Anatomy and Physiology Notes

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Anatomy and Physiology Notes

Human Anatomy and Physiology Notes
Lecture 1- Introduction, Structure, Function and Cell Theory
Overview
● When referring to anatomy/Physiology of the body, we assume that we are talking about a
healthy 22-year old male weighing 155 lbs or a health young female weighing 125 lbs
Topics of Anatomy
● Is the study of the structure of body parts
● Must memorize
● Major subdivisions of anatomy are:
○ Gross anatomy- the study of large body structures
■ Regional anatomy- study of structures in particular region
■ Systematic anatomy- body structure is studied system by system
■ Surface anatomy- study of internal structures as they relate to the overlying skin
surface
○ Microscopic anatomy- study of structures that can’t be seen with naked eye
■ Cytology- considers cells of body
■ Histology- the study of tissues
○ Developmental anatomy- traces structural changes that occur in body throughout
lifespan
■ Embryology- study of developmental changes that occur before birth
Topics of Physiology
● Physiology is the study of the functions of the body
● Must understand
● Subdivisions concern the individual specific organ systems
○ Eg; neurophysiology, cardiovascular physiology, etc.
● Often focuses on events at a cellular/molecular level
Complementarity of Structure and Function

● Anatomy and physiology is inseparable as function always reflects structure
● This key concept is called the principle of complementarity of structure and function
● You can infer function from structure
Scientific Reductionism
● Reductionism- an approach to understanding the nature of complex things by reducing them to
simpler things
● Physiology often focuses on events at the cellular or molecular level because the body’s
abilities depend on individual cells
Levels of Structural Organization

● Body’s organization ranges from atoms (smallest unit) to the entire organism (largest unit)
● Levels of structural organization (simplest to most complex): chemical, cellular, tissue, organ,
organ system and organismal
○ Chemical level- atoms combine to form molecules
○ Cellular level- Cells are made up of molecules
○ Tissue level- Groups of similar cells with a common function
■ 4 types- epithelial, muscle, connective, and nervous
○ Organ level- Structure composed of two (usually 4) tissue types. Specialized functional
structure for specific activity
○ Organ System level- organs work together to accomplish a common purpose
○ Organismal level- An organism is made up of numerous organ systems
Organ Systems of the Body

Organ System Function
Integumentary ● Forms external body covering + protects deep tissues from

System injury
● Synthesizes vitamin D and houses external stimuli receptors and
sweat and oil glands
● Hair, nails, skin
Skeletal System ● Protects + supports organs

● Provides framework that muscles use for movement
● Blood cells synthesized in bones and bones store minerals
Muscular System ● Allows manipulation of the environment, locomotion and facial

expression
● Maintains posture + produces heat
Nervous System ● Fast- acting control system of body

● Responds to external and internal stimuli by activating
appropriate muscles and glands
Endocrine System ● Glands secrete hormones which regulate numerous body

processes
Cardiovascular ● Blood vessels transport blood and other nutrients to the body
System ● Heart pumps the blood
Lymphatic System ● Picks up fluid leaked from blood vessels and returns it to blood
● Disposes of debris in the lymphatic stream
● Houses white blood cells (lymphocytes) involved in immunity
Respiratory ● Keeps blood supplied with oxygen + removed carbon dioxide

System ● Gas exchange occurs through walls of air sacs in lungs
Digestive System ● Breaks down food to be absorbed by blood

● Indigestible foods eliminated as feces
Urinary System ● Eliminates nitrogenous wastes from body

● Regulates water, electrolyte and acid-base balance of the blood
Reproductive ● Male and female

System ● Used for production of offspring
Parts of a Cell: Structure and Function

● Refer to the following lecture 1 powerpoint for condensed chart
● https://uottawa.brightspace.com/d2l/le/content/117592/viewContent/2387015/View
Organelles In The Cytoplasm

● Is the region between nuclear and plasma membranes
● Consists of cytosol, organelles and inclusions (stored nutrients, secretory products, pigment
granules)
Organelle Structure Function
Mitochondria ● Rodlike, double-membrane ● Site of ATP synthesis

structure ● Powerhouse of the cell
● Outer membrane is smooth ● Has own DNA and replicates
● Inner membrane folded into independently
projections called cristae
● Gel-like matrix in the center
Ribosomes ● Consist of 2 subunits, each ● Synthesize proteins for various cell

composed of ribosomal RNA and needs
protein ● Can switch back and forth between free
● Free or attached to rough ER floating and bounded states
Rough ER ● Outer surface is covered in ● External face synthesizes proteins

ribosomes ● Integral proteins and phospholipids are
● Membranous also manufactured
● Coils through the cytoplasm ● Proteins are bound in vesicles and
transported to the Golgi apparatus and
other sites
Smooth ER ● Is continuous with rough ER ● Does not make proteins

● Membranous system of sacs and ● Contains enzymes that are responsible
tubules for synthesizing lipids and steroids,
● No ribosomes lipid metabolism and drug
detoxification
Golgi ● Stack of flattened membranes and ● Packages, modifies and segregates

Apparatus vesicles proteins and lipids made in the rough
● Located near the nucleus ER for secretion from the cell, inclusion
in lysosome and incorporation in plasma
membrane
Peroxisomes ● Membranous sacs of catalase and ● Oxidase enzyme detoxifies many toxic
oxidase enzymes substances and neutralizes free radicals
● Catalase (important enzyme) breaks
down hydrogen peroxide
Lysosomes ● Membranous sacs containing acid ● Sites of intracellular digestion

hydrolases ● Work best in acidic conditions
● Spherical
Microtubules ● Cylindrical ● Support cell and gives it shape

● Composed of tubulin proteins ● Involved in intracellular and cellular
movements
● Form centrioles, cilia and flagella
Microfilament ● Fine filaments composed of ● Involved in muscle contraction and

s protein actin other intracellular movements
● Helps form cell’s cytoskeleton
Intermediate ● Protein fibers ● Make up the stable cytoskeleton

Filaments ● Composition varies ● Resist mechanical forces acting on cell
Centrioles ● Paired cylindrical bodies ● Organize microtubule network during

● Each composed of nine triplets of mitosis to form the spindle and asters
microtubules ● Form the bases of cilia and flagella
Inclusions ● Varied ● Are storage for nutrients, wastes and

cell products
Cellular Extensions
Part Structure Function
Cilia ● Short, cell surface projections ● Their coordinated movement

● Composed of nine pairs of creates a current that propels
microtubules surrounding a substances across cell surfaces
central pair
Flagellum ● Like cilium, but longer ● Propels the cell in its

● Only present in sperm cells in environment
humans
Microvilli ● Tubular extensions of plasma ● Increase surface area for

membrane absorption
● Contain bundle of actin fibers
Parts Of The Nucleus

Part Structure Function
Nucleus ● Largest organelle ● Control center of cell

● Surrounded by nuclear envelope ● Transmits genetic information
● Contains fluid nucleoplasm, nucleoli ● Provides instructions for protein
and chromatin synthesis
Nuclear ● Double membrane ● Separates nucleoplasm from

Envelope ● Porous cytoplasm
● Outer membrane is continuous with ER ● Regulates passage of substances to
and from the nucleus
Nucleolus ● Dense, spherical bodies ● Site of ribosome subunit synthesis

● Non-membrane bound
● Composed of RNA and proteins
Chromatin ● Granular and threadlike ● Fundamental units are nucleosomes

● Composed of 30% DNA, 60% histone ● Packs DNA into a small volume to
proteins and 10% RNA chains fit the nucleus
● Protects DNA structure and sequence
Maintaining life
● Necessary Life Functions
○ All organisms carry out specific vital functional activities necessary for life
○ Include maintenance of boundaries, movement, digestion, responsiveness, metabolism,
excretion, reproduction and growth
● Survival Needs -Requirements for maintaining life
○ Nutrients- taken through diet. Contain chemical substances required for energy and cell
building
○ Oxygen- chemical reactions that release energy from foods are oxidative reactions that
require oxygen
○ Water- accounts for 60-80% of body weight. Provides necessary environment for
chemical reactions
○ Body Temperature- Normal body temperature is required for chemical reactions to
occur. Muscular system activity generates the most heat.
○ Atmospheric Pressure- breathing and gas exchange are dependent on appropriate
atmospheric pressure.
Lecture 2- Types of Tissues
Processes Needed to Make Us Multicellular

● Cell division
● Work with environment
● Movement
● Cell differentiation
● Cell-Cell interactions
○ Direct interactions, endocrine and nervous system signal interactions
Tissues
● Are groups of cells that are similar in structure and perform a common function
● Are four basic types of tissues in the body
○ Epithelial, connective, nervous and muscle tissue
● In short, epithelial tissue covers, connective tissue supports, nervous tissue controls and muscle
tissue provides movement
Overview of 4 Basic Tissues

Tissue Function Location
Nervous Tissue Internal communication Brain, spinal cord, nerves
Muscle Tissue Contracts to cause movement Skeletal muscles

Cardiac muscles
Muscles of the walls of hollow organs
(smooth muscles)
Epithelial Forms boundaries, protects, secrets, Lining of digestive track and hollow
Tissue absorbs, filters organs
Glands
Skin Surface
Connective Supports, protects, binds other tissues Bones, tendons, fat and other soft
Tissue together padding tissue
Preparing Human Tissues for Microscopy

● Histology- the study of tissues
● Microscopy allows for the study of tissue structure
● Before viewing, the specimen must undergo the following steps in order;
○ It must be fixed (preserved)
○ Then, it must be cut into sections (slices) thin enough to transmit light/electrons
○ Finally, it must be stained to enhance contrast (light microscopy)
● Transmission electron microscopy (TEM)
○ Tissue sections are stained with heavy metal salts that deflect electrons in the beam to
different extents, providing contrast
● Scanning electron microscopy (SEM)
○ Provided 3-D pictures of an unsectioned tissue surface
Type 1- Epithelial Tissue

● Is a sheet of cells that covers a body surface or lines a body cavity
● Two forms occur in the body
○ Covering and lining epithelium
■ Forms outer layer of the skin
■ Lines the open cavities of the urogenital, digestive and respiratory systems
■ Covers the walls and organs of the closed ventral body cavity
○ Glandular epithelium
■ Covers the glands of the body
■ Secretory tissue in glands (eg; salivary glands)
● All substances received or given off by the body have to pass through the epithelium
○ Selective barrier- intestinal epithelium allows passage of certain substances
■ Bidirectional, highly regulated in healthy and not healthy tissue
■ Gut microbiome
● Accomplishes many functions including;
○ Protection, absorption, secretion, filtration, excretion, sensory reception
Special Characteristics of Epithelium

● Has five distinguishing characteristics
○ Polarity, specialized contacts, supported by connective tissue, avascular but innervated,
ability to regenerate
● Polarity
○ Apical surface (top) is exposed to surface or cavity
■ Most are smooth, but some have microvilli (increases surface area) or cilia
○ Basal surface (bottom), faces inwards towards body
■ Attached to basal lamina, an adhesive sheet that holds the basal surface of
epithelial cells to underlying cells
● Acts as a selective filter that determines which molecules diffusing from
underlying tissue are allowed in
○ Cell polarity is crucial for normal cell physiology and tissue homeostasis
● Specialized Intercellular Contacts
○ Cells need to fit closely together to form barrier
○ Specialized contact points bind adjacent epithelial cells together (between cells)
○ Lateral contacts include;
■ Tight junctions- prevents substances from leaking between cells (regulated)
■ Adherens junctions- mediate cell-cell adhesion via protein cadherins
■ Desmosomes- prevents cells from pulling apart in tissues subjected to

mechanical stress (made up of protein desmoglein)
■ Gap junctions- mediated by connexions. Intercellular communication.
○ Use different connecting proteins and attach to different filaments
● Supported by Connective Tissue
○ All epithelia have a basement membrane (BM)
■ reinforces , resists stretching, tearing
■ Defines epithelial boundary
■ Consists of
● Basal lamina- layer of extracellular matrix (ECM) proteins
● Reticular lamina- deep to the basal lamina. Network of collagen III
fibers
○ Disease Relevance- cancerous epithelial cells can penetrate BM and invade underlying
tissues, resulting in spread of cancer (metastasis)---> 90% of cancer deaths
● Avascular, But Innervated
○ Avascular- no blood vessels
■ Nutrients and oxygen diffuse from connective tissue
○ Innerved- have nerves
● Regeneration
○ Epithelials cells have high regenerative capacities
○ Simulated by loss of apical-basal polarity and broken lateral contacts
○ Some cells exposed to friction, hostile substances ---- > damage
■ Must be replaced
■ Requires adequate nutrients and cell division
■ Wound healing
Classification of Epithelia
● Based on two factors
● Number of cell layers (1 or more)
○ Simple epithelia- a single layer thick
○ Stratified epithelia- are two or more layers thick
■ Protection is a major role
■ New cells regenerate from below
● Basal cells divide and migrate to surface
■ More durable than simple epithelia
● Shape of cells
○ Squamous: flattened and scale-like (humans have a lot of them)
■ Most of the cells in the outer layer of skin
■ Passages of respiratory and digestive tracts
■ Lining of hollow organs
○ Cuboidal: box-like, cube
○ Columnar: tall, column-like

■ In stratified epithelia, shape can vary in each layer, so classified according to
shape in apical layer
Types of Simple Epithelium

Type and Image Description Function Location
Simple Squamous -Single layer of flattened cells. -Allows materials to pass via -Kidney glomeruli
-Disk shaped nuclei diffusion and filtration in sites -Air sacs of lungs
-Sparse cytoplasm where protection is not key, but -Lining of heart
-Simplest of epithelia rapid diffusion is. -blood vessels
-Two types based on location -Secretes lubricating substances -lymphatic vessels
*(see bottom for note)* in serosae (lining of ventral body -Serosae
cavity)
Simple Cuboidal -Single layer of cube like cells -Secretion and absorption -Kidney tubules
-spherical central nuclei --Ducts and
secretory portions of
small glands
-Ovary surface
Simple Columnar -Single layer of tall cells with -Absorption -Non-ciliated line
round to oval nuclei -Secretion of mucous, enzymes most of digestive
-Many have microvilli and other substances tract (stomach to
-Some have cilia -Ciliated type propelles mucous rectum), gallbladder,
-Layer may contain (or reproductive cells) and excretory ducts
mucus-secreting unicellular of some glands
glands (goblet cells) -Ciliated type lines
small bronchi,
uterine tubes, and
some parts of uterus
Simple -Single layer of cells differing -Secrete substances, mainly -Ciliated type lines
Pseudostratified in height mucous trachea and most of
Columnar -Nuclei seen at different levels -Propulsion of mucus by ciliary upper respiratory
-May contain mucus-secreting action tract
cells and have cilia -Nonciliated type in
male sperm-carrying
ducts and ducts of
large glands
*There are two types of simple squamous epithelium based on location;

● Endothelium
○ Lines inner surface of blood vessels, lymphatic vessels, and the heart
○ Derived from ectoderm and endoderm in the early embryo
● Mesothelium
○ Form serous membranes- surround pericardium, peritoneum, pleura, and internal
reproductive organs (covers outer surface)
○ Derived from mesoderm
○ Two membrane system with fluid in between
Types of Stratified Epithelium

Type/Image Description Function Location
Stratified Squamous -Thick, composed of several layers -Protects -Located in areas of high
-Most widespread underlying tissue wear and tear
-Basal cells are cuboidal or columnar and in areas subject to -Keratinized cells found
metabolically active abrasion in skin and
-surface cells are flattened (squamous) nonkeratinized found in
-Keratinized type- surface cells are dead moist linings of mouth,
and full of keratin. Basal cells are active esophagus and vagina
in mitosis and produce superficial layer
cells
Stratified Cuboidal -Very rare -Found in some sweat

-Typically only two layers thick and mammary glands
Stratified Columnar -Very rare in the body -Transitional areas

-only apical layer is columnar between two other types
of epithelia
-Small amounts found in
pharynx, male urethra,
and lining of some
glandular ducts
Transitional -Resembles both stratified squamous and -Stretches readily -Lines the ureters,
stratified cuboidal -Permits stored bladder and part of the
-Basal cells are cuboidal or columnar urine to distend urethra
-Surface cells dome-shaped or squamous urinary organ
like depending on the degree of organ
stretch
Glandular Epithelia
● Gland-one or more cells that make and secrete an aqueous fluid called a secretion
● Classified by two factors
○ Site of product release
■ Endocrine- internally secreting
■ Exocrine- externally secreting
○ The relative number of cells forming the gland
■ Unicellular (eg. goblet cells) or multicellular (eg. salivary)
Formation of Multicellular Exocrine & Endocrine Glands

● Multicellular epithelial glands form by invagination (inward growth) of an epithelial sheet
● Exocrine glands retain the connecting cells, which form a duct that transports secretions to the
epithelial surface
● Endocrine glands lose their ducts during development
○ Secrete hormones into the interstitial fluid which ten enter the blood
Formation of Multicellular Endocrine & Exocrine Glands

● Multicellular epithelial glands form by invignation (inward growth) of an epithelial sheet
○ Exocrine glands retain connecting cells which form a duct that transports secretions into
the epithelial surface
○ Endocrine glands lose their ducts during development.
■ They secrete hormones into interstitial fluid which then enter the bloodstream
Endocrine Glands
● Ductless
● Secretions are released into surrounding interstitial fluid which is then infused into the
bloodstream
● Hormones- chemicals that travel through the lymph or blood to target organs that respond in a
characteristic way
● Major glands of the endocrine system include;
○ Pineal gland, pituitary gland, pancreas, ovaries/testes, thyroid and parathyroid glands,
hypothalamus, adrenal glands
Exocrine Glands
● Secretions are released onto body surfaces (eg skin) or into body cavities
● More numerous than endocrine glands
● Secrete products into ducts
● Can be multicellular or unicellular
● Examples include mucus, sweat, oil, etc
● Unicellular Exocrine Glands
○ Only important ones are mucus cells and goblet cells
○ Found in epithelial linings of intestinal and respiratory tracts
○ All produce mucin, a sugar protein that can dissolve in water to form mucus which is a
slimy, protective coating
● Multicellular Exocrine Glands
○ Composed of duct and secretory unit
○ Usually surrounded by supportive connective tissue that supplies blood and innervation
■ Connective tissue can form capsule around gland or extend into gland, dividing
it into lobes
○ Classified by structure and modes of secretion as shown in the image below
○ Main modes of secretion
■ Merocrine glands- secrete products via exocytosis
■ Holocrine glands- secretory cell ruptures, releasing secretions and dead cell
fragments
■ Apocrine- accumulate products within, but only apex rupture (not sure if this
type occurs in humans)
Type 2- Connective Tissue

● Is the most abundant and widely distributed or primary tissues
● Major functions: binding and support, protection, insulation, storing reserve fuel and
transporting substances
● 4 main classes;
○ Connective tissue proper, cartilage, bone, blood
Overview of Classes of Connective Tissue

(refer to ppt and textbook for chart)
Common Characteristics of Connective Tissue

● Three characteristics differentiate connective tissue from other types of primary tissue
○ Have common embryonic origin: mesenchyme
○ Have varying degrees of vascularization
■ Eg; cartilage is avascular (lack of blood vessels) and bone is highly vascular
(consists of vessels)
○ Cells are suspended/embedded in extracellular matrix (ECM)
■ ECM supports cells so that they can bear weight, tension and abuse
Structural Elements of Connective Tissue

● All connective tissues have 3 main elements
○ Ground substance, fiber and cells
● Ground substance and fiber together make up the ECM
● Composition and arrangement of these three elements vary in different types of connective
tissues
● Ground Substance
○ Gel-like material that fills space between cells
■ Is the medium through which solutes diffuse between blood capillaries and cells
○ Components
■ Interstitial fluid
■ Cell adhesion proteins
■ Proteoglycans (sugar proteins) composed of protein core + large polysaccharides
■ Water trapped in varying amounts, affecting the viscosity of ground substance
● Fibers
○ 3 types of fiber provide support
■ Collagen
● Strongest, most abundant type
● Tough; provides high tensile strength
■ Elastic Fibers
● Networks of long, thin elastin fibers
● Allow for stretch and recoil
■ Reticular
● Short, fine, highly branched collagenous fibers
● Branching forms networks that offer more ‘give’
● Cells
○ “-blast” cells
■ Immature form of cell that secretes ground substance and ECM fibers
■ Fibroblasts- found in connective tissue proper
■ Chondroblasts- found in cartilage
■ Osteoblasts- found in bone
■ Hematopoietic- stem cells in bone marrow

○ “-cyte” cells
■ Mature, less active form of ‘-blast’ cell- is part of, and helps maintain the health
of the ECM
○ Other cell types
■ Fat cells- store nutrients
■ White blood cells- tissue respond to injury (neutrophil, lymphocyte)
■ Mast cells- initiate local inflammatory response against foreign microorganisms
they detect (1st line of defence)
■ Macrophages - phagocytic cells that eat dead cells; microorganisms that
function in the immune system
● Refer to ‘Areolar connective tissue: A prototype’ slide in ppt for image of all components
Types of Connective Tissue Proper

Type/Image Description Function Location
Loose Areolar -Most widely distributed -Universal packing material -Widely distributed
connective tissue between other tissues under epithelia of the
-Gel-like matrix containing -Wraps and cushions organs body
all 3 fiber types and some -Plays an important role in -Eg: forms lamina
white blood cells inflammation propria of mucous
-Macrophages and fat cells are membranes, surrounds
contained in spaces organs and capillaries
-Loose fibers allow for increased
ground substance, which can act as
water reservoir
Loose Adipose -Includes white (reserve) and -Shock absorption -Under the skin in
brown fat (creates heat) -Insulation subcutaneous tissue
-Similar to areolar tissue, but -Supports and protects organs -Around kidneys and
greater nutrient storage -Provides reserve energy storage eyeballs
-Cells are called adipocytes -Within abdomen
-Sparse matrix -In breasts
-Richly vascularized
-Nucleus pushed to side by
large fat droplets
Loose Reticular -Loose network of reticular -Fibers form internal skeleton -Lymphoid organs
fibers in ground substance (stroma) that supports other cell (lymph nodes, bone
-Reticular (fibroblast) cells types including blood cells, mast marrow, spleen)
lie on the fibers cells and macrophages
Dense Regular -Primarily thick, parallel -Very high tensile strength from -Tendons
collagen fibers that are one direction -Ligaments
slightly wavy -Attaches muscles to bones -Aponeuroses (flat
-Major cell type is fibroblast tendon)
-Few elastin fibers
-Poorly vascularized
Dense Irregular -Primarily irregularly -Withstands tension from many -Dermis of skin
arranged collagen fibers directions -Fibrous capsules of
-Forms sheets rather than -Provides structural strength organs or joints
bundles -Submucosa of
-Some elastin fibers digestive tract
-Fibroblast is the main cell
type
Dense Elastic -Dense regular connective -Allows tissues to recoil after -Walls of large
tissue stretching arteries
-Contains a high proportion -Maintains pulsilate flow of blood -Certain ligaments
of elastin fibers through arteries associated with
-Aids passive recoil of lungs vertebral column
following inspiration -Walls of bronchial
tubes
Cartilage
● Tough, yet flexible material that lacks nerve fibers
● Matrix secreted from chondroblasts (during growth) and chondrocytes (when adult)
○ Chondrocytes found in cavities called lacunae
○ 80% water, packed with collagen fibers and sugar proteins
● Is avascular
○ Receives nutrients and support from surrounding layer of dense, irregular CT
■ Perichondrium provides to chondroblasts and chondrocytes
Types of Connective Tissue Cartilage

Tissue/Image Description Function Location
Cartilage Hyaline -Most abundant -Supports and reinforces -Forms most of

-Amorphous (lacking structure), but -Serves as resilient cushion embryonic skeleton
firm matrix -Resists compressive stress -Covers the ends of long
-Appears as shiny bluish glass bones in joint cavities
-Collagen fibers form imperceptible -Forms costal cartilages
network of the ribs, nose, trachea
-Chondrocytes lie in lacuna and larynx
-Surrounded by perichondrium
Cartilage Elastic -Similar to hyaline, but more elastic -Maintains the shape of a -Supports the external
fibers in the matrix structure while allowing ear
great flexibility -Epiglottis
Fibrocartilage -Matrix similar to, but less firm than -Tensile strength allows it to -Intervertebral discs
that in hyaline absorb compressive shock -Pubic symphysis
-Thick collagen fibers predominate -Discs of knee joints
-Strong
Clinical Relevance of Cartilage

● Avascular cartilage loses ability to divide with age, so injuries heal slowly
○ Common in individuals with sports injuries
● Later in life, cartilage can calcify or ossify (become bony), causing chondrocytes to die
Bone
Type Description Function Location
Bone -2 types of bone: compact bone and -Supports and protects -Bones
spongy bone -Provides levers for the
-Hard, calcified matrix containing muscles to act on
many collagen fibers - Stores calcium, other
-Osteocytes lie in lacunae minerals and fat
-Very well vascularized -Marrow inside bones is
-Calcium cannot be synthesized by site of blood cell formation
the body. Bones are the source of (hematopoiesis)
Ca -Bones supply extra
calcium when needed
Blood
Type Description Function Location
Blood -Red and white blood -Transports respiratory -Contained within

cells in a fluid matrix gasses, nutrients, blood vessels
(plasma) water, wastes and other
-Platelets substances
-Derived from
mesoderm
Lecture 3- Plasma Membrane, Diffusion, Transport and Osmosis
Chapter 3 Readings (Refer to ppt)
Plasma Membrane
● Aka the cell membrane
● Acts as a barrier separating the intracellular fluid from the extracellular fluid
● Is a selective barrier- plays a role in cell activity by controlling what enters and leaves the cell
● Consists of membrane lipids that form a flexible lipid bilayer
Plasma Membrane Lipids

● The plasma membrane consists of;
○ 75% phospholipids which consist of 2 parts;
■ Phosphate heads- are polar and hydrophilic
■ Fatty acid tails- are nonpolar and hydrophobic
○ 5% glycolipids
■ Lipids with sugar groups attached to outer membrane surface
○ 20% cholesterol
■ Increases membrane stability
■ Component of lipid rafts
Phospholipids
● Are amphipathic- contain both a hydrophilic and hydrophobic region
● In the digestive system, phospholipid containing bile salts help break up ingested lipids using
amphipathic properties
Plasma Membrane Proteins

● Specialized membrane proteins float through the fluid plasma membrane, resulting in constantly
changing patterns
○ Referred to as fluid mosaic
○ Surface sugars form glycocalyx
● Membrane structures help hold cells together through cell junctions
Functions Of The Plasma Membrane
Function Description
Physical Barrier -Encloses the cell

-Separates ICF from ECF
Selective Permeability -Determines which substances enter or exit the cell
Communication -Plasma membrane proteins interact with specific chemical

protein messengers and relay messages to the cell interior
outside-in and inside-out
Cell Recognition -Cell surface carbohydrates allow cells to recognize each

other
Building Blocks of the Plasma Membrane

Name Description
Phospholipids -Form the basic structure of the membrane

-Hydrophobic tales prevent water-soluble substances from
crossing
-Form a boundary
Cholesterol -Stiffens membrane

-Further decreases water solubility of the membrane
-Typical 4 ring steroid structure
Proteins -Determines what functions the membrane can perform

-Many roles in communication, transport, signalling, joining
cells, etc.
-Proteins with different shapes have different functions
Carbohydrates -Act as identity molecules

-Allow for cell recognition during development so
cells can sort themselves out into organs and tissues
-Allow immune cells to recognize good and bad substances
-Found only on the outer surface of the membrane
-Together, all carbohydrates on outside of cell form a coating
called the glycocalyx
Membrane Proteins
● There are 2 types;
○ Integral membrane proteins
○ Peripheral membrane proteins
● Allow communication (signal transduction) with environment (bi-directional)
● Have specialized functions
● Some float freely in the plane of the membrane while others are immobilized into cellular
structures
● How do we know a protein is free to move in the plasma membrane?
Fluid Mosaic Model

● Fluorescence recovery after photobleaching (FRAP) was used to help refine the fluid mosaic
model
Modification of Membrane Fluidity

● Membrane fluidity can be modified by a number of factors
● Cholesterol (up to 20%) and glycolipids (5%) and other specialized lipids can modify
membrane fluidity
● Form lipid rafts- undergo endocytosis (refer to tb)
● Level of phospholipid saturation also have an effect
Classes of Membrane Proteins

● Integral Membrane proteins
○ Firmly inserted into membrane
○ Most are transmembrane proteins
○ Have both hydrophobic and hydrophilic regions
■ Hydrophobic areas interact with lipid tails
■ Hydrophilic areas interact with water
○ Function as transport proteins (channels and carriers), enzymes or receptors
● Peripheral Membrane Proteins
○ Loosely attached to integral proteins or anchored to the membrane with covalently
attached lipid groups
○ Can be on inner or outer part of the PM
○ Include filaments on intercellular surface which used for plasma membrane support
○ Function as;
■ Enzymes
■ Signal transduction (DEFINE)
■ Scaffold proteins (DEFINE)
Functions of Membrane Proteins

Function Description
Transport -A protien that spans the membrane may provide a hydrophilic

channel across the membrane that is selective for a particular
solute
-Some transport proteins hydrolyze ATP as an energy source to

actively pump substances across the membrane
Receptors for Signal -A membrane protein exposed to the outside of the cell may have a
Transduction binding site that fits the shape of a specific chemical messenger,
suh as a hormone
-When bound, the chemical messenger may cause a change in
shape in the protein that initiates a chain of chemical reactions in
the cell
Enzymatic Activity -A membrane protein may be an enzyme with its active site
exposed to substances in the adjacent solution
-A group of several enzymes in a membrane may catalyze
sequential steps of a metabolic pathway
Cel-Cell Recognition -Some glycoproteins serve as identification tags that are

specifically recognized by other cells
-Important for cell-cell communication
Attachment to -Elements of the cytoskeleton and the ECM may anchor to

Cytoskeleton and ECM membrane proteins
-Helps maintain cell shape, fixes the location of certain membrane
proteins and plays a role in cell movement
Cell-Cell Adhesion -membrane proteins of adjacent cells may be hooked together in

various kinds of intercellular junctions
-Some membrane proteins (cell adhesion molecules or CAMs) of
this group provide temporary binding sites that guide cell
migration and other cell-cell interactions
Glycolax
● Consists of sugars (carbohydrates) sticking out of the cell surface
○ Some sugars are attached to lipids (glycolipids) and some to proteins (glycoproteins)
● Every cell has different patterns of this ‘sugar coating’
○ Functions as specific biological markers for cell to cell recognition
○ Allows immune system to recognize self vs. nonself
Cell Junctions
● Are specialized areas of the plasma membrane
● There are three types
○ Gap Junctions
■ Communicating junctions
■ Allow ions and small molecules to pass from cell to cell
■ Particularly important in heart cells and embryonic cells
■ Eg: Channels formed by connexons
■ Found in epithelial cells and some nerve cells
○ Desosomes
■ Anchoring junctions
■ Bind adjacent cells together like molecular velcro
■ Help keep cells from tearing apart
■ Eg: Linker proteins (cadherins)
○ Tight junctions
■ Impermeable junctions
■ Form continuous seals around the cell
■ Prevent molecules from passing between cells
■ Eg: Interlocking junction proteins
Gap Junctions
● Transmembrane proteins (connexons) form tunnels or gaps between cells that allow small
molecules to pass from cell to cell
● Used to spread ions, simple sugars or other small molecules between cells
● Allows electrical signals to be passed quickly from cell to cell
○ Mostly in cardiac and smooth muscle cells, but sometimes in neurons
Transport of Substances Across the PM

● Many substances constantly move across the plasma membrane
○ Some molecules pass through easily and some do not
● The plasma membrane is selectively permeable, allowing only certain molecules to pass
through
● Two essential ways substances can cross the PM;
○ Active Transport: energy (ATP) is required
○ Passive Transport: No energy is required
Passive Membrane Transport

● Requires no energy input
● Three types of passive diffusion:
○ Simple diffusion
○ Facilitated diffusion
○ Osmosis
● All types involve diffusion- the natural movement of molecules from areas of high
concentration to areas of low concentration
○ Also referred to as moving down a concentration gradient
How Does Diffusion Work?

● All molecules have random, high-speed movement due to their intrinsic kinetic energy
● Movement results in collisions between molecules
● Molecules in higher concentration areas collide more, resulting in molecules being scattered to
lower concentration areas- called diffusion
Factors That Affect Diffusion

● The speed of diffusion is affected by 3 factors
○ Concentration
■ The greater the difference of concentration between the two areas, the faster the
diffusion occurs
○ Molecular Size
■ Smaller molecules diffuse faster
○ Temperature
■ Higher temperatures increase kinetic energy which results in faster diffusion
● Equilibrium is reached when there is no net movement of molecules in one direction
Diffusion Across the Plasma Membrane

● Molecules have a natural drive to diffuse down concentration gradients that exist between
extracellular and intracellular areas
● Nonpolar, hydrophobic lipid core of the PM prevents the diffusion of some substances and
creates concentration gradients by acting as a selectively permeable barrier
● Molecules that are able to diffuse passively through the membrane include:
○ Lipid-soluble and nonpolar substances
○ Very small molecules that can pass through membrane or membrane channels
○ Referred to as simple diffusion
● Flux- movement of molecules in any certain direction
Simple Diffusion
● Nonpolar lipid-soluble (hydrophobic) substances diffuse directly through phospholipid bilayer
○ Eg: oxygen, carbon dioxide, steroid hormones, fatty acids
● Small amounts of very small polar substances, such as water, can also pass through
Facilitated Diffusion
● Larger, non-lipid soluble or polar molecules can cross membrane but only with the support of
carrier molecules
○ This is facilitated diffusion
● There are 2 types:
○ Carrier- mediated
○ Channel- mediated
● Use different types of integral membrane proteins
● Impact different properties to transport
Carrier-Mediated Facilitated Diffusion

● Certain hydrophobic molecules are transported passively down their concentration gradient by
carriers (transmembrane proteins)
● Each carrier transports specific substances
● Binding of molecule causes carrier to envelope and change shape (conformational change) that
results in the molecule being moved across the membrane
● The rate of transport of a substance across a membrane depends on three factors
○ Find in Textbook
Channel Mediated Facilitated Diffusion

● Some integral membrane proteins form channels that allow ions to diffuse across the
membrane
● Ion channels show selectivity for particular types of ions based on channel diameter, charged
residue lining pore, water of hydration
Osmosis
● Is a special name for the net movement of water across a selectively permeable membrane
● Water diffuses across plasma membranes
○ Some through lipid bilayer (some sneak past ist hydrophobic barriers)
○ Mostly through specific water channels called aquaporins (AQP’s)
■ Prevent the passage of ions and other solutes. Only water can go through
■ Impermeable to charged species
● Flow occurs when the water concentration is different on the two sides of the membrane
● Water is a polar molecule but it rapidly diffuses across the plasma membrane of most cells
● A liter of pure water is 55.5 M (moles/mass)
● The addition of solute lowers the water concentration
○ This depends on the number of solute particles, not the chemical nature of the solute
Osmolarity
● Osmolarity: measures the total number of solute particles in a solution
○ Water concentration varies with number of solute particles- solute particles displace
water molecules
■ When solute concentration goes up, water concentration goes down and vise
versa
● Osmolarity= molarity x the number of particles that a substance forms in water
○ 1 M solution of glucose is 1 Osm
○ 1 M solution NaCl ionizes to Na+ and Cl- (2 particles) is 2 Osm (osmolar or osmol/L)
○ 1 M solution of MgCl2 is 3 Osm
○ A 3 Osm solution may have 1 M glucose and 2 M NaCl
● Water moves by osmosis from areas of high solute (high water) to areas of low solute (low
water) concentration
● ICF and ECF is approximately 300 mOsm
Osmotic Pressure
● When a solution containing solutes separated from pure water by a semi-permeable membrane
(permeable to water but not solutes), the pressure that must be applied to the solution to prevent
the net flow of water into it is called the osmotic pressure
● The greater the osmolarity of a solution, the greater its osmotic pressure and the lower its water
concentration
Membrane Permeability Influences Diffusion and Osmosis

● When solutions of different osmolarities are separated by a membrane permeable to all
molecules, diffusion of solutes and osmosis of water occur
○ Cross membrane until equilibrium of solutes and water is reached
● At equilibrium: Same concentration of solutes and water molecules on both sides, with equal
volume on both sides
● When solutions of different osmolarities are separated by a membrane that is only permeable to
water, only osmosis (not diffusion) will occur until equilibrium is reached
● Water will have net movement across membrane until its concentration (or osmolarity) is the
same on both sides
● Results in volume changes on both sides
○ Low solutes side volume decreases
○ High solute side volume increases
○ Only molecules that cannot cross the PM cause volume changes
Osmosis
● Movement of water involves pressures:
○ Hydrostatic pressure: outward pressure exerted on cell side of the membrane caused
by increases in volume of cell due to osmosis
■ Also referred to as ‘back pressure’
○ Osmotic pressure: inward pressure due to the tendency of water to be pulled into a cell
with higher osmolarities
■ The more solutes inside a cell the bigger the pull on water to enter, resulting in
higher osmotic pressures inside the cell
● When hydrostatic pressure equals osmotic pressure, no further net movement of water occurs
○ Water trying to get out equals water trying to get in
● Plant cells are surrounded by strong cell walls that act to limit hydrostatic pressure levels which
in turn limit osmotic pressure
○ Plant cells can only fill with so much water then stop
● Animal cells don’t have cell walls, therefore they cannot limit hydrostatic and osmotic pressures
○ Animal cells will burst if too much water is taken in
● Water can also leave a cell, causing it to shrink
● Changes in cell volume can disrupt cell function
Extracellular Osmolarity and Cell Volume

● Na+, Cl-, K+
● Will not be tested on the last point
Tonicity
● Refers to the ability of a solution to change the shape or tone of cells by altering the cells
internal water volume
○ Isotonic solution has the same osmolarity of non-penetrating solutes as inside the cell,
so volume remains unchanged
○ Hypertonic solution has a higher osmolarity non-penetrating solutes than inside the cell,
so water flows out of the cell, resulting in cell shrinking (crenation)
○ Hypotonic solution has a lower osmolarity non-penetrating solutes than inside the cell,
so water flows into the cell, resulting in cell swelling
■ Can lead to cell bursting (lysis)
Lecture 4- Membrane Transport and Resting Membrane Potential
Carrier-Mediated Transport
● Integral membrane proteins move via conformational changes
● Three factors determine the magnitude of solute flux through a mediated transport system:
○ The extent to which the binding sites are ‘saturated’ (maximally occupied)
○ The number of transporters in the membrane
○ The rate at which the conformational change occurs
● There are many types of transporters that are specific for a substance or class of substances
● Transpor fewer molecules per unit time than ion channels
○ Saturable binding
● There are two types of mediated transport:
○ Facilitated diffusion
○ Active transport
Facilitated Diffusion and Active Transport

● Facilitated diffusion- net flux proceeds across a membrane from higher to lower concentration
● Simple diffusion- flux limited only by the concentration gradients
● In carrier-mediated transport, flux depends on the number of available characters
● Active transport- net movement of solute from a lower concentration to a higher concentration
requires continuous input of energy from ATP
● There are two means of coupling ATP to active transport:
○ The direct use of ATP in primary active transport
○ The use of an electrochemical gradient to drive secondary active transport
Primary Active Transport- Na+/K+ ATPase

● ATP hydrolysis provides the energy for primary active transport
● Transporters are ATPases- enzymes that hydrolyze ATP (break down using water)
○ Eg: Na+/K+ ATPase pump
● How it works:
○ The transporter (with bound ATP) binds 3 Na+ on inside of cell (low affinity for K+)
○ ATPase activated. Autophosphorylation.
○ Conformational change and release of Na+ to outside
○ Increased affinity for K+ allows two K+ to bind
○ Dephosphorylation and return to original conformation. Release of K+ to inside.
● Each ATP hydrolysis moves 3 Na+ outside and 2 K+ inside the cell
A Chemical and Electrical Gradient
● The pumping activity of the Na+/K+ ATPase establishes and maintains the characteristic
distributions of K+ and Na+
● The unequal distribution of ions creates an electrical potential across the PM
○ Can be used for transport and/or signalling
● Thus the Na+/K+ ATPase establishes an electrochemical gradient that:
○ Can be used to do work (eg: transport of other solutes)
○ Is the basis for electrical impulses in neurons
● Na+/K+ ATPase pump uses 10-40% of ATP in cell under resting conditions to maintain
gradient
Secondary Active Transport

● Uses the stored energy of an electrochemical gradient to move both an ion and second solute
across a membrane
○ The creation of the electrochemical gradient depends on the primary active transporter
● Secondary active transport- movement of ion down its electrochemical gradient coupled to
the transport of another molecule
○ These transporters have binding sites for an ion (usually Na+) and the co-transported
molecule
Symporters and Antiporters

● In secondary active transport, the movement of Na+ is always downhill (from high to low)
● Co-transport (symport): the ion and second solute move across the membrane in the same
direction
● Countertransport (antiport): the ion and second solute move in opposite direction
Vesicular Transport
● Involves the transport of large molecules, particles, and fluids across the membranous sacs
called vesicles
● Required cellular energy (usually ATP)
● Vesicular transport processes include;

○ Endocytosis: transport into cells.
■ Are 3 different types: phagocytosis, pinocytosis and receptor-mediated
endocytosis
○ Exocytosis: transport out of cell
○ Transcytosis: transport into, across, then out of the cell
○ Vesicular trafficking: transport from one area or organelle to another
Endocytosis
● Involves formation of protein-coated vesicles (bend membrane)
● Usually involves receptors: therefore can be a very selective process
○ Substance being internalized must be able to bind to its unique receptor
● Once a vesicle is pulled inside the cell, it may:
○ Fuse with lysosome OR
○ Undergo transcytosis
● Some pathogens are capable of hijacking receptors for transport into cell (some take advantage
of acidic lysosome environment)
Phagocytosis
● Type of endocytosis that is referred to as ‘cell eating’
● Membrane projections called pseudopods form and flow around solid particles that are being
engulfed, forming a vesicle that is pulled into a cell
○ Formed vesicle is called a phagosome
○ Phagocytosis is used by macrophages and other white blood cells
● The phagosome combines with a lysosome and its contents are digested
● The vesicle has receptors capable of binding to microorganisms or solid particles
Pinocytosis
● Fluid phase endocytosis or ‘cell drinking’
○ Plasma membrane infolds, bringing extracellular fluid and dissolved solutes inside the
cell
○ Fuses with endosome
● Used by some cells to sample environment
● Is the main way in which nutrient absorption occurs in the small intestine
● Membrane components are recycled back into membrane
● No receptors used, so it is non-specific
Receptor-Mediated Endocytosis
● Involves endocytosis and transcytosis of specific molecules
● Substances bind to specific receptor proteins, enabling the cell to ingest and concentrate specific
substances in protein coated vesicles
● Many cells have receptors embedded in clathrin-coated pits which will be internalized along
with the specific molecule bond
○ Eg: enzymes, low density lipoproteins (LDL), iron and insulin
○ Contain viruses, diphtheria, and cholera toxins may also be taken into the cell this way
● Substances may be released inside cell or digested in a lysosome
● Caveolae have smaller pits and different protein coat from clarathin, but still capture specific
molecules and sometimes use transcytosis
● Associated with lipid rafts (cholesterol and other special lipids)
Overview of Endocytosis By Protein-Coated Pits

1. Coated pit ingestes substance
2. Protein coated vesicle detaches from plasma membrane
3. Coat proteins are recycled to PM
4. Uncoated vesicle fuses with sorting vesicle called an endosome
5. Transport vesicle containing membrane components moves to the plasma membrane for
recycling
6. Fused vesicle may:
a. Fuse with lysosomes for digestion of its contents
b. Deliver its contents to the plasma membrane on the opposite side of the cell
(transcytosis)
Exocytosis
● Process by which material is ejected from the cell
○ Usually activated by cell-surface signals or changes in membrane
● Substances being ejected in secretory vesicles
● Protein on vesicle called v-SNARE finds and hooks up to target t-SNARE proteins on
membrane
● Eg: hormones, neurotransmitters, mucus, cellular wastes
Overview of Exocytosis
1. The membrane bound vesicle migrates to the plasma membrane
2. Proteins at the vesicle surface (v-SNARES) bind with t-SNARES (plasma membrane proteins)
3. The vesicle and plasma membrane fuse and a pore opens up
4. Vesicle contents are released to the cell exterior
Cell Environment Interactions

● Cells interact with the environment by responding directly to other cells, or indirectly to
extracellular chemicals
● Interactions always involve glycolax
○ Cell adhesion molecules (CAMs)
○ Plasma membrane receptors
Cell Adhesion Molecules (CAMs)

● Every cell has thousands of sticky glycoprotein CAMs projecting from membrane
● Functions:
○ Anchor cell to extracellular matrix or to each other
○ Assist in the movement of cells past one another
○ Attract WBCs to injured or infected areas
○ Stimulate synthesis or degradation of adhesive membrane junctions (eg: tight
junctions)
○ Transmit intracellular signals to direct cell migration, proliferation, and specialization
Plasma Membrane Receptors

● Membrane receptor proteins serve as binding sites for several chemical signals
● Contact signalling: cells that touch, recognize each other by each cell’s unique surface
membrane receptors
○ Used in normal development and immunity
● Signal Transduction: interaction between receptor and ligands (chemical messengers) that
cause changes in cellular activity
○ Ligands can be small molecules or proteins
○ In some cells, binding triggers enzyme activation; in others, it opens chemically gated
ion channels causing changes in excitability
■ Eg of ligands: neurotransmitters, hormones and paracrines
○ Some ligands can cause different responses in different cells depending on chemical
pathway that the receptor is part of
○ When ligand binds, receptor protein changes shape (conformation) and becomes
activated
○ Some activated receptors become enzymes; others act directly to open or close ion gates,
causing changes in excitability
○ All transduce signals across the PM
G Protein-Coupled Receptors (GPCRs)

● Activated G protein-coupled receptors indirectly cause cellular changes by attracting G
proteins, which in turn can affect ion channels, activate other enzymes or cause release of
internal second messenger chemicals such as cyclic AMP or calcium
● GPCR Superfamily: >800 7 TM - receptors participate in diverse physiological and
pathological functions
○ 36% of marketed pharmaceuticals target human GPCRs
● Orphan GPCRs: ‘Endogenous’ ligands of more than 140 GPCRs unknown, leaving the natural
functions of those GPCRs in doubt, a great source of drug targets
Overview of G Protein-Coupled Receptors (GPCRs)

1. Ligand (1st messenger) binds to the receptor causing it to change shape and activate.
2. Activated receptor binds to a G protein and activates it. G protein changes shape, causing it to
release GDP and bind GTP
3. Activated G protein activates (or inactivates) an effector protein by causing its shape to change
4. Activated effector enzymes catalyze reactions that produce 2nd messengers in the cell.
(common second messengers: cyclic AMP and C^21)
5. Second messengers activate other enzymes or ion channels. Cyclic AMP usually activates
protein kinase enzymes
6. Kinase enzymes activate other enzymes. They transfer phosphate groups from ATP to specific
proteins and activate a series of other enzymes that trigger various metabolic reactions and
structural changes in the cell.
Lecture 5- Fundamentals Of The Nervous System
Neurons
● Neurons (nerve cells) are highly specialized cells that are the basic functional units of the
nervous system
● Special characteristics:
○ Excitable cells (conduct electrical impulses)
○ Extreme longevity (lasts a person’s lifetime)
○ Postmitotic (don't undergo mitosis) with fe exceptions
○ Higher metabolic rate: require continuous supply of oxygen and glucose
● All neurons have a cell body and one or more slender processes
Neuron Cell Body

● Also called the perikaryon or soma
○ 5-140 um diameter
● Biosynthetic center of neuron
○ Synthesizes proteins, membranes, chemicals
○ Rough ER (chromatophilic substance or nissl bodies)
● Contains spherical nucleus with nucleolus
● Neurons in the SN contain a pigment related to melanin
● PM is usually part of receptive region that receives input from other neurons
● Most neuron bodies are located in the CNS but some are in PNS
○ Nuclei: clusters of neuron cell bodies in the CNS (gray matter)
○ Ganglia: clusters of neuron cell bodies in PNS
Neuron Processes
● Arm like processes that extend from cell body
○ CNS contains both neuron cell bodies and their processes
○ PNS contains chiefly neuron processes

● Tracts (usually white matter)
○ Bundles of neuron processes in CNS
● Nerves
○ Bundles of neuron processes in PNS
● Two types of processes:
○ Dendrites
○ Axon
Dendrites
● Neurons can contain 100s of these short tapering, diffusely branched processes
● Same organelles as in stroma
● Receptive (input) region of neuron
● Convey incoming messages toward cell body as graded potentials (short distance signals)
● In many brain areas, finer dendrites are highly specialized to collect information
○ Contain dendritic spines, appendages with bulbous or spiky ends
○ Shape modified by activity- basis for learning and memory
Axon Structure
● Each neuron has a single axon that starts a cone shaped area close to the soma called axon
hillock
● Axons can be short or extremely long (>1 meter)- called nerve fibers
● Can have occasional branches (at 90 degrees) called axon collaterals
● Axons branch profusely at their end (terminus)- as many as 10,000 terminal branches
● Distal endings are called axon terminals or terminal boutons
Axon Function
● Axon is the conducting region of neuron (conducts information away from the cell body)
● Generates nerve impulses and transmits them along axolemma (axon cell membrane) to axon
terminal, a region that secretes neurotransmitters, which are released into the extracellular
space
○ Can excite or inhibit neurons it contacts
● Communicates with many different neurons at the same time
● Axons rely on cell bodies to renew proteins and membranes
● Quickly decay of cut or damaged (PNS vs CNS)
Axonal Transport
● Axons have efficient internal transport mechanisms
● Molecules and organelles are moved along axons by motor proteins and cytoskeletal elements
○ Anterograde: away from the cell body
■ Eg: mitochondria, cytoskeletal elements, membrane components, enzymes,

synapses-specific proteins, neurotransmitters, certain mRNA’s
○ Retrograde: towards cell body
■ Eg: organelles to be degraded, signalling molecules, viruses and bacterial toxins
Peripheral Myelination
● Formed by schwann cells
○ Wraps around axon
○ One cell forms one segment of the myelin sheath
● Outer collar of perinuclear cytoplasm: peripheral bulge containing nucleus and most of
cytoplasm
● Plasma membranes have less proteins
● No channels or carriers, so good electrical insulators
● Cell adhesion molecules (CAMs) bind to adjacent myelin membranes
● Myelin Sheath Gaps
○ Gaps between adjacent schwann cells
○ Sites where axon collaterals can emerge
○ Also called nodes of ranvier
● Non-Myelinated Fibers
○ Thin fibers not wrapped in myelin
○ Surrounded by schwann cells but no coiling
○ One cell may surround 15 different fibers
Central Myelination
● Myelin sheaths in the CNS
○ Formed by processes of oligodendrocytes, not whole cells
○ Each cell can wrap up to 60 axons at once
○ Also forms nodes of ranvier
○ No outer collar of perinuclear cytoplasm
○ Speeds up transmission of action potentials
○ Thinnest fibers are unmyelinated, but covered by long extensions of adjacent neuroglia
○ White matter: regions of the brain and spinal cord with dense collection of myelinated
fibers
■ Usually fiber tracts
○ Gray matter: mostly neuron cell bodies and unmyelinated fibers
Membrane Potential in Neurons

● Like all cells, neurons have a resting membrane potential- charge difference across PM
● Neurons can rapidly change the membrane potential due to the presence of additional ion
channels
● Neurons (and muscle cells) are excitable cells as they can conduct electrical potentials
Basic Principles of Electricity

● Opposite charges attract each other and will move towards each other if not separated by some
barrier
● Separate charges have the ability to do work- called electrical potential
● Difference in charge between two points is called potential difference (or just potential)
○ Measured in volts (v)
● Movement of electrical charge: current
● Ohm’s Law: Current is proportional to the potential difference and inversely proportional to
the resistance I=V/R
● Insulator: a substance with high electrical resistance
● Conductor: substances with low electrical resistance
Summary: Establishing The RMP

● The Na+/K+ ATPase establishes a concentration gradient and generates a small negative
potential
○ Pump uses up to 40% of the ATP produced but the cell
● Greater net movement of K+ than Na+ makes the membrane potential more negative on the
inside
● At a steady state, ion fluxes through the channels and activity of the pump balance each other
Distribution of Ions in a Nerve Cell

● Na+, K+ and Cl- are present in the highest concentration and membrane permeability to each is
independently determined
● Na+ and K+ play the most important role in generating the resting membrane potential but Cl-
is a factor in some neurons
● Remember: Na+ Cl- out, K+ in
● The resting membrane potential is established and determined mainly by Na+/K+ ATPase
activity
● Cl- is NOT on Exam
Equilibrium Potential
● The magnitude of the membrane potential depends mainly on two factors
○ (Ion) differences out vs in
○ Membrane permeability to different ions (# of open channels for each ion)
● A membrane that is permeable only to K+ (refer to this slide in lecture 5 ppt)
● The membrane potential at which the flux due to concentration difference becomes equal in
magnitude but opposite in direction is called the equilibrium potential for that type of ion
● Consider a membrane permeable only to Na+ (refer to this slide in lecture 5 ppt)
The Nernst Equation

● The Nernst equation describes the equilibrium potential for any ion species
○ The electrical potential necessary to balance a given ionic concentration gradient across
a membrane so that the net flux on that ion is zero
● Eion= 61/Z log (Co/Ci)
○ Eion = equilibrium potential in mV
○ Ci = intracellular ion concentration
○ Co = extracellular ion concentration
○ Z = valence of the ion
○ 61 is a constant that takes into account the universal gas constant, the temperature, and
the Faraday electrical constant
● When there is a bigger number over a smaller number, the log will be positive and when
there is a smaller number over a bigger number, the log will be negative
○ ENa= 61/1 log (145/15) = +60 mV
○ EK= 61/1 log (5/15) = -90 mV
● At typical concentrations (table 6.2), Na+ flux through open channels will drive the membrane
potential towards +60 mV while K+ flux will bring it towards -90 mV
○ In an actual nerve at rest, there are many more open K+ channels than Na+ channels;
chloride permeability generally falls in between
○ PK= 1, PNa= 0.04, PCl= 0.45
Forces Influencing Na+ and K+ At Resting Membrane Potential

● At a resting membrane potential of -70 mV, both the concentration gradient and electrical
potential favour inward movement of Na+ while K+ concentration gradient opposes the
electrical potential
● The greater permeability and movement of K+ maintains the resting membrane potential at a
value near EK
Resting Membrane Potential

● Electrical potential energy produced is by the separation of oppositely charged particles across
the plasma membrane in all cells
○ The difference in electrical charge between two points is referred to as voltage
● A voltmeter can measure the potential difference across the membrane of a resting neuron of
-70 mv
● Resting membrane voltages range from -50 to -100 mV in different cells
○ The membrane is said to be polarized
● All cells under resting conditions have a potential difference with the inside being more
negative
○ This is called the resting membrane potential
● ECF has a voltage of zero
● If the potential across the membrane is 70 mV and the ICF has excess negative charge, then
membrane potential is -70 mV
● Membrane potential is determined by two factors:

○ Differences in ionic composition of ICF and ECF
○ Differences in plasma membrane permeability to each ion
Charge Separation Across Plasma Membrane

● Voltage occurs only at the membrane surface
○ Rest of cell and extracellular fluid are neutral
● There is a tiny excess of negative ions along inner surface of PM and positive ions around the
outside
● Excess charges collect at the plasma membrane and are attracted to each other
● The actual number of charges that are separated is small compared to the total number of
charges in the cell
● Only a very thin shell of charge difference is needed to establish a membrane potential
Resting Membrane Potential

● Depends on:
○ Differences in K+ and Na+ concentrations inside and outside cells
○ Differences in permeability of the PM to these ions
● Energy is required to keep opposite charges separated across a membrane
Leak Channels Generate the Resting Potential

● Leak Channels: (not ligand-gated) are integral membrane proteins that are selective ion
channels- some are always open
● In a neuron at rest, there are many more open K+ leak channels than Na+ leak channels
● So PM is 25 times more permeable to K+ than Na+
● This brings RMP closer to the K+ equilibrium potential
● therefore , the RMP depends mostly on K+
○ Na+/K+ ATPase maintains resting membrane potential by maintaining concentration
gradients
Forces Influencing Na+ and K+ At Rest

● At a resting membrane potential of -70 mv, both the concentration gradient and electrical
potential favour inward movement of Na+, while the K+ concentration gradient opposes the
electrical potential
● The greater permeability and movement of K+ maintains the resting potential at a value near EK
Types of Ion Gated Channels

● Chemically Gated (ligand gated) Channels
○ Open only with binding of specific chemical (eg: neurotransmitter)
● Voltage Gated Channels
○ Open and close in response to changes in membrane potential
● Mechanically Gated Channels

○ Open and close in response to physical deformation of receptors, as in sensory receptors
● When gated channels open, ions diffuse quickly:
○ Along chemical concentration gradients from high to low concentration
○ Along electrical gradients, towards opposite electrical charge
Changes in Membrane Potential

● Membrane potential changes when:
○ Concentrations of ions across membrane change
○ Membrane permeability to ions change (opening of channels)
● Changes produce two types of signals
○ Graded potentials
■ Incoming signals operating over short distance
○ Action potentials
■ Long- distance signals of axons
● Changes in membrane potential are used as signals to receive, integrate and send information
● Terms describing membrane potential changes relative to RMP:
○ Depolarization: a decrease in membrane potential (moves towards zero and above)
■ Inside of membrane becomes less negative than RMP
■ Probability of producing impulses increases
○ Hyperpolarization: increase in membrane potential (away from zero)
■ Inside of a membrane becomes more negative than RMP
■ Probability of producing impulse decreases
Graded Potentials
● Short lived, localized changes in membrane potential
○ The stronger the stimulus, the more voltage changes and the further current flows
● Triggered by stimulus that opens gated ion channels
○ Results in depolarization or hyperpolarization
● Named according to location and function
○ Receptor potential (generator potential): graded potentials in receptors of sensory
neurons
○ Postsynaptic potential: neuron graded potential
● Once gated ion channel opens, depolarization spreads from one area of membrane to the next
● Current flows but dissipates quickly and decays
○ Graded potentials are signals only over short distances
Action Potentials
● Principle way neurons send signals
○ Means of long distance neural communication
● Only occur in muscle cells and axons of neurons (excitable cells)
○ Aka nerve impulse

● Involves opening of specific voltage-gated channels
● Brief reversal of membrane potential with a change in voltage of 100 mV
● Don’t decay with distance like graded potentials
Voltage-Gated Channels
● An action potential (AP) results from changes in ion permeability due to the function of
voltage-gated Na+ and K+ channels
● APs are generally very rapid (can be a few milliseconds) and can have frequencies of several
hundred/second
● Voltage-gated channels give the membrane the ability to generate and propagate APs
Action Potential Mechanism

● Absolute refractory period
○ The period of time during which a second action potential ABSOLUTELY cannot be
initiated, no matter how large the applied stimulus is
● Relative refractory period
○ the period shortly after the firing of a nerve fiber when partial repolarization has
occurred and a greater than normal stimulus can stimulate a second response
● Refer to ppt 5 and textbook
Action Potentials are All or Nothing

● APs occur in excitable membranes with voltage gated sodium channels (VGSCs)
● These channels open as the membrane depolarizes, causing a positive feedback opening of more
VGSCs and moving the membrane potential towards ENa
● Local anesthetics (procaine and lidocaine) block VGSCs
● The number of ions that cross the membrane during an AP is extremely small compared to the
total number of ions in the cell, producing only small changes in the IC ion concentrations
Threshold Potential
● Depolarization triggers an AP only when the membrane potential exceeds a threshold potential
(55 mV)
● Regardless of the size of the initial stimulus, if the membrane reaches threshold, the APs
generated are all the same size
● APs propagate without any change in size from one site to another along a membrane
○ Don’t decay like graded potentials
● Because it its ‘all or none’, a single AP cannot convey the information about the magnitude of
the initial stimulus
● How is information encoded in the nervous system?
Action Potential Propagation

● The (local) current entering during an AP is sufficient to easily depolarize the adjacent
emembrane to the threshold potential
● The propagation of the AP from the dendrites to the axon terminal is typically one way because
the absolute refractory period follows along the wake of the moving AP
● The speed of propagation depends on fiber diameter (why?) and whether or not the fiber is
myelinated
Saltatory Conduction
● In myelinated nerve fibers, APs undergo saltatory conduction
● Action potentials jump from one node of ranvier to the next as they propagate along a
myelinated axon
● Multiple Sclerosis- Myelin breakdown
Lecture 6- Synapses
The Synapse
● Nervous system works because information flows from neuron to neuron
● Neurons are functionally connected by synapses- junctions that mediate information transfer
○ From one neuron to another neuron
○ OR from one neuron to an effector cell
● Presynaptic neuron: neuron conducting impulses towards synapse (sends information)
● Postsynaptic neuron: neuron transmitting electrical signal away from synapse (receives
information)
○ In PNS may be a neuron, muscle cell or gland cell
● Most neurons function as both
Synaptic Connections
● Axodendritic: between axon terminals of one neuron and dendrites of another
● Axosomatic: between axon terminals of one neuron and soma (cell body) of others
● Less common connections;
○ Axoaxonic: axon to axon
○ Dendrodendritic: dendrite to dendrite
○ Somatodendritic: dendrite to soma
● Two main types of synapses
○ Chemical synapse
○ Electrical synapse
Chemical Synapses
● Is the most common type
● Specialized for release and reception of chemical neurotransmitters
● Typically composed of two parts:
○ Axon terminal of presynaptic neuron: contains synaptic vesicles filled with

neurotransmitter
○ Receptor region on postsynaptic neuron’s membrane: receives neurotransmitter
■ Usually on dendrite or cell body
○ Two parts separated by fluid-filled synaptic cleft
● Electrical impulse changed to chemical across synapse, then back into electrical
● Transmission across synaptic cleft
○ Synaptic cleft prevents nerve impulses from directly passing from one neuron to the
next
○ Chemical event (as opposed to an electrical one)
○ Depends on release, diffusion, and receptor binding of neurotransmitters
○ Ensures unidirectional communication between neurons
Events at Chemical Synapses

1. AP arrives at presynaptic terminal
2. Voltage-gated Ca2+ channels open in response to depolarization.
3. Ca2+ influx causes synaptic vesicles to fuse with PM and release neurotransmitter (NT) by
exocytosis.
4. NT diffuses across the synaptic cleft and binds to its specific postsynaptic receptor(s)
5. Neurotransmitter binding opens ion channels resulting in a graded potential
6. NT effects are terminated by several mechanisms
a. Reuptake by transporters
b. Diffusion away from synapse
c. Enzymatic degradation
Synaptic Delay
● Time needed for neurotransmitter to be released, diffuse across the synapse, and bind to
receptors
○ Can take anywhere from 0.3 to 5.0 ms
● Rate-limiting step of neural transmission
● Transmission of AP down axon can be very quick, but synapse slows transmission to
postsynaptic neuron down significantly
● Not noticeable, because these are still very fast
Electrical Synapses
● Less common than chemical synapses
○ Don’t have delays like chemical synapses
● Neurons are electrically coupled
○ Joined by gap junctions that connect the cytoplasm of adjacent neurons
○ Communication is very rapid and may be unidirectional or bidirectional
○ Found in some brain regions responsible for eye movements or hippocampus in areas
involved in emotions and memory
○ More abundant in embryonic nervous tissue, heart and smooth muscle
Postsynaptic Potentials
● Neurotransmitter receptors cause graded potentials that vary in strength based on:
○ Amount of neurotransmitter released
○ Time neurotransmitter stays bound to its receptor
● Depending on the effect of chemical synapse, there are two types of postsynaptic potentials
○ EPSP: excitatory postsynaptic potentials
○ IPSP: inhibitory postsynaptic potentials
Excitatory Synapses and EPSPs

● Neurotransmitter binding opens chemically gated channels (usually a non-selective cation
channel)
● Allows simultaneous flow of Na+ and K+ in opposite directions
● Na+ influx greater than K+ efflux, resulting in a depolarizing graded potential called excitatory
postsynaptic potential (EPSP)
● EPSPs trigger AP at axon hillock if the membrane potential reaches threshold
○ This triggers opening of voltage-gated Na+ channels
● Glutamate is the major excitatory NT in the CNS
● Occur on dendritic spines
Inhibitory Synapses and IPSPs

● Neurotransmitter binding to receptor opens chemically gated channels that allow influx/efflux
of ions that cause a hyperpolarization.
○ e.g. K+ channel or Cl- channel
● Moves the membrane potential further away (more negative) from threshold to fire an AP
○ Called in inhibitory postsynaptic potential (IPSP)
● Glycine and gamma-aminobutyric acid (GABA) are inhibitory neurotransmitters in the CNS
Integration and Modification of Synaptic Events

● Postsynaptic
○ Often, a single EPSP cannot induce an AP, but EPSPs can summate (add together) to
influence postsynaptic neuron
■ IPSPs can also summate
○ Most neurons receive both excitatory and inhibitory inputs from thousands of other
neurons
■ Only if EPSPs predominate and bring to threshold will an AP be generated
● There are two types:

● Temporal summation
○ One or more presynaptic neurons transmit impulses in rapid-fire order
○ First impulse produces EPSP, and before it can dissipate another EPSP is triggered,
adding on top of first impulse
● Spatial summation
○ Postsynaptic neuron is stimulated by a large number of terminals simultaneously
○ Many receptors are activated, each producing EPSPs, which can then add together
Activity Dependant Synaptic Potentiation

● Repeated use of synapse increases ability of presynaptic neuron to excite postsynaptic neuron
● Presynaptic changes
○ Ca2+ concentration increases in presynaptic terminal - release of more neurotransmitter -
more EPSPs in postsynaptic neuron
● Postsynaptic changes
○ Potentiation can cause Ca2+ voltage-gated channels to open in postsynaptic neuron. Ca2+
activates kinases, leading to more effective response to subsequent stimuli
○ NMDA receptors (Mg2+ block removed by strong depolarization)
○ Long-term potentiation involved in learning and memory
Comparison of Graded and Action Potentials

● Refer to ANP Reference Charts
Developmental Aspects of Neurons

● Nervous system originates from neural tube and neural crest (ectoderm)
○ The neural tube becomes CNS
● Neuroepithelial cells of neural tube proliferate into number of cells needed for development
● Neuroblasts
○ Become amitotic and migrate to final positions
○ Sprout axons to connect with targets and become neurons
● Growth cone: structure at tip of axon that allows it to interact with its environment via:
○ Cell surface adhesion proteins (laminin, integrin, and nerve cell adhesion molecules,
or N-CAMs), which provide anchor points
○ Neurotrophins that attract or repel the growth cone
○ Nerve growth factor (NGF), which keeps neuroblast alive
○ Filopodia are growth cone processes that follow signals toward target
Developmental Aspects of Neurons

● Once axon finds its target, it then must find the right place to form synapse (Roger Sperry -
Chemoaffinity Hypothesis)
● About two-thirds of neurons die before birth
○ If axons do not form a synapse with their target, they are triggered to undergo apoptosis
(programmed cell death)
○ Many other cells also undergo apoptosis during development
● During postnatal development, learning reinforces certain synapses and prunes away others
(activity-dependent synaptic plasticity)
Lecture 7- Skeletal Muscles

Muscles- The Basics
● Are responsible for all of our movements- limbs, trunk, bowel
● Is the largest tissue by weight (40-50% of body mass)
○ Are 640 different muscles in the human body
● Come in various shapes and sizes but all have the same primary function
● Major functions
○ Control of body movement
○ Stabilize body position (postural muscles)
○ Regulate organ volume (sphincters)
○ Move substances within the body (cardiac, smooth muscle)
○ Produce heat
● Key properties
○ Excitability- receive and respond to stimuli (conduct APs)
○ Contractility- shorten forcibly
○ Extensibility- can be stretched
○ Elasticity- resume resting strength
Comparison of Skeletal, Cardiac and Smooth Muscle

● Distinguished by shape, number & position of nuclei, presence of striations, and
whether they are under voluntary or involuntary control
● Skeletal
○ Elongated cells with multiple peripheral nuclei; striations (light and dark bands)
○ Under voluntary control.
● Cardiac
○ Branching cells with a single central nucleus; striated
○ Under involuntary control.
● Smooth
○ Single nucleus within a spindle-shaped cell.
○ Do not have visible striations.

○ Smooth muscle is under involuntary control.
Organizational Levels of Skeletal Muscles

● Skeletal muscle is what you cook on the barbecue on summer evenings
● Each skeletal muscle is a separate organ composed of hundreds to thousands of muscle cells
called fibers
○ Muscle fibers are bundled together in fascicles.
● Control of skeletal muscle is primarily voluntary – you can consciously control them - but
also involuntary – postural muscles and diaphragm.
● Tubes within tubes organization
● In order from smallest to largest:
○ Myofilaments - Myofibril- Muscle cell- Fascicle- Muscle
Internal Structure of a Skeletal Muscle

● Skeletal muscles:
○ contractile cells
○ connective tissue
● Connective tissue:
○ Epimysium – surrounds
muscle
● Perimysium
○ surrounds fascicles (bundles
of muscle cells).
○ Conduit for nerves and blood
vessels.
● Endomysium separates and
electrically insulates the muscle
cells.
○ Strength and support for
entire muscle. Is continuous with the tendons.
Microscopic Organization of a Skeletal Muscle Fiber

● The majority of space within a muscle fiber is taken up by the contractile elements - the thin
and thick filaments - which are bundled into structures called myofibrils.
● A single muscle fibers contain numerous mitochondria and many nuclei
● The myonuclei lie just beneath the sarcolemma (plasma membrane)
Fusion of Myoblasts Into Muscle Fibers

● Multinucleated muscle fibers: from fusion of > 100 myoblasts.
● After fusion, the muscle fiber is terminally differentiated (postmitotic)
○ Number of muscle fibers set before birth

● Postnatal growth achieved by enlargement of existing muscle fibers (hypertrophy) by
increasing the number of contractile filaments (myofibrils) = more forceful contractions.
○ Hypertrophy also occurs in response to strength training
● Satellite cells:
○ Stem-like cells that persist in mature skeletal muscle.
○ Injury - proliferate and fuse with one another or with damaged fibers to regenerate
functional muscle.
○ Contribute to growth, repair, and hypertrophy
○ Contained within the endomysium (basal lamina)
Transverse (T) Tubes

● Extensions of sarcolemma (PM) - penetrate into the cytoplasm,
● Pass between and surround each myofibril
● Connect with other T-tubules
● Each T-tubule is flanked by SR on either side (triad)
● Function: Conduct APs from surface into the interior
● Note: The inside of the T-tubule is continuous with the ECF!
Sarcoplasmic Reticulum (SR)

● Specialized smooth ER that surrounds each myofibril (bundles of myofilaments).
○ Connected together across fiber width but not length.
○ Stores Ca2+ in enlarged regions called terminal cisternae.
○ Initiates muscle contraction by releasing Ca2+ from the terminal cisternae when
prompted by the T-tubular system.
Structure of Myofibril
● The myofibrils that are packed into a muscle cell are composed of individual contractile
elements called myofilaments.
● Two types of myofilaments:
○ The thin filament is composed mainly of the protein actin
○ The thick filament is made up chiefly of the protein myosin
Arrangement of Myofilaments in a Myofibril

● The arrangement of thick and thin filaments forms light and dark alternating bands (striations)
along the myofibrils.
○ A bands (Dark) = length of thick filaments.
○ I bands (Light) = only thin filaments; corresponds to distance between adjacent thick
filaments.
■ I band bisected by Z line = a protein disc which anchors the thin filaments and
connect adjacent myofibrils.
○ H zone, region between the thin filaments.

○ M line, located in the center of H zone, consists of protein fibers that connect
neighboring thick filaments.
● Sarcomere – Region of myofibril between 2 successive Z lines : minimal contractile unit
Organization of Thick and Thin Filaments

● Within a sarcomere, thick and thin filaments interdigitate so that in cross section they are seen
to form a hexagonal lattice,
● 6 thin filaments arrayed around each thick filament.
● Thick filaments are also arranged hexagonally to each other.
Structure of Myofilaments
● Refer to slide in ppt for diagram
The Sliding Filament Theory

● The changing relationships of the thick and thin filaments can be observed when comparing
electron micrographs of shortened and lengthened skeletal muscle in longitudinal section.
● Proposed by Hugh Huxley et al. (1957) based on the evidence that the I band length decreases
while the A band length remains almost constant during muscle shortening. The filaments
themselves do not change length, only position: Z lines come together
● The sliding filament theory states that the contraction of a muscle cell occurs as the thin
filaments slide past the thick filaments. During contraction, the sarcomere shortens and the thin
and thick filaments overlap to a greater degree, thereby shortening the entire muscle.
Length-Tension Relationship
● Forcefulness of muscle contraction depends on the length of the sarcomeres before contraction
begins.
● Tension produced by a sarcomere should be proportional to the number of cross-bridges or the
amount of overlap between the thick and thin filaments.
● Muscle fiber will develop the greatest tension when there is optimal overlap between the thick
and thin filaments
The Cross Bridge Cycle

•This is a cycle, a continuous process, so you could
imagine any step as the starting point.
•Ca+ allows myosin to bind to actin
Cross Bridge Cycle I

•ATP is bound to the myosin head.
•Myosin is in its low energy (E) conformation (45° to
thin filament) and is not attached to actin.
Cross Bridge Cycle II

•Myosin is an ATPase: Enzyme hydrolyzes ATP into ADP and Pi.
•ADP + Pi, remain in the ATP-binding site
•This leaves the myosin head in a high E state, with a preferred orientation of 90° to the thin
filament.
● Note: When muscles are at rest, myosin is "energized" with ADP and Pi bound.
Cross-Bridge Cycle III

● Energized myosin can now bind to the myosin binding sites on actin.
Cross-Bridge IV
● Myosin head binding to thin filament causes a conformational change- releases ADP and Pi and
changes orientation of thick filament to a low E state (45°) to thin filament.
● Transition to low E state produces a force and lateral movement of thick and thin filaments:
Power Stroke. Myosin heads rotate towards center of sarcomere –heads oriented in different
directions on either side of the M line.
Cross-Bridge Cycle I
● Binding of ATP to the myosin head causes detachment of the cross-bridge from actin.
Overview of Cross-Bridge Cycle

1. ATP binding to myosin head causes detachment of the cross-bridge from actin.
2. Myosin hydrolyzes ATP into ADP and Pi, which remain in the ATP-binding site. Myosin head
is in high-energy state, 90° to thin filament. Myosin energized in resting muscle.
3. Myosin binds to actin forming a cross-bridge.
4. ADP and Pi released, changes the preferred orientation of the thick filament to a lower-energy
state at 45° to the thin filament. Transition to lower-energy state produces force and lateral
movement of the thick and thin filaments. (POWER STROKE). Myosin heads rotate towards
the center of the sarcomere.
Note: Cross-bridge formation is asynchronous resulting in smooth & sustained force production
whose magnitude depends on the number of interacting sites and the force generated by each site. Each
of the 600 myosin heads in a thick filament attach and detach ~5x/sec.
Overview of Contraction
● Cross bridge movement results in sliding of thick and thin filaments past one another.
○ Hundreds of simultaneous cross-bridge movements shorten the sarcomere.
○ The width of the I-band decreases as the thick filaments slide towards the Z-lines of the
sarcomere.
● The sliding filament model predicts the repetitive formation of cross-bridges between the
myosin and actin filaments, shortening of the sarcomeres along the length of the myofibrils,
shortening of the fiber and muscle, and consequent force development.
Regulation of Contraction By Ca2+

● Contraction driven by ATP hydrolysis but triggered by Ca2+ release from SR.
● Contraction requires Ca2+ entry into the cytoplasm, while relaxation requires its removal.
● Triad consists of one T-tubule lying between two adjacent terminal cisternae.
Role of Ca2+ In The Contraction Mechanism

● At low intracellular [Ca2+] (~10-7 M):
○ Tropomyosin blocks the binding sites on actin.
○ Myosin cross bridges cannot attach to binding sites on actin
○ The relaxed state of the muscle is enforced
● At high intracellular [Ca2+] (~10-3 M):
○ Troponin binds Ca2+ and undergoes a conformational change
○ This change uncovers the myosin binding sites on actin so that cross bridges can form
Regulation of Contraction by Ca2+ Release

● Tropomyosin prevents the myosin heads from binding to actin in relaxed muscle when ATP is
present, but Ca2+ is low.
Regulation of Contraction by Ca2+ (Return to SR)

● Active transport (requires ATP) of Ca2+ back into SR by specialized ion pumps in the SR
membrane called Sarco[endo]plasmic reticulum Ca2+ ATPases (SERCAs)
● Ca2+-binding proteins (calsequestrin, calreticulin, etc.) act as a buffer for Ca2+ storage – lower
free [Ca2+] in the lumen.
Roles of ATP in Skeletal Muscle Contraction

1. Hydrolysis of ATP by myosin energizes the cross bridges, providing the energy for force
generation (Power Stroke)
2. Binding of ATP to myosin disconnects the myosin head from actin, allowing the cross bridge
cycle to repeat.
3. Hydrolysis of ATP by SERCAs provides the energy for active transport of Ca2+ back into the
SR, which lowers the [Ca2+] in the cytoplasm, ending the contraction and allowing the muscle to
relax.
Rigor Mortis
● ATP hydrolysis required for contraction and relaxation.
● After death, membranes become leaky. Ca2+ leaks into cytosol - allows myosin to bind actin.
ATP synthesis has ceased, - so cross-bridges cannot detach from actin.
● Rigor mortis (rigidity of death) muscles cannot contract or stretch. Begins 3-4 hrs after death
and lasts ~24 hrs
The Neuromuscular Junction (NMJ)

● Skeletal muscles innervated by motor neurons of the somatic nervous system.
○ Axons of these neurons travel in nerves and branch profusely as they enter muscles.
● Neuromuscular junction (NMJ): point of contact between a motor neuron and the muscle
fibers it innervates ~0.01% of the muscle fiber’s surface area.
○ As a rule, each muscle fiber is innervated by a single axon and has only one NMJ,
located approximately midway along the fiber’s length.
● NMJs consist of:
○ Axonal endings - synaptic vesicles contain the neurotransmitter acetylcholine (ACh).
○ The motor end plate - a specialized part of the sarcolemma (highly folded) that
contains nicotinic acetylcholine receptors (nAChRs)
○ Separated by the synaptic cleft.
○ A basal lamina (endomysium) is present in the synaptic cleft.
○ nAChRs concentrated at crests of postsynaptic folds (10,000/um2), in precise register
with presynaptic release zones.
Role of ACh Receptors

● nAChR - ligand-gated ion (cation selective) channel.
● ACh binding causes nAChRs to open, allowing cations (Na+ and K+) to flow through.
○ Produces a local depolarization - end plate potential (EPP).
● EPP causes a conformational change in voltage-gated Na+ channels, which are concentrated in
the postsynaptic membrane (~10 fold higher than elsewhere on the sarcolemma). These begin
(and propagate) AP.
● ACh in the synaptic cleft diffuses away or is broken down by acetylcholinesterase (AChE)
into acetic acid and choline - prevents continued contraction in the absence of additional stimuli
● AChE inhibitors include neostigmine (treatment of myasthenia gravis, glaucoma) and potent
nerve agents (sarin, VX and Novichok)
Summary of Events At The NMJ

1. AP arrives at the axon terminal of motor neuron.
2. Voltage-gated Ca2+ channels open. Ca2+ enters axon terminal.
3. Ca2+ entry causes acetylcholine (ACh) release by exocytosis.
4. ACh diffuses across the synaptic cleft and binds to ACh receptors (AChRs) on the
sarcolemma.
5. ACh binding opens ligand-gated ion channels that passage of Na+ and K+ which produces a
local depolarization called the end plate potential (EPP)
6. ACh effects are terminated by its breakdown in the synaptic cleft by acetylcholinesterase
(AChE) and diffusion away from the junction.
Summary of Excitation- Contraction Coupling

● Refer to ANP reference charts
Twitch Contractions and Wave Summation

● Mechanism: additional Ca2+ released form SR promotes continued contraction
Three Phases of a Muscle Twitch

● A muscle twitch includes three phases: actual times for each phase varies with specific muscle
being studied
● Latent Phase
○ Sarcolemma and T-tubules depolarize. Ca2+ released into cytosol and cross-bridges
begin to cycle.
○ The number of cross-bridges not sufficient to visibly shortened muscle.
○ Lasts < 5 msec
● Contraction Phase
○ Sarcomeres shorten as a result of myosin cross-bridge cycling. Continues until peak
tension.
○ Speed with which this phase occurs depends on the load being lifted and the fiber type:
fast- or slow-twitch
● Relaxation Phase
○ Ca2+ in cytosol rapidly decreases - actively transported back into terminal cisternae.
○ Cross-bridge cycling decreases and stops.
○ Tension (force) is reduced, allowing the muscle to return to its original length.
Motor Units
● A motor neuron and all the muscle cells it stimulates is called a motor unit.
● Each neuron branches forming neuromuscular junctions with several muscle cells.
○ Different motor units may contain different numbers (and types as we will see) of
muscle cells and produce differing degrees of force.
● Strength of a contraction proportional to the size of the motor units and the number activated.
Recruitment
● The stimulation of additional motor units for increased strength of contraction is called
recruitment.
○ Weakest motor units recruited first, followed by stronger ones.
● Allows for smooth, graded contractions of whole muscles
● Motor neurons to a whole muscle fire asynchronously. Alternating activity delays onset of
muscle fatigue.
● Even during a “maximal voluntary contraction'', not all motor units activated.
Muscle Tone
● Continuous, passive partial contraction of the muscles, or resistance to passive stretch at rest -
steady state condition for muscles.
● Small groups of motor units are alternatively active and inactive in a constantly shifting pattern.
○ Helps keep muscles firm, but it does not result in a contraction strong enough to produce
movement. Without nerve stimulation, for example if the motor nerve is cut or damaged,
the muscle loses all tone and become flaccid.
● Sudden pull or stretch: the body automatically increases muscle's tension, a reflex that helps
guard against danger as well as helping maintain balance.
● Both extensor and flexor muscles are involved in the maintenance of a constant tone while at
rest. In skeletal muscles, this helps maintain a normal posture.
● Cramps - changes in muscle tone in flexors or extensors
Types of Muscle Contractions

● The force exerted on an object by a contracting muscle: tension
● The force exerted on the muscle by an object: load
● Tension and load are opposing forces.
● Isotonic (concentric) contraction
○ The peak tension exceeds the resistance of the load resulting in shortening of the muscle
and movement of the load.
■ Occurs when you use your muscles to successfully push or pull an object.
● Eccentric contraction
○ The muscle lengthens.
● Isometric contraction
○ Peak tension < load. the muscle contracts and develops tension, but its length does not
change.
○ This occurs when a muscle attempts to push or pull an immovable object.
Overview of ATP Synthesis in Muscle

● When ATP supplies are low, muscle cells use three processes to synthesize additional ATP
○ Hydrolysis of creatine phosphate
○ Glycolysis
○ The Krebs cycle (citric acid cycle) and oxidative phosphorylation
Creatine Phosphate
● At rest, muscle fibers produce more ATP than they need. Excess ATP used to synthesize
creatine phosphate or phosphocreatine (PCr).
● Creatine kinase (CK) transfers high energy phosphate group to creatine.
○ Muscle cells use this PCr to store energy. Normal metabolism cannot produce energy as
quickly as a muscle cell can use it, so an extra storage source is needed.
● During exercise, PCr serves as an immediate reserve of high-energy phosphate groups, which is
used to replenish ATP - for the first few seconds of intense activity.
● Elevation of CK in blood is an indication of muscle damage. Clinically, CK is assayed in blood
tests as a marker of myocardial infarction (heart attack), rhabdomyolysis (severe muscle
breakdown), muscular dystrophy, and acute renal failure.
Anaerobic Respiration- Glycolysis

● Initial way of using glucose (all cells). Used exclusively by muscle cells when insufficient O2
available for aerobic metabolism. Produces less ATP than aerobic metabolism but advantage:
doesn't require oxygen.
● Takes place in the cytoplasm, not in the mitochondria
○ Advantage in muscle cells responsible for quick bursts of speed or strength.
● Like most chemical reactions, glycolysis slows down as its product, pyruvic acid, builds up
○ To reduce this effect, pyruvic acid is converted to lactic acid (fermentation).
● When intensive exercise, such as sprinting, begins, glycolysis is activated. The breakdown of
muscle glycogen supplements the energy produced from ATP and PCr stores
● In theory, the amount of glycogen in the muscles could maintain a sprint for at least 80 seconds
but even the world's best sprinters can only maintain peak speed for about 20 seconds.
Something is limiting them.
Muscle Fatigue
● Muscle fatigue is the inability of a muscle to contract forcefully after prolonged activity even
though the muscle is still receiving stimuli from the nerve.
● The exact cause of muscle fatigue is still not clear but several factors may contribute. The lack
of ATP is not one of them - The [ATP] in fatigued muscle is only slightly lower than in resting
muscle.
● Several contributing factors: (don’t need to know)
○ Conduction failure due to buildup of K+ in T-tubules – leads to depolarization and Na+
channel inactivation.
○ Lactic acid buildup – acidification may alter Ca2+ uptake to and release from SR.
○ Inhibition of cross-bridge cycling due to the build-up of ADP and Pi
○ Fatigue may protect muscle cells from permanent exercise-induced damage due to the
onset of rigor.
Aerobic Respiration- Oxidative Phosphorylation

● The product of glycolysis, pyruvic acid, is converted to acetyl-CoA can be metabolized
aerobically by a pathway known as the Krebs Cycle to provide additional energy.
○ Other products from fats and protein can be converted into energy via Kreb’s cycle.
● Aerobic respiration takes place exclusively in the mitochondria.
● Energy is released in the form of ATP (36 ATP/glucose) and, especially, as high energy
electrons.
○ These high energy electrons are sent to the electron transport system which produces the
vast majority of the ATP.
○ Requires O2 and produces CO2 and H2O as products.
● Used for endurance activities since it can go on for hours.
Summary of ATP Production in Muscle Fibers

Features of White, Fast-Twitch Muscle Fibers

● Also called glycolytic fibers or Type IIB fibers
● Large in diameter
● Light in colour due to reduced myoglobin
● Surrounded by a few capillaries
● Relatively few mitochondria
● High glycogen content
Metabolism in White, Fast-Twitching Fibers

● Muscles with many white muscle fibers are well suited for activities requiring power and speed
for a short duration
● Mainly use glycolysis, which synthesizes ATP quickly
● Rapid cross-bridge cycling results in fast contractions
○ Therefore, also called fast-twitch glycolytic fibers
● Powerful due to large numbers of microfilaments (large diameter)
● Fatigue rapidly
Features of Red, Slow-Twitch Fibers

● Also called slow, oxidative fibers or Type I fibers
● About half the diameter of white fibers
● Dark red in colour due to large quantity of myoglobin
● Surrounded by many capillaries
● Numerous mitochondria
● Low glycogen content
Metabolism in Red, Slow-Twitch Fibers

● Muscles with a high number of red muscle fibers are especially suited for activities requiring
endurance and continuous contraction
● Mainly use Krebs cycle and oxidative phosphorylation for synthesizing ATP
● Cross bridge cycling occurs relatively slowly
○ Therefor called slow-twitch oxidative fibers
● Fatigue resistant, high endurance
Intermediate, Fast-Twitch Fibers

● Also called Type IIa fibers or Fast Oxidative-glycolytic
● As their name suggests, these fibers have properties of both white fast-twitch and red
slow-twitch muscle fibers.
● They are pinkish in color due to their intermediate levels of myoglobin.
● Fast acting myosin and mostly use oxidative phosphorylation to generate ATP, but they have
some glycogen stores.
● Moderately resistant to fatigue.
Muscle Fiber Types

● Three main types of skeletal muscle fibers - identified by: size, speed of contraction,
endurance:
○ Red slow-twitch
○ White fast-twitch
○ Intermediate fast-twitch
● Most muscles: one fiber type predominates; but mix of types gives range of contractile speed
and fatigue resistance.
● All fibers in motor unit are of same type - determined
○ in large part by innervation.
Lecture 8- Smooth Muscle
Smooth Muscle
● Found in places we can’t control
● Found in walls of most hollow organs:
○ Respiratory, digestive, urinary, reproductive, circulatory (not in the capillaries)
○ Not found in heart – heart contains cardiac muscle
● Most smooth muscle organized into sheets of tightly packed fibers
● Most organs contain two layers with fibers oriented at right angles to each other.
Smooth Muscle Layers

● Longitudinal layer: fibers run parallel to the long axis of organ; contraction causes organ to
shorten
● Circular layer: fibers run around circumference of organ; contraction causes lumen of organ to
constrict
● Alternating contraction and relaxation of layers mixes and squeezes substances through lumens
of hollow organs
Smooth vs. Skeletal Muscle Fibers

● Spindle-shaped fibers
● Thinner and shorter than skeletal muscle fibers
● Single, central nucleus
● No striations and no sarcomeres (hence name) but do contain overlapping thick and thin
filaments
● Involuntary control
● Lacks connective tissue sheaths
○ No epimysium or perimysium
○ Contains endomysium only (made by cells themselves)
Innervation of Smooth Muscle

● Innervated by the autonomic nervous system
● Axons contain varicosities (bulbous swellings) instead of neuromuscular junctions
● Varicosities store and release neurotransmitters into a wide synaptic cleft referred to as a
diffuse junction
○ Refer to ppt for diagram and image
Types of Smooth Muscle

● Smooth muscle varies in different organs by:

○ Muscle fiber arrangement and organization
○ Innervation
○ Responsiveness to various stimuli
● All smooth muscles is categorized as either:
○ Unitary (single-unit)
○ Multi-unit
Unitary Smooth Muscle

● Aka visceral muscle
● Found in all hollow organs except heart
● Possess all common characteristics of smooth muscle:
○ Arranged in opposing (longitudinal and circular) sheets
○ Innervated by varicosities
○ Often exhibit spontaneous action potentials
● Electrically coupled by Gap junctions so the cells act in coordinated fashion (like a single unit)
called a syncytium
● Respond to various chemical stimuli
● Can generate its own electrical energy
Myogenic Activity
● Single unit smooth muscle is self-excitable
○ Does not require nervous stimulation for contraction
○ Clusters of specialized smooth muscle cells called pacemaker cells display spontaneous
activity
○ These cells do not contract but cause the syncytium to contract via gap junctions.
○ Membrane potential changes in cyclical manner
● Two types of spontaneous depolarizations
○ Pacemaker potentials
○ Slow-wave potentials
Multiunit Smooth Muscle

● Located in large airways in lungs, large arteries, arrector pili muscles, and iris of the eye
● Very few gap junctions, and spontaneous depolarization is rare
● Similarities with skeletal muscle:
○ Consists of independent muscle fibers
○ Innervated by autonomic nervous system, forming motor units
○ Graded contractions occur in response to neural stimuli that involve recruitment
● Controlled by the autonomic nervous system and hormones
Intermediate Filament-Dense Body Network

● Intermediate filaments: (non-contractile) form lattice-like arrangement that resists tension

● Dense bodies: proteins that anchor filaments to sarcolemma at regular intervals
○ Dense bodies and intermediate filaments harness the pull generated by myosin cross
bridges
● Caveolae - contain many Ca2+ channels to allow rapid influx of extracellular Ca2+
● Gap junctions - electrically connect smooth muscle cells
Mechanism of Smooth Muscle Contraction

● Thick and thin filaments in myofilaments are arranged diagonally in a muscle fiber
● Connected to intermediate filaments and each other by dense bodies (contain protein called
a-actinin)
● Muscle fiber contracts in corkscrew manner (shortens along length)
Arrangement of Thick and Thin Filaments

● Thick filaments are fewer and have myosin heads along entire length
● Ratio of thick to thin filaments (1:13) lower than in skeletal muscle (1:2)
● Myosin heads along entire thick filament but oriented in opposite direction on either face
● Thin filaments move in opposite direction
Mechanism of Smooth Muscle Contraction
Regulation of Contraction by Ca2+

● Don’t need to memorize too much, just realize that it’s different from skeletal muscle
● No T tubules in smooth muscle fibers
● Less elaborate SR
○ SR stores Ca2+ but most calcium used for contraction comes from ECF
● Caveolae contain many voltage-gated Ca2+ channels to allow rapid influx of extracellular Ca2+
● Ca2+ entry triggers Ca2+ release from SR – called Calcium-induced calcium release
● Smooth muscle contains tropomyosin but it does not prevent myosin binding to actin
○ Does not contain troponin
● Ca binds calmodulin
2+
● Activated calmodulin activates myosin light chain kinase

● Activated myosin kinase phosphorylates and activates myosin head
● Crossbridge formation with actin
● Relaxation:
○ Detachment from calmodulin
○ Active transport of Ca2+ into SR and ECF
○ Dephosphorylation of myosin to inactive myosin
E-C Coupling (Smooth Muscle)

1. Calcium ions (Ca2+) enter the cytosol from the ECF via voltage-gated or non-voltage-gated
Ca2+ channels, or from the SR
2. Ca2+ binds to and activates calmodulin.
3. Activated calmodulin phosphorylates and activates the enzyme myosin light chain kinase
(MLCK).
4. The activated kinases catalyze the transfer of phosphate (phosphorylation) to myosin,
activating the myosin ATPase.
5. Activated myosin forms cross bridges with actin of the thin filaments just as in skeletal muscle
and shortening begins.
● Refer to ppt for diagram
Energy Efficiency of Smooth Muscle Contraction

● Slower to contract and relax but maintains contraction for prolonged periods (up to 3 seconds)
with little energy cost
○ Slower ATPases
○ Each crossbridge lasts longer (uses less energy per time)
○ Ca2+ pumped out more slowly
● Most smooth muscles maintain moderate degree of contraction constantly without fatiguing
○ Referred to as smooth muscle tone
● Makes ATP via aerobic respiration pathways
Regulation of Contraction
● Don’t always need AP for calcium release, can use graded potential also
○ Ca regulates contraction
● Neural
○ Neurotransmitter binding causes either or all-or-none action potential
■ Results in increases in Ca2+ concentration in sarcoplasm
■ Response depends on the neurotransmitter released and the type of receptor

molecules
● One neurotransmitter can have a stimulatory effect on smooth muscle in
one organ, but an inhibitory effect in a different organ
● Hormones and local chemicals
○ Some smooth muscle cells have no nerve supply
■ Depolarize spontaneously or in response to chemical signals (e.g. hormones) that
bind to GPCRs
■ Other chemical factors including histamine, high CO2, low pH, low oxygen
enhance or inhibit Ca2+ entry into the cytoplasm.
○ Some smooth muscles respond to both neural and chemical stimuli
Special Features of Smooth Muscle Contraction

● Stress-relaxation response: spontaneously contracts when stretched BUT only briefly, then
adapts to new length
○ Retains ability to contract on demand
○ Enables organs such as stomach and bladder to temporarily store contents
○ Length and tension changes
● Can contract when between half and twice its resting length
○ Allows organ to have huge volume changes without becoming flabby when relaxed
Summary of Smooth Muscle Contraction

● Slow, synchronized contractions
○ Rate and intensity of contraction may be modified by neural and chemical stimuli
● Contraction in smooth muscle is similar to skeletal muscle contraction in following ways:
○ Actin and myosin interact by sliding filament mechanism
○ Final trigger is increased intracellular Ca2+ level
○ ATP energizes sliding process
○ Contraction stops when Ca2+ is no longer available
● Refer to ppt for diagram
Comparison of Skeletal and Smooth Muscle


Anatomy and Physiology Notes

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Anatomy and Physiology Notes

Human Anatomy and Physiology Notes

Lecture 1- Introduction, Structure, Function and Cell Theory

Complementarity of Structure and Function

Levels of Structural Organization

Organ Systems of the Body

Integumentary ● Forms external body covering + protects deep tissues from

Skeletal System ● Protects + supports organs

Muscular System ● Allows manipulation of the environment, locomotion and facial

Nervous System ● Fast- acting control system of body

Endocrine System ● Glands secrete hormones which regulate numerous body

Respiratory ● Keeps blood supplied with oxygen + removed carbon dioxide

System ● Gas exchange occurs through walls of air sacs in lungs

Digestive System ● Breaks down food to be absorbed by blood

Urinary System ● Eliminates nitrogenous wastes from body

Reproductive ● Male and female

Parts of a Cell: Structure and Function

Organelles In The Cytoplasm

Organelle Structure Function

Mitochondria ● Rodlike, double-membrane ● Site of ATP synthesis

Ribosomes ● Consist of 2 subunits, each ● Synthesize proteins for various cell

Rough ER ● Outer surface is covered in ● External face synthesizes proteins

Smooth ER ● Is continuous with rough ER ● Does not make proteins

Golgi ● Stack of flattened membranes and ● Packages, modifies and segregates

Lysosomes ● Membranous sacs containing acid ● Sites of intracellular digestion

Microtubules ● Cylindrical ● Support cell and gives it shape

Microfilament ● Fine filaments composed of ● Involved in muscle contraction and

Intermediate ● Protein fibers ● Make up the stable cytoskeleton

Centrioles ● Paired cylindrical bodies ● Organize microtubule network during

Inclusions ● Varied ● Are storage for nutrients, wastes and

Cilia ● Short, cell surface projections ● Their coordinated movement

Flagellum ● Like cilium, but longer ● Propels the cell in its

Microvilli ● Tubular extensions of plasma ● Increase surface area for

Parts Of The Nucleus

Nucleus ● Largest organelle ● Control center of cell

Nuclear ● Double membrane ● Separates nucleoplasm from

Nucleolus ● Dense, spherical bodies ● Site of ribosome subunit synthesis

Chromatin ● Granular and threadlike ● Fundamental units are nucleosomes

Lecture 2- Types of Tissues

Processes Needed to Make Us Multicellular

Overview of 4 Basic Tissues

Nervous Tissue Internal communication Brain, spinal cord, nerves

Muscle Tissue Contracts to cause movement Skeletal muscles

Preparing Human Tissues for Microscopy

Type 1- Epithelial Tissue

Special Characteristics of Epithelium

■ Desmosomes- prevents cells from pulling apart in tissues subjected to

○ Columnar: tall, column-like

Types of Simple Epithelium

*There are two types of simple squamous epithelium based on location;

Types of Stratified Epithelium

Stratified Cuboidal -Very rare -Found in some sweat

Stratified Columnar -Very rare in the body -Transitional areas

Formation of Multicellular Exocrine & Endocrine Glands

Formation of Multicellular Endocrine & Exocrine Glands

Type 2- Connective Tissue

Overview of Classes of Connective Tissue

Common Characteristics of Connective Tissue

Structural Elements of Connective Tissue

■ Hematopoietic- stem cells in bone marrow

Types of Connective Tissue Proper

Types of Connective Tissue Cartilage