony of repetitive low-intensity activities can also lead to central fatigue.
Increased oxygen consumption is necessary to recover from exercise. This excess postexercise oxygen consumption
(EPOC) is often due to an oxygen deficit created during strenuous exercise. EPOC may also be necessary to process
nutrients depleted during exercise and can be due to increased body heat caused by exercise. Increased body heat
accelerates oxygen-consuming chemical reactions. Epinephrine is released during exercise as a component of the
sympathetic involvement. This hormone increases oxygen consumption and can lead to EPOC.
There are three types of skeletal muscle fibers, based on differences in ATP hydrolysis and synthesis: slow-oxidative
(type I) fibers, fast-oxidative (type IIa) fibers, and fast-glycolytic (type IIx) fibers. Fast fibers have a higher myosin
ATPase activity than slow fibers do. This allows fast fibers to have a faster crossbridge cycling rate (15–40 ms to peak
tension) than do slow fibers (50–100 ms to peak tension). Oxidative fibers (red fibers) are better equipped to utilize
oxidative phosphorylation for the synthesis of ATP; therefore these fibers are more resistant to fatigue. Glycolytic
fibers (white fibers) are equipped with high content of glycolytic enzymes and less oxidative capacity. These fibers
tend to have explosive contractions, but are easily fatigued.
Muscle fibers adapt considerably in response to the demands placed on them. Regular aerobic exercise will promote
metabolic changes within oxidative muscle fibers, allowing them to more efficiently use O2 and therefore better endure
prolonged exercise without fatiguing. Muscle fibers can also respond to regular bouts of anaerobic, short-duration,
high-intensity resistance training by undergoing hypertrophy. This is an enlargement of the muscle due to increased
synthesis of actin and myosin filaments in the fast-glycolytic fibers. It is also possible for the interconversion of fast-
glycolytic and fast-oxidative fibers depending on the type of regular exercise the muscle endures. It is not possible for
the interconversion of fast and slow fibers.
Muscles may atrophy (decrease mass) due to: (1) disuse atrophy, seen in muscles not used for long periods of time,
such as when a limb is in a cast; (2) denervation atrophy, if the nervous input to the muscle is lost, atrophy will ensue;
and (3) age-related atrophy or sarcopenia, which begins at about age 40 with progressive loss of motor neurons,
especially those innervating fast-glycolytic fibers.

CONTROL OF MOTOR MOVEMENT

Multiple neural inputs influence motor neuron output. Three levels of input to motor neurons control their output: (1)
input from afferent neurons (spinal cord); (2) input from primary motor cortex from cell bodies known as pyramidal
cells (corticospinal or pyramidal motor system); and (3) input from the brainstem (extrapyramidal or multineuronal
motor system).
Muscle receptors provide afferent information needed to control skeletal muscle activity. There are two types of muscle
receptors. (1) Muscle spindles (known as intrafusal fibers), which monitor muscle length and are involved in certain
reflex movements (knee-jerk reflex). Each muscle spindle is innervated by the gamma-motor neuron, whereas the
extrafusal (ordinary contractile cells) are innervated by the alpha motor neuron. Two types of afferent sensory endings
terminate on the intrafusal fibers, primary (annulospiral) endings and secondary (flower-spray) endings. These sensory
components play a role in the stretch reflex mechanism. (2) Golgi tendon organs, which monitor changes in muscle
tension and send this information to the conscious regions of the brain to make one aware of the tension in the
particular muscle. Recent work refutes the idea that Golgi tendon organs illicit protective spinal reflexes and halt
further contractions, but instead they function as a pure sensor for relaying tension information.

SMOOTH AND CARDIAC MUSCLE

Smooth and cardiac muscle share some basic properties with skeletal muscle. All three types of muscle have a
specialized contractile apparatus made up of thin actin filaments and thick myosin filaments, which interact in response
to increases in intracellular Ca++. They also directly use ATP to power contractions. There are specific structural
differences among the three muscle types. Smooth muscle cells are small and unstriated. There are three filament types:
(1) thick myosin filaments, which are longer than in skeletal muscle; (2) thin actin filaments, which contain
tropomyosin but no troponin; and (3) intermediate filaments, which do not participate directly in contraction, but are
involved with supporting the cytoskeletal framework of the cells. These myofilaments are not arranged in sarcomeres
and thus show no banding (not striated). With no Z lines for anchoring the thin filaments, smooth muscle utilizes dense
bodies, which are located by the intermediate fibers. The thin and thick filaments are oriented slightly diagonally from
side to side in the cells in a diamond-shaped lattice. Thick filaments are arranged such that crossbridges are present
along the entire length of the thick filament, providing a longer distance that thin filaments can be pulled along the
thick filament.
Smooth muscle cells are turned on by Ca2-dependent phosphorylation of light chains of proteins associated with the
myosin heads. Cross bridging cannot occur unless the myosin light chain kinase has been activated.

Muscle Physiology 59
Smooth muscle can be grouped into two categories depending on the pattern of contractile activity. Phasic smooth
muscle contracts in bursts and is most abundant in hollow organs that push contents through them, such as the digestive
organs. Tonic smooth muscle is partially contracted at all times. This contractile state is called tone. There is no
bursting of activity, but rather there is incremental increase/decrease in contractile intensity. The smooth muscle in the
walls of arterioles is tonic. Multiunit smooth muscle has multiple independently functioning units that must be
stimulated by autonomic neurons to contract (neurogenic). Multiunit smooth muscle is found: (1) in walls of large
blood vessels; (2) in small airways to the lungs; (3) in the muscle of the eye that adjusts the lens; (4) in the iris of the
eye, which adjusts pupil size; and (5) at the base of hair follicles, which is responsible for producing “goose bumps.”
Single-unit smooth muscle cells form functional syncytia, which is when multiple cells function as if they are one large
unit. These cells are myogenic or self-excitable because they produce one of two spontaneous depolarizations:
(1) pacemaker potentials, or (2) slow-wave potentials. Gradation of single-unit smooth muscle contraction is dependent
on the number of crossbridges formed rather than the number of motor units recruited (as in skeletal muscle).
Because of the arrangement of myosin along the thick filaments, smooth muscle can still develop tension when
stretched. This is evident as the urinary bladder fills and stretches the smooth muscle in the wall, and can still develop
tension when stimulated. While smooth muscle will relax when gradually stretched, it responds to a sudden stretch by
initially increasing tension (likely due to passive rather than active contraction), then quickly adjusting to the new
length (stress relaxation response).
Smooth muscle is slow and economical as the ATPase activity of the smooth muscle is much slower than in skeletal
muscle.
Cardiac muscle blends features of both skeletal and single-unit smooth muscle. The general intracellular architecture is
like skeletal muscle, while the Ca++ functions like single-unit smooth muscle. Cardiac muscle is striated, but
contractions are involuntary. Unique to cardiac muscle is that the cells are joined together in a branching network.

LIST OF KEY TERMS

Skeletal muscle structure - ryanodine receptor - oxidative phosphorylation

- myoblasts - dihydropyridine receptor - aerobic exercise
- striated - crossbridges - endurance-type exercise
- contractile component - power stroke - myoglobin
- thick filament - rigor complex - anaerobic exercise
- contractile proteins - relaxation - high-intensity exercise
- myofibrils - insertion - glycolysis
- myosin - origin - glycogen
- thin filament Skeletal muscle mechanics - lactate
- actin - tension - fatigue
- regulatory proteins - fatigue - central fatigue
- tropomyosin - asynchronous recruitment - neuromuscular fatigue
- troponin - motor recruitment - EPOC
- sarcomere - twitch - oxygen debt
- A band - twitch summation Skeletal muscle fiber types
- H zone - tetanus - slow-oxidative (type I)
- I band - length-tension relationship - fast-oxidative (type IIa)
- M line - optimal length - fast-glycolytic (type IIx)
- Z line - load - red fiber
- sarcoplasmic reticulum - concentric contraction - white fiber
- lateral sacs/terminal cisternae - eccentric contraction - atrophy
Skeletal muscle contraction - isometric contraction - denervation atrophy
- neurogenic - isotonic contraction - disuse atrophy
- alpha motor unit - load-velocity relationship - hypertrophy
- sliding-filament theory - lever Control of motor movement
- excitation-contraction - fulcrum - corticospinal motor system
coupling - load arm - pyramidal motor system
- neuromuscular junction - power arm - extrapyramidal system
- motor end-plate Skeletal muscle metabolism - spastic paralysis
- transverse tubule - ATP - flaccid paralysis
- foot-protein Ca2+ channels - creatine phosphate - hemiplegia

60 Chapter Eight
 - quadriplegia - multineuronal motor system - neurogenic
- paraplegia - primary endings - single-unit smooth muscle
- dystrophin - secondary endings - visceral smooth muscle
- muscular dystrophy - coactivation - functional syncytium
- utrophin - stretch reflex - myogenic activity
- intrafusal fibers - patellar tendon reflex - self-excitable
- extrafusal fibers - Golgi tendon organs - visceral smooth muscle
- muscle spindles Smooth muscle - slow-wave potentials
- extrafusal fibers - dense bodies - stress-relaxation response
- intrafusal fibers - phasic smooth muscle - latch phenomenon
- alpha motor neurons - tonic smooth muscle Cardiac muscle
- gamma motor neurons - multiunit smooth muscle

LECTURE HINTS AND SUGGESTIONS

1. This chapter offers an excellent opportunity to illustrate the structure-function relationship between the striated
nature of skeletal muscle and its contractility. Relate the striations to the arrangement of myosin and actin in the
sarcomeres. Describe the A band and I bands of the sarcomere. Relate the arrangement of the muscle proteins to
the sliding-filament model of skeletal muscle contraction. Models of sarcomeres are available from biological
supply companies.

2. Electron micrographs of muscle fibers, myofibrils, and myofilaments are particularly useful here to enhance
instruction.

3. Demonstrate isometric versus isotonic contraction. See any general physiology lab manual for procedures. An
electronic muscle stimulator can also be used.

4. Show and/or record various physiological muscle responses; e.g., tetanus and muscle twitch. The gastrocnemius
muscle and sciatic nerve from a frog leg attached to a kymograph or physiograph is the classical approach.
Recordings can also be made through the use of a physiograph. See any general physiology lab manual for
procedures.

5. The importance of ATP in muscle contraction can be demonstrated using live muscle tissue and altering ATP,
potassium, and magnesium levels. Kits are available from biological supply companies.

6. It is important to get the students physically involved in the discussion. Use a student volunteer to demonstrate
skeletal muscle contractions and point out the important events as they happen. If available, the students can use
the muscle grip tester to demonstrate strength of contraction as well as muscle fatigue.

7. A clinical dimension can be added by describing motor impairments resulting from trauma at various muscle
innervation sites; e.g., polio and tetanus.

8. Describe the general functions of muscles to the students. They must understand that muscles do more than
contract. For example, posture, heat production, and glycogen storage are also functions of muscles.

9. Use various cuts of meat to show arrangement of fascicles within muscles. Relate “dark” and “white” meat of
chicken to the various types of muscle fibers.

10. Be sure to remind students of the learning resources available on the textbook website and of the literature-
searching capability of InfoTrac®.

Muscle Physiology 61
AUDIOVISUAL AIDS

Videos/Films
Following are films that may be suitable for presentation in your class.

The Brooks/Cole Human Physiology Video Library.

Mark A. Paternostro, 2011. Clip #2: How Muscles Hold Tension.
(Request this supplement from your local Cengage Sales Representative.)

http://ffh.films.com

Bones and Muscles, 15 min.

This program stresses the importance of maintaining one’s strength for good health. On a field trip to a
biopharmaceutical company, we learn of a new treatment for osteoporosis that promotes bone growth.

Human Facial Expression, 15 min.

This program examines the muscle movement responsible for human expressions.

Muscles, 20 min.
In this program, the diverse nature of muscle tissue is examined, from its gross structure to its detailed
microstructure, where chemical energy is harnessed to produce movement.

Motor Development, 25 min.

This program presents positioning and handling techniques for both the hypertonic profile child, who is
easily overstimulated and has stiffening of the limbs, and the hypotonic profile child, who has flaccid muscle
tone and decreased movement.

Moving Parts, 26 min.

This program uses dramatizations to illustrate the interactions of muscles with other body systems.

Muscular System at Work: The Inner Athlete, 24 min.

Topics include muscles and movement; cardiac, smooth, and skeletal muscle; detailed structure of a skeletal
muscle; types of muscle contraction and movement; muscles and posture; homeostasis; and the important
roles played by skin, hair, nails, and glands.

Steroids and Sports, 19 min.

This program analyzes the effects of using anabolic steroids for sports gains on muscle and other body organs.

The Physiology of Exercise, 15 min.

This program examines changes in the body as individuals exercise.

http://www.biol.sc.edu/~vogt/comp-phys.html

Comparative Physiology, streamed video courses. Free.

http://www.carolina.com

Human Machine, eyewitness video, 30 min.

Investigate the most complex machine of all—a light, flexible, immensely strong framework that laughs, cries,
and blushes. Travel inside the body to explore the vital organs that power our lives, and examine the
remarkable instruments through which we sense the world.

http://www.youtube.com/watch?v=aYCUMbzSKfY

Smooth Muscle. YouTube video. 1 min.

Images of smooth muscle grown in culture. Related videos available.

62 Chapter Eight
Software
Biochemistry of Muscle, EI, covers contraction and relaxation.
BIOPAC Muscular Function, BIP, hardware and software to measure EMG, strength, summation, etc.
BIONET, http://www.bioscience.heacademy.ac.uk/hosted.
Dynamics of the Human Muscular System, EI, presents muscle types and muscle contraction.
Electronic Image Collection for Cell Biology, http://www.ramex.com.
Exercises in Muscle Contraction, EI, animated exercises in muscles stimulation and contractions.
Flash: Skeletal Muscles, PLP, surveys more than 200 muscles.
Interactions CD-ROM Series for Anatomy and Physiology, Muscular and Skeletal Systems Disk #3, JH.
Location, CBS, reviews functions of bones and muscles and covers types of muscles and joints.
MRI of the Musculoskeletal System, RAM, an interactive CD.
Muscular System, PLP, three-part program covering human muscular system.
Neuromuscular Concepts, PLP, covers skeletal muscle contraction beginning with neuromuscular junction.
PHYSIOEX, AW, simulated lab exercises, http://www.physioex.com.
Skeletal Muscle Anatomy and Physiology, PLP, covers three muscle types, lever systems, and sliding-filament theory.

Relevant Educational Websites

http://ajpcon.physiology.org/
Homepage of the American Journal of Physiology.

http://nmrc.bu.edu/
Homepage for the Neuromuscular Research Center at Boston University. This page has hyperlinks to
information pages.

http://www.3dotstudio.com/zz.html
Animation of the contraction of a sarcomere.

http://www.neuro.wustl.edu/neuromuscular/mother/acetylcholine.htm
Jump-off page with hyperlinks to detailed examination of the function of ACh and ACh receptors.

http://muscle.ucsd.edu/
Homepage for the University of California San Diego muscle physiology lab. Students can hyperlink to
detailed descriptions of many important concepts of muscle anatomy and physiology.

http://www.uic.edu/classes/phyb/phyb516/smoothmuscle.htm
A detailed analysis of smooth muscle function with hyperlinks to illustrations and other information.

http://thalamus.wustl.edu/course/spinal.html
A discussion of spinal cord and motor neurons, and includes photomicrographs and cartoons.

http://lessons.HarveyProject.org/development/muscle/menu.html
Animated tutorial of muscle structure and sarcomere function.

http://imc.meded.com
A subscription site with in-depth discussions of physiology.

Relevant Organizations Providing Educational Resources

The ALS Association
27001 Agoura Rd., Ste. 150
Calabasas Hills, CA 91301
http://www.alsa.org

Muscle Physiology 63
International Society for Musculoskeletal and Neuronal Interactions
http://www.ismni.org/

Muscular Dystrophy Association

3300 East Sunrise Dr.
Tucson, AZ 85718
http://www.mdausa.org

Musculoskeletal Transplant Foundation

125 May Street
Edison, NJ 08837
(732) 661-0202
http://www.mtf.org/

National Institute of Arthritis and Musculoskeletal and Skin Diseases

National Institutes of Health
1 AMS Circle
Bethesda, MD 20892-3675
http://www.nih.gov/niams

Answers to End of Chapter Essays

1. Levels of organization from larger to smaller include whole muscle, muscle fiber (cell), myofibril
(specialized intracellular structure), sarcomere (functional unit of skeletal muscle), thick and thin
filaments (cytoskeletal elements), myosin and actin (protein molecules).

2. The striated appearance, or banding pattern, of skeletal muscle is due to the alternating bands of thick and
thin filaments. The A band (dark band) is made up primarily of thick filaments and some overlapping thin
filaments. It lies in the center of the sarcomere. Within the A band is a central H zone, which is the
portion of the A band where the thin filaments do not exist. The I band consist of the thin filaments not
part of the A band. On either side of the I band, between sarcomeres, are dense Z lines.

3. The functional unit of skeletal muscle is the sarcomere. It is the smallest component of the muscle that
can perform all of the functions of the whole muscle.

4. Each thick filament is composed of several hundred myosin molecules packed together. The protein’s
tails are intertwined with each other in the center of the thick filament and the heads are staggered along
the ends of the thick filament. Heads are radially arranged around individual thick filaments with 60
degrees between adjacent heads. Sets of myosin molecules come together so that one side is a mirror
image of the other. The thin filaments are composed primarily of actin, with regulatory proteins troponin
and tropomyosin wrapped around the actin polymers. Each actin is a small, spherical unit, and multiple
actin molecules are attached to one another end to end into two strands. The strands are then twisted
together (like a strand of pearls twisted together). Tropomyosin is a threadlike protein that lies in the
grooves of the actin strand. Troponin is a small three-subunit protein that lies at regular intervals along the
actin strand.

5. The thin filaments on each side of the sarcomere slide inward to its center. As they do so, they slide past
the stationary thick filaments that lie at the center of the sarcomere. As the two filaments slide past one
another, the Z lines on each side are brought closer together and the entire sarcomere shortens. This is
referred to as the sliding filament theory because the thick and thin filaments slide past each other.
Crossbridge powerstrokes provide the energy for the movement associated with sarcomere shortening.
When myosin and actin connect at the crossbridge, the head of the myosin bends inward toward the center
of the sarcomere, pulling the thin filament and the attached Z line toward the center, shortening the
sarcomere.

64 Chapter Eight
6. Calcium is the link between excitation and contraction. In skeletal muscle, calcium is released from the
sarcoplasmic reticulum upon stimulation from action potentials passing down the transverse tubules. The
calcium binds to troponin, causing a conformational change. As troponin changes shape it causes
tropomyosin to change its shape, physically moving out of its blocking position. Myosin crossbridge can
now attach to the exposed actin binding sites. The binding triggers the crossbridge to bend and the
powerstroke occurs. In smooth muscle, upon stimulation, calcium moves from the ECF into the cytosol.
Calcium binds to calmodulin, which in turn activates myosin light chain kinase. This enzyme
phosphorylates a myosin crossbridge, which can now bind with actin.

7. Two factors can be changed to produce graded contractions: the number of muscle fibers contracting in
the muscle and the tension developed by each contracting fiber.

8. A motor unit is one motor neuron and all the muscle fibers it activates. Small motor units (those with a
smaller number of muscle fibers) are associated with muscles that produce delicate, precise movements.
Large motor units containing a large number of cells (1,500–2,000) are associated with coarse, powerful
contractions. In motor unit recruitment, more and more motor units are recruited to contract to produce
more powerful contractions.

9. Twitch summation and tetanus can be achieved by increasing the frequency at which a muscle is
stimulated to contract. This causes the muscle to contract before it can completely relax so that
contractions are added, or summed up, on each other. This action produces a stronger contraction than a
single stimulation. In tetanus, the muscle is stimulated so quickly that it does not have the chance to relax
at all between stimulations and a smooth, maximal strength contraction occurs.

10. Maximal contraction strength can be achieved when a muscle is at its optimal length before contracting.
This optimal length is usually the resting length of the muscle found in the body. When the muscle is
shortened or stretched beyond this optimal length it produces a weaker contraction. This is because when
stretched or shortened fewer thin-filament binding sites are exposed to thick-filament crossbridges.

11. In isotonic contractions muscle tension remains constant as the muscle length changes. During isokinetic
contraction, the velocity of shortening remains constant as the muscle length changes. In isometric
contraction muscle length remains constant as tension develops.

12. ATP provides energy for the powerstroke of the crossbridge, promotes crossbridge detachment so that the
cycle can be repeated, and provides energy for the active transport of calcium into the sarcoplasmic
reticulum during relaxation. Creatine phosphate is the first energy system utilized at the beginning of
muscle activity. It donates a phosphate group to ADP to form ATP. Oxidative phosphorylation takes
place in the mitochondria and provides the largest amount of ATP to the contracting muscle. It is aerobic
and relatively slow because it involves a large number of steps. Glycolysis reactions occur in the cytosol
and are anaerobic, so they can rapidly provide energy to produce ATP in the absence of oxygen.
However, significantly less ATP is produced compared to oxidative processes.

13. The three types of skeletal muscle fibers are slow oxidative (type I), fast oxidative (type IIa), and fast
glycolytic (type IIx). Slow oxidative fibers have slow myosin-ATPase activity and are relatively slow to
contract but resistant to fatigue. Because they are oxidative they have many mitochondria, and have a
large blood flow and myoglobin content. This causes the fiber to have a red color. Fast oxidative fibers
are similar to the slow oxidative type except that the myosin-ATPase activity is higher and they are more
susceptible to fatigue. The fast glycolytic fibers have fast myosin-ATPase activity and are thus fast to
contract but easily fatigued. Because they are glycolytic and not oxidative they have few mitochondria
and a smaller blood flow and myoglobin count. This causes the fiber to have a white color.

Muscle Physiology 65
 14. The corticospinal tracts originate in the primary motor cortex and send axons that terminate on motor
neurons in the spinal cord. This system influences the final common pathway for motor activity. The
multineuronal motor system includes many regions of the brain including the reticular formation of the
brain stem. This area, in turn, receives input from the cortex, cerebellum, and basal nuclei. This system
controls overall body posture involving involuntary movements of large muscle groups of the limbs and
trunk.

15. Muscle spindles detect changes in the length of the muscle. They consist of a collection of specialized
muscle fibers called intrafusal fibers. At the center of these fibers is a non-contractile element innervated
by afferent sensory neurons. These fibers lie within spindle-shaped connective tissue capsules that lie in
parallel to the extrafusal fibers. The sensory fibers are activated when the muscle is stretched. Golgi
tendon organs are found in the tendons of muscles and respond to changes in muscle tension. Golgi
tendon organs consist of afferent nerve endings intertwined with bundles of connective tissue fibers that
make up the tendon.

16. Phasic smooth muscle contracts in bursts of activity caused by action potentials. It produces alterations in
contraction and relaxation. This muscle is found in the walls of hollow organs of the digestive system and
helps mix and propel food during digestion. Tonic smooth muscle is usually partially contracted at all
times. This is called tone. This type of muscle is found in the walls of arterioles and the tone contributes
to maintenance of blood pressure in these vessels.

17. Multiunit smooth muscle consists of multiple discrete units that can act independently of one another.
These units must be stimulated independently to produce contractions. Single-unit smooth muscle consists
of muscle fibers that contract simultaneously as a single unit. Gap junctions are responsible for coupling
the cells together into one unit (functional syncytium).

18. In neurogenic muscle, contractions are initiated only upon stimulation from nerves that innervate the
muscle. Multiunit smooth muscle is neurogenic. In myogenic muscle, electrical activity in pacemaker
cells spreads to other cells via gap junctions and the entire unit contracts. Single-unit smooth muscle is
myogenic.

19. Because gap junctions ensure that all muscle fibers contract simultaneously in single-unit smooth muscle,
it is impossible to vary the number of fibers that contract at any time. Graded contractions can be
produced by altering the tension produced in the muscle. This is accomplished by varying the cytosolic
calcium concentration. As calcium levels increase, more crossbridges are activated and a greater tension
develops.

20. In general, smooth muscle is slower and more energy efficient than skeletal muscle. Myosin-ATPase
activity is slower in smooth muscle, and smooth muscle relaxes more slowly than skeletal muscle because
calcium is removed at a slower rate from the cytosol. Despite being slower, smooth muscle can generate
the same force per unit area as skeletal muscle, but does so with less energy expenditure. Since
crossbridge cycling is slower in smooth muscle it uses less ATP over time.

21. Cardiac muscle is similar to skeletal muscle in that it is striated, and contains troponin and tropomyosin as
well as sarcoplasmic reticulum and transverse tubule system. It also has a similar length-tension
relationship. Cardiac muscle is similar to single-unit smooth muscle in that some of the calcium that is
necessary for contraction enters from the ECF, it relies on pacemaker activity to initiate contractions, and
cardiac cells are interconnected via gap junctions so electrical activity easily spreads between cells,
resulting in a synchronous contraction.

66 Chapter Eight
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 miles distant from Platæa. It cannot, in fact, have ever
entered into the calculations of the Greek generals.

As regards the three passes which lead on to the field, can we, if
we bear in mind the condition of the Greek army at the time of the
movement, suppose for a moment that the Greek generals would
move to a position farther away from the passes than they were
before the movement began? Surely their natural course would be to
retire to the higher ground, where they would be in comparative
safety from the Persian cavalry, and where, in the position I have
indicated, they would be exactly between the passes on the Platæa-
Megara and Platæa-Athens roads respectively, and where, too, they
would be in easy communication with both by way of the rocky part
of the Kithæron slope.
The last piece of evidence bearing on this point is afforded by the
remarkable tale of Amompharetos, the Spartan officer who refused
to move from the second position with the rest of the Lacedæmonian
force. The reason which prompted (H. ix. 53) him in his refusal is
stated by Herodotus to have been the disgrace of retreating before
the enemy.
The feeling is hardly comprehensible if the movement was to be
to the “Island” in the plain; it is comprehensible if it was to be towards
Kithæron, since he might well suspect that the real design of
Pausanias was to retreat through the mountain into Attica.

At the time at which this resolution was taken the Greeks were for
the moment a beaten army. What might have happened had the
proposed movement ever become an actual fact cannot of course be
said, but it is extremely probable that a leakage through the passes
would have commenced, by which its numbers would have been so
rapidly reduced that an absolute withdrawal from Bœotia would have
shortly become necessary. Had that been the case, Northern Greece
must have been lost. It would have been very difficult after what had
happened to keep together so composite a force in an advanced but
purely defensive position. As far as the Greek generals were
concerned, their design, doubtless, was to remain on the defensive
in Bœotia, now that their offensive strategy had not met with
success, and to starve the enemy out of the country. Alexander of
Macedon had reported to them the approaching failure of the
Persian supplies.
They had failed in attempting to take the offensive in a country
which was peculiarly adapted for the operations of cavalry. They had
been led by the success they had achieved in the first position into
the mistake of remaining in a position which could only be justified by
an active offensive, not recognizing that their success on that
occasion had been due to the wrong use which Mardonius or
Masistios had made of their numerous cavalry. The passes formed,
as it were, their immediate base; and they had not recognized that
by leaving an interval between themselves and that base they
exposed their line of communications to the attacks of an arm
against whose mobility they had nothing to match, and had rendered
themselves assailable from all sides by a mode of assault which
made it impossible for them to retaliate in adequate fashion. It was
their first experience of the possibilities of cavalry action on a large
scale; and it is noticeable that in the many subsequent years of
warfare with Persia in that century they never again willingly exposed
themselves to the possibility of such a reverse as they had suffered
in the second position at Platæa.
The curious piece of land, with its curious name of the “Island,”
which was fixed upon by the Greek commanders as the goal of the
proposed movement, is easily recognizable at the present day,
owing to the accuracy with which Herodotus describes it. It is one of
those ridges which extend northwards into the plain from the foot of
the rocky slope of Kithæron; but it is remarkable among them for two
peculiarities. The streams which bound it on
LAST HOURS ON
THE ASOPOS
either side have their sources on the mountain
RIDGE. slope above it within but a few yards of one
another, so that it may be said to be all but
surrounded by water; and the ordinarily rounded back of these
northward stretching ridges is varied in this case by a steep and very
noticeable natural mound which rises upon it. It is nearly two miles
due south of the ridge which formed the Greek second position, and
about one mile due east of the town of Platæa. As a position it
offered the distinct advantage of being within touch of the Platæa-
Athens and Platæa-Megara passes, and of affording the army
comparative immunity from cavalry attack.
The Greek commanders were disappointed in their hope or
expectation that the cavalry attack would not be renewed on that
day, on the morning of which the Council of War was held. The
Persians had plain evidence of the effectiveness of the method they
were employing; and for the whole of a second day the Greeks had
to support a series of assaults without being able to inflict any
proportionate damage on the foe. Those who have been in the plains
of Bœotia during the heat of the late summer and early autumn will
best appreciate the terrible hardships which the army suffered. For at
least forty hours it must have been wholly cut off from water, and for
a considerable part of that time must have been undergoing severe
exertion and supporting severe losses under a burning sun upon that
dusty, rolling plain. It is evident indeed that the Persian cavalry
withdrew during the night; but they had, before doing so, rendered
the Gargaphia spring unusable. The longed-for darkness came at
last, and at the appointed hour the movement began. It began,
indeed,—but it ended vary differently from what had been intended.
With the exception of the Spartans and Athenians, the Greek army,
without waiting for further orders, started off in haste at the time
appointed, —but not to the “Island,” so says Herodotus. They never
even thought of so doing, but were glad to take refuge just outside
208
the town of Platæa, around the temple of Hera.
It is quite plain that Herodotus believed that the centre of the
Greek army fled in something resembling a panic, and neither
obeyed, nor intended to obey, orders. When the story is examined it
is found to contain several features which are very difficult to
reconcile with the colouring which Herodotus has given to it. If these
Greeks intended to fly, why did they not make directly for the Platæa-
Megara pass, which must have been open? Why did they go to
Platæa?
Having gone thither, why did they not seize upon the town?
 It has been suggested that in going thither they were carrying out
orders, and that the position at the Heræon was part of the position
of which the “Island” was another part. The nature of the ground
between the two positions, and their distance apart, does not render
this very probable.
The probability is that, like so many stories of incidents in Greek
history, this particular one has suffered from the exaggeration of
some of its true details, and the loss of others.
It is not strange that the events of the last two days’ fighting on
the Asopos ridge should have created disposition to panic
throughout the Greek army. Doubtless such a disposition did exist,
and showed itself in the hurried departure of a large portion of the
army. But even Herodotus does not say that the departure was
premature. So far, the tale may give a true representation of the facts
of the case. What in all probability took place
RETIREMENT OF
GREEK CENTRE.
was that the retiring force started without
taking due precautions to ascertain the exact
line of retreat, or to provide itself with guides who could lead it amid
the darkness of the night to the position which had been appointed
as the rendezvous of the army. If that were so, a mistake was not
merely possible, but almost inevitable. In that country of rise and fall
which lies between Kithæron and the Asopos, one slope resembles
another, and one stream is similar in character to its neighbours, so
that there is a conspicuous absence of those landmarks which may
serve as a guide to one who traverses it by night.
It is not difficult to imagine their position when they found
themselves at the Heræon. They must have arrived there in the
night, and may have thought it well to wait until dawn disclosed the
true position of the rallying point of the Greek army. From the
Heræon the “Island” would be full in sight, when the advance of
daylight had rendered the surrounding country visible. But the dawn
can have brought but little comfort to them. It is more easy to realize
the scene that met their eyes than the intensity of this anxiety which
that scene aroused. Away to the east the “Island”—empty,
unoccupied. To the north-east the Persian army pouring in streams
over the ridge which they had evacuated during the night, and from
the hollow beneath that ridge two clouds of dust arising, sole
indication of the terrible struggle which was going on beyond the
intervening ridges. This is not merely a picture of the imagination and
no more. It is what these men must have seen from their position at
Platæa; for the narrative of Herodotus indicates with sufficient
clearness the time and the locality of the battles which were fought
on that fatal morning by the right and left wings of the Greek army.
It would be unjustifiable to pass a decisive judgment on the
conduct of those Greeks. The extent of their cowardice is not indeed
calculable. The tradition which Herodotus followed tended manifestly
to depict the whole attitude of the Peloponnesian Greeks, both on
this and other occasions in the war, in the most unfavourable light.
There can be little doubt that the tradition was Athenian in origin.
It is not, however, necessary to assume that in this or any other
particular instance the truth was distorted in order to cloke Athenian
failings. The animosities of that later time at which Herodotus wrote
his history are quite sufficient to account for the discoloration of
facts, and in this particular instance there does not appear to be any
valid reason for supposing that the Athenians had anything to
conceal.
Much had happened during the night of which the Greek centre
was wholly unaware.
Pausanias on the right wing, on seeing that the troops in the
centre had begun the movement of retirement, gave orders to the
Spartans to do the same. He supposed that the other Greeks would,
according to orders, withdraw to the “Island;” and, as has been seen,
it is probable that the latter had every intention of so doing.
A question now arises which is of very great importance with
regard to the history of the battle. Was it the intention of Pausanias
that his Spartans on the right wing should likewise withdraw direct to
the “Island,” or did he intend to carry out some other operation
before joining the rest of the army in that position? Herodotus, whose
attention is at this point in his narrative distracted by the not
unimportant tale of Amompharetos, makes no direct statement on
the subject, but he furnishes sufficient indirect evidence to render it
almost certain that the march of the Spartans neither was, nor was
intended to be, directed straight to the “Island,” but aimed first at the
accomplishment of a design which was at the moment of the utmost
importance to the army. It is further noticeable that the indirect
evidence of Herodotus is peculiarly supported by the light which is
thrown upon the further incidents of the battle by the topography of
the field.
In describing the resolution to move to the “Island,” Herodotus, it
will be remembered, says that it was the intention of the Greek
commanders to send half the army to relieve the provision trains
which were blocked in the passes; and the other evidence shows
that the Spartans’ movement from the second position had this
209
immediate object in view.
When, therefore, Pausanias saw the Greek
OBSTINACY OF
AMOMPHARETOS.
centre was starting for the “Island,” he gave
orders to his division of forty thousand
Spartans and helots to move also, intending to march towards the
pass on the Platæa-Athens road, and, possibly, towards
Dryoskephalæ pass also. For the moment, however, his plan was
completely upset by the obstinacy and insubordination of one of his
captains, Amompharetos, who, so Herodotus says, commanded the
210
Pitanate division of the Lacedæmonian force. The refusal to obey
was evidently based on the supposition that the withdrawal from the
211
position was nothing less than a preliminary to a retreat. Orders
and expostulations proving alike unavailing, persuasion was tried,
but with no better effect. The whole night seems to have been
wasted in the dispute, for it was not until day was dawning that
Pausanias moved away, reluctant to leave the regiment to its fate,
but hoping that when Amompharetos found himself alone he would
make up his mind to follow. The Tegeans accompanied the Spartans
in their movement.
That movement is described in some detail by Herodotus. While
the Athenians, who had been suspiciously waiting for the Spartans to
move, turned down towards the plain, the latter kept to the hills, and
 212
made for the slopes of Kithæron, from fear of the Persian cavalry.
Amompharetos, on seeing that the rest of the Spartan army was
leaving him in good earnest, started slowly after it, which movement
being perceived by Pausanias, he stopped the main body in order to
wait for him. He had by that time advanced ten stades, or a little
more than a mile, from his previous position, and he waited at a
place which is described as having been near the river Moloeis, in
what was called the Argiopian country, where was a temple of
Eleusinian Demeter.

The Temple of Eleusinian Demeter, etc.

Herodotus, in the words recently quoted, gives the following
facts:⁠—
(1) That Pausanias, with the main body of the Spartans,
advanced ten stades, and then waited for Amompharetos and his
regiment.
(2) That he waited at the river Moloeis, in what is called the
Argiopian country.
(3) That there is a temple of Eleusinian Demeter in that part.
To which may be added from an earlier chapter:⁠—
(4) Pausanias, after failing to persuade Amompharetos, starts at
dawn: his movements Herodotus describes as having been as
follows:⁠—

H. ix. 56.
“He gave the signal and led away the whole
of the rest of his force through the hills; the
Tegeans also followed him.... For the Lacedæmonians
intended to keep to the hilly ground and the foot-slopes of
Kithæron, because they feared the cavalry.”
Circumstances (1) and (2) may be taken
TEMPLE OF
ELEUSIAN
together.
DEMETER. This is the first mention of the river Moloeis.
 Nothing is known of the position of the Argiopian country; but it
may be surmised that it had some more or less striking characteristic
which caused it to be called by a special name.
Any identification of these two geographical features must be a
matter of extreme uncertainty.
After my stay at Kriekouki in 1892–1893, I was disposed to
identify the brook A 5 with the river Moloeis. I am inclined, after
revisiting the country in August, 1899, to think that the brook A 6 is
more probably the stream mentioned by Herodotus. It is the largest
of the streams which traverse the field, save, of course, the Asopos
itself, and Leake, in his map, assigns to it the name of the main
stream. Though the Spartan army after advancing ten stades from
the Asopos ridge would not be actually on this brook, yet, even
granting to the measurement given a greater accuracy than perhaps
it can claim, the position attained might be described as περὶ
ποταμὸν Μολόεντα.
The matter, however, is so uncertain, that it is impossible to insist
on either of these two identifications against the other.
The name Argiopian has been supposed to mean “white rock,”
and, adopting this interpretation, one visitor to the region has
discovered that there is a patch of white rock on the side of Kithæron
which is a prominent object from the plain. As we were in August,
1899, encamped in the plain near the head of brook A 5, I had ample
opportunity for observing the north slope of Kithæron. No such patch
was then visible on the dull grey-green rock slopes of the mountain.
As to the interpretation of the name I am, I confess, sceptical. The
interpretation of place-names in Greece, as elsewhere, is a most
dangerous though fascinating pastime; and it is but too often the
case that the most obvious interpretation is the most incorrect. All, I
think, that can be safely assumed in the present instance is that the
Argiopian country was a stretch of land marked off in some way from
its surroundings, and I venture to think that either the Long Ridge or
the Plateau has sufficient peculiarities, quâ ground, to render it
possible that either of them might in ancient times have been known
by a distinctive local name.
 The temple of Eleusinian Demeter is a more important landmark
in the history of the battle.
I venture to think that the ruins of the church of St. Demetrion are
probably on or near the site of that temple, and that the τέμενος of
the temple was on the flat top of that mound of the Long Ridge. The
reasons which may be adduced for this identification are as
follows:⁠—
(a) The Christians, in cases in which they adopted an ancient
temple, or the site of an ancient temple, for a church, were apt, in
dedicating the new sanctuary, to make a pious pun on the dedication
of the pre-existing pagan shrine. This kind of nomenclature has been
peculiarly common in Greece.
(b) It is true that Pausanias mentions several temples of
Eleusinian Demeter as having existed in this immediate
neighbourhood, no one of which can be supposed to have stood in
this position.
But it is also clear that the temple mentioned by Herodotus is not
any one of these mentioned by Pausanias.
Cf. Paus. ix. 4, 2, at Platæa; ix. 4, 3, at Skolos; ix. 9, 1, Grove of
Demeter at Potniæ on the road from Thebes to Platæa, ten stades
from Thebes, i.e. about sixty stades from Platæa.
The omission of Pausanias to mention the one to which I refer
may possibly be accounted for by the fact that it lay, if I am right, not
near the great road from Athens to Thebes, viâ Dryoskephalæ, but
near the alternative and probably much less frequented route
between the two towns by way of the Platæa-Athens pass.
(c) Pausanias (ix. 2, 6) says: “The trophy of the battle of Platæa
which the Greeks set up stood about fifteen stades from the city.”
It would be in accordance with Greek custom that this trophy
should be set up in that part of the field where the decisive
engagement took place. If, as I take it, it was fought just south of the
hill on which the ruined church of St Demetrion stands, a trophy
placed on or near that site would be almost exactly two miles from
the town of Platæa, or a little more than seventeen stades.
 (d) This hypothesis as to the site of the temple receives peculiar
confirmation, as I shall show, from the account given by Herodotus of
the subsequent incidents of the battle, but it will, I think, be best to
take the order of the incidents as given by Herodotus.
In Plutarch’s life of Aristides (xi.) is a topographical detail which
would present considerable difficulty, were it not accompanied by
other details which show the absolute valuelessness of Plutarch as
an authority on the topographical question. He is speaking of the
time at which a move from the first position was contemplated: “Near
Hysiæ Kithæron is a quite ancient temple called (the shrine) of
Eleusinian Demeter and Kore.” He says further on that the rocky
slope of Kithæron meets the plain near the temple, and then adds
that the Heroön of Androkrates was there also, surrounded by a thick
grove of trees.
Pausanias mentions (ix. 2, 1) a temple of Apollo among the ruins
of Hysiæ, but says not a word of any temple of Eleusinian Demeter
being found there.
But the reference to the position of the Heroön of Androkrates
shows that either Plutarch has made a great error, or that his
language is hopelessly inexact.
If latter be assumed to be the case, the temple of Demeter to
which I have referred might, I think, be described as being near what
I take to be the site of Hysiæ. It might even, owing to the height of
Kithæron, be described as underneath that mountain. As to the
position of the temple being near the edge of the ύπωρέη, it may be
pointed out that the site of the modern Kriekouki is a part of the rocky
mountain-side which projects far into the plain, but stops far short of
the church of St. Demetrion.
But even the utmost looseness of language will hardly account for
the assertion that the Heroön of Androkrates was near this temple,
still less that it was near the site of Hysiæ.
The explanation of the difficulty is, I take it, that Plutarch was a
biographer and not an historian; and consequently he may not have
thought, nay, he evidently did not think it necessary to deal intimately
with the topography of the battle. There is very little topographical
detail given by him, though as a Bœotian he had most probably
traversed one or more of the roads which cross the field. He makes
no mention of the νῆσος, either directly or indirectly:⁠—a most strange
omission if he insisted at all on topographical detail.
There is in the passage above referred to a clause which is in
direct conflict with Herodotus’ narrative. Plutarch speaks of τὰς
ὑπωρείας ... ἄφιππα ποιούσας τὰ καταλήγοντα καὶ συγκυροῦντα τοῦ
πεδίου πρὸς τὸ ἱερόν, whereas Herodotus describes a cavalry attack
as having taken place upon this ground. Furthermore,—and this is a
most important point if Herodotus’ account of the fighting be correct,
—the temple of Demeter certainly lay on the Persian line of retreat
after they had been defeated by the Spartans; that is to say, it was
almost certainly between the place where the final action took place
or began and the Persian camp.
Again, I think that the words of Herodotus, in describing the first
phase of the second position of the Greeks as “near the Spring of
Gargaphia and the τέμενος of Androkrates the Hero, through hills of
no great height and level country,” can only mean that the τέμενος
was on the left of the Greek line; for the ἀπέδος χῶρος can only be
the plain between Platæa and the Asopos which comes from
Leuktra, and this position for the τέμενος accords with the details
given by Thucydides. How, then, the τέμενος be described as near
the temple of Demeter, if the latter was near Hysiæ?
On the question of the value of Plutarch’s evidence relative to the
battle, I agree with Colonel Leake:⁠—
“It is scarcely worth while to advert to the particulars in
which the other ancient authors who have related this
great event differ from Herodotus; Diodorus and Plutarch
lived so long afterwards that they cannot have much
weight against the testimony of the contemporary
historian. The former does not, however, deviate from it in
any important point, and the contradictions of the latter are
undeserving of much respect, as being those of a Bœotian
angry with Herodotus for having spoken freely of the
 disgraceful conduct of his countrymen, and thinking no
mode of exculpation so effectual as that of throwing
general discredit on the historian’s accuracy.” (“Travels in
Northern Greece,” vol. ii. p. 353.)

The movement of the Spartans was directed towards the pass on the
Platæa-Athens road. It was evidently their object to avoid as much
as possible the unimpeded ground of the plain on which the Persian
cavalry could act, and with this intent they seem to have diverged
slightly from the direct route to the pass in order to gain the northern
end of that rocky bastion of Kithæron whereon the modern Kriekouki
stands, a point at which the rocky mountain foot projects for some
distance into the plain. They would thus take a
SPARTANS LEAVE
THE SECOND
direction S.S.E. from their position on the
POSITION. Asopos ridge, and must have passed the now
useless Gargaphia spring on their way. After
proceeding thus for ten stades, or about a mile and a furlong, they
waited for Amompharetos and his company, whose reluctant
movement from the late position they would be able to see from the
moment that it began. It must therefore have been near the head
waters of the two streams which run northwards on either side of the
Long Ridge that this delay took place; and either the easternmost of
these streams or the larger brook which skirts the site of Kriekouki
on the west would seem to be the river Moloeis of which Herodotus
speaks. Looking northward from that position, the highest point of
the Long Ridge would rise in front of them in the form of a flat-topped
table-land, bounded on either flank by the steep-sided stream
213
valleys, with the temple of Eleusinian Demeter, of which
Herodotus speaks, upon its summit. The flat table-land itself formed
doubtless that sacred precinct of the goddess of which Herodotus
has something more to tell. Somewhere in the neighbourhood was
the Argiopian country of which the historian speaks, all trace of
whose name and identity has long vanished into the past. Possibly
the plateau to the east of the Long Ridge bore that name in ancient
times.
 It was in the position which they had thus attained that they were
fated to fight the decisive engagement by which the long-drawn
struggle was won. They never reached that rocky projection where
214
Kriekouki now stands.
 H. ix. 57.
“Those with Amompharetos joined them, and the
whole of the barbarian cavalry assailed them,” says
Herodotus, in one significant sentence.
In the gathering light of the dawn the Persians beyond the Asopos
had noticed that the ridges which the Greeks had occupied for more
than a fortnight past were vacant, and had sent forward their cavalry
to overtake the retreating enemy. Amompharetos seems to have just
managed to reach the main body before the attack fell; but the
reunited force had no time to move before the cavalry was upon it.
The ground on which it stood at the time was as unfavourable as it
well could be. The bottom of the ridge lying west of Kriekouki sinks
into the depression in a long gentle slope, affording a terrain
admirably suited to the operations of the Persian cavalry; and it was
on this ground that the Spartans were obliged to make their stand.
The object of the cavalry attack was evidently to prevent the Greeks
from attaining, before the Persian infantry should have time to come
up, the strong position which the rocky slopes of Kithæron would
afford. Mardonius made the fatal mistake of supposing that the
215
Greeks were a broken as well as a beaten army, and his sole
anxiety seems to have been to render the victory complete. Had he
been able to appreciate the real circumstances of the case, had he
been content to abide by the tactics he had employed on the
previous days, the whole history of the battle might have been
changed.

H. ix. 59. Having despatched his cavalry in pursuit of the

Greeks, Mardonius lost no time in hurrying forward his
216
infantry across the Asopos on the track of the retreating foe. It is
interesting to notice that it was against the Lacedæmonians and
Tegeans alone that his efforts were directed, the line of retreat of the
Athenians on the right wing of the Greeks being hidden from him by
217
the hills.
It was with the Persian infantry that Mardonius led the way; but
the rest of the army followed with much speed and little order, eager
to take part in the rout of the enemy with whom they had been so
long face to face. They must have ascended the long gradual slope
of the theatre which is formed by the curve of the Asopos ridge on its
north front, and, arrived at its summit, they would see on the low
ground southward the large mass of the Lacedæmonian contingent
engaged by their cavalry.
Pausanias had no illusions as to the serious
BEGINNING THE
nature of the cavalry attack, but sent an urgent
GREAT FIGHT.
message to the Athenians, who were at this
time making straight for the Island, to come to his help with all
218
speed. The message is recorded, after the manner of Herodotus,
in what purport to be the words actually used. Without unduly
insisting upon the letter, its spirit is remarkable. The Spartans are
represented as having placed the worst construction upon the
conduct of the Greek centre in withdrawing from the position on the
Asopos ridge before they had done so themselves. And yet it will be
remembered that Herodotus expressly says that these “runaways”
did not leave that position until the hour which had been originally
fixed for that movement. But here these allies are represented as
cowards and traitors to their friends. The whole question of the
conduct of the Greek centre at this stage of the battle is an
exceedingly difficult one; the only thing which seems to be clear is
that those Greeks who bore the burden and heat of this day of
decisive battle rightly or wrongly regarded them as having yielded to
a panic of fear. The condemnation was probably unjust; but it is not
possible to demonstrate conclusively that it was so.

H. ix. 61.
The Athenian help never reached the Spartans, and
so they were left to fight the battle alone. It was no
small combat. Over fifty thousand Greeks on the one side were face
to face with what must have been numerically a still more formidable
force—a force which may have been indeed several times their own
in number. The Persian cavalry adhered to the tactics which had
been so effective on the previous days, avoiding close fighting, and
making the assault with arrows and missile weapons from behind a
barrier of shields. Many of the Lacedæmonians fell, and many more
were wounded in this unequal combat, in which they must have been
able to inflict but little damage on their mobile opponents. The
sacrifices were unfavourable; and so the Greeks did not take the
offensive. Pausanias must have been wholly in agreement with the
sacrifices, knowing well, it may be apprehended, that his only
chance of salvation lay in offering a firm front to the attack, and in
preventing the assailant horse from finding its way through the line of
Greek shields. The Persian infantry was, of course, by this time
aiding the assault, and it must indeed have been their shields which
were set up to form the extemporized stockade. This went on for a
long time. Pausanias was probably waiting till the press of the enemy
became sufficiently great to deprive their front ranks of that mobility
from which the Greek hoplite had most to fear. When this state of
things supervened he let his men loose upon the foe, after offering
up a prayer to Hera, which that goddess answered by rendering the
sacrifices favourable. It is probable that the attention which the
Greek commander paid to sacrificial omens was due rather to their
effect on the minds and courage of the common soldiers than to any
undue trust which he placed in them as indications of the tactical
policy to be pursued.

H. ix. 62.
The Tegeans were the first to attack the mass of
barbarians opposed to them, and after an interval the
Lacedæmonians did the same. They first had to break through the
barricade of shields; then the battle became a fierce hand-to-hand
struggle. Herodotus bears testimony to the fact that the Persians
were no whit inferior in courage, but being without heavy defensive
armour, they were no match for the most experienced and best
Cf. ix. 62, ad
drilled infantry in Greece. They evidently made fierce
fin. efforts to break through the iron-clad line of the
enemy, charging singly and in groups. But it was all of
no avail; not even the most desperate courage could compensate for
the immense disparity of weapons, and the men who had “made the
tour of Asia on foot,” and had fought and conquered the numerous
peoples of the East, found themselves utterly unable to cope with an
army drawn from one small corner of the Western Continent.
Mardonius had made a terrible miscalculation when he launched his
comparatively light-armed infantry against the Greek hoplites. He
H. ix. 63.
paid for his mistake with his own life. Surrounded by a
picked body of Persians, he was himself in the midst
 of the mêlée, and was naturally marked out by
FLIGHT OF THE
PERSIANS.
the Greeks for attack. At last he fell, but not
until the assailants had paid dearly for their
success.

PLATÆA: PANORAMA FROM SCENE OF LAST FIGHT.

1. Hills North of Kopais.
2. Mount Kandili (Mekistos) in Eubœa.
3. Hills between Kopais and Euripus.
4. Mount Dirphys in Eubœa.
5. Valley of Stream A 4.
6. Temenos of Demeter.
7. Valley of Stream A 5.
8. The Plateau.
[To face page 502.

It is not difficult at the present day to follow the course of this

219
struggle. After their commander’s fall the Persians gave way, and
were pushed down the gradual slope towards the stream valleys
220
which ran on either side of the precinct of Demeter, affording
them direct lines of retreat to their camp. The stream of fugitives
would divide where the rise of the ground towards the precinct
begins, and would naturally take the easier and quicker routes
H. ix. 65.
afforded by these valleys. They fled in disorder to their
camp and stockade. Herodotus notes with pious
emphasis the fact that, though the fight had raged near the temple,
none of the sacrilegious destroyers of her temple at Eleusis fell
within the consecrated ground about this shrine of Demeter. On
more materialistic grounds it would be extremely unlikely that men
flying for their lives in panic flight should ascend the hill whereon the
sanctuary and its precinct stood, when easier avenues of retreat
221
were ready for them on either side of it.
Such were the fortunes of what had been the Greek right wing. It
is now necessary to follow the movements of the Athenians who had
been stationed on the left of the second position.
It is clear that the two days’ cavalry attack had driven back the
Greek left from the Asopos; and, in the final phase of its tenure of the
second position, the army must have been massed on the summit of
the Asopos ridge; so that the left would be at no great distance from
the right.

H. ix. 54. The Lacedæmonian delay in starting upon the

movement of retirement seems to have aroused
suspicion in the minds of the Athenians, if Herodotus’ narrative is to
be believed. He does not mention any specific cause for the distrust
in the present instance, other than the delay caused by the obstinacy
of Amompharetos, attributing it rather to general grounds: “They
knew the Lacedæmonians had a way of intending one thing and
saying another.” The conduct of the Athenians is not consistent with
the tale of suspicion; and the tale itself probably does something less
than justice to the Athenian motives for remaining at their post of
danger. It is noticeable that when they do move they take a wholly
different line to the Spartans, showing that they did not suspect them
of any intention of deserting the army, and consequently the only
conceivable motive they can have had in waiting must have been the
desire that the withdrawal should be covered, if necessary, by a
sufficient number of the Greek army. As, however, the Spartans
started just before daylight, and the Persians did not immediately
discover the movement, the Athenians evidently thought that there
would be no danger in carrying out the original design; so, while their
allies made their way towards the passes, they took the easiest route
to the Island.
They apparently started on their movement about the same time
as the Lacedæmonians. Their course can be traced with fair
certainty. From their position on the Asopos ridge north of the ruins
of the church of St John, they did not take the direct line for the
Island, which would have led them past the site of that modern
sanctuary, but descended to the western slope of the ridge to the
222
Platæan plain.
On reaching it they would pass up the course of that stream which
223
Herodotus identifies with the upper Asopos, and would be
marching in a straight line for the Island. The
ATHENIANS AND
route adopted was indeed but little longer than
MEDIZED GREEKS.
the direct route, and would be easier, as it lay
H ix. 60.
along the level plain. It must have been at some point
in the course of this march that the message from
Pausanias reached them, announcing the attack of the Persian
cavalry, and urgently entreating them to come to his assistance, or at
H. ix. 61.
any rate to send their bowmen to his help. On
receiving this the Athenians diverged from the line of
march, and started towards the place where the Spartans were
being sore pressed by the cavalry. Shortly after this change of
direction had taken place they were attacked by the medized Greeks
who were with the Persian force, and were thus prevented from
assisting the Spartans. The circumstance that this fight seems to
have been out of sight of that part of the Greek army which was at
Platæa, renders it probable that the Athenians had turned from the
plain along the line of the modern track which leads from Leuktra to
Dryoskephalæ, and that the medized Greeks, coming up over the
Asopos ridge in the neighbourhood of the church of St. John,
overtook them in the hollow near the springs of Apotripi, where a
H. ix. 67.
fierce engagement took place. Although the rest of the
medized Greeks voluntarily played the coward on this
occasion, the Bœotians showed that they had not espoused the
Persian cause to no purpose, and for a long time maintained an
obstinate struggle, losing three hundred of their best and bravest in
the battle. In the end they had to give way, and fled in disorder to
Thebes.

H. ix. 68.
The whole Persian army was now in full flight, save
the cavalry, which made a brave and, for the time
being, a successful effort to cover the retreat, though it could not
prevent the pursuing Greeks from inflicting severe losses upon the
fugitives.

H. ix. 69.
It was at this time, says Herodotus, that the Greeks
at Platæa first heard that a battle had taken place, and
that Pausanias and his men had won the day. They must have spent
an anxious morning on that rocky slope near the Heræon, not
venturing to move, because uncertain what they ought to do, and
knowing nothing of their friends save that they were not at the Island,
where, according to arrangement, they should have been. They may
perhaps have seen the Athenians moving along the foot of the
Asopos ridge shortly after dawn, and the Persian army as, a little
later, it streamed across that ridge, must have been plainly visible;
but from the moment when the Athenians disappeared into the
hollow near the springs of Apotripi, until they received this message,
all they can have seen or heard must have been the clouds of dust
rising from two points in the depression beneath the northern ridges,
and the confused and distant noise of an engagement. They seem to
have played an inglorious, if not a cowardly, part in sitting still while
these things were going on; but if it be borne in mind that the little
they did see of the movements of their friends and of the enemy on
the fatal morning was wholly inexplicable by the orders they
themselves had received, or by anything that had happened within
their knowledge, their inaction may seem natural. Herodotus is not
inclined to put the best construction on the conduct of those who
were drawn largely from those Greek states which had advocated
what seemed to him the cowardly blunder of confining the defence to
the Isthmus.
The receipt of the message, therefore, put them for the first time
in possession of certain information as to what had happened since
they had parted from their allies on the Asopos ridge on the