GAS EXCHANGE

Cellular respiration External respiration

● Oxidative process within cells ● Exchange of O2 and CO2 between organism

● Glycolysis, oxidative phosphorylation, etc. and environment

● Uptake of molecular O2 from the environment and the discharge of CO2 to the environment
● Works with Partial pressure gradients
○ The pressure exerted by a particular gas in a mixture of gases
○ Net movement of gas at an exchange surface:
■ A gas always undergoes net diffusion from a region of high partial pressure to a
region of lower partial pressure
● Conditions for gas exchange vary depending on respiratory medium (source of O2) - air or water
● Water has lower O2 content, greater density, and greater viscosity so aquatic animals must expend more
energy for gas exchange
● Respiratory surfaces​ - part of animal’s body where gas exchange occurs (for diffusion to be effective):
○ Moist​ (b/c PM of cells must be in contact w/ aqueous solution)
○ Thin​ (path for diffusion is short)
○ Large surface area​ (area for diffusion is large)

Respiratory Organs
1.) Cutaneous respiration​ (direct diffusion)
● Relatively simple animals - sponges, cnidarians, flatworms
● Every cell in the body is close enough to external env. That gases can diffuse quickly between any
cell and the env.
● Thin, moist epithelium (or their PM integument) constitutes a respiratory organ
● Earthworms and some amphibians
○ Skin serves as a respiratory organ
○ Dense network of capillaries just below the skin facilitates the exchange of gases between
the circulatory system and env.

2.) Gills​ or ​Branchia​ (external or internal)

● External gills - Sea star, Marine worms
Internal gills - Crustaceans, fish (but gills still exposed to external
env.)
● Gills - outfoldings of the body surface that are suspended in the
water
● Distribution of gills over the body can vary considerably
● Often have a total surface area much greater than that of the rest of the body’s exterior
● Ventilation
○ Movement of respiratory medium over respiratory surface
○ Maintains the partial pressure gradients of O2 and CO2 across the gill that are needed for gas
exchange
○ Two ways of ventilation:
■ Move gills through water
● E.g. crayfish and lobsters have paddle-like appendages that drive currents of water
over gills
 ■ Move water over gills
● E.g, mussels and clams move water with cilia
○ In fishes
■ Motion of swimming or coordinated movements of mouth and gill covers to ventilate their
gills
■ Current of H2O enters mouth of fish → H2O passes through slits in the pharynx → H2O
flows over gills and exits body

● Countercurrent exchange​ (in fishes - maximizes gas exchange)

○ Exchange of a substance or heat between 2 fluids flowing in opposite directions - blood and water
○ Blood flows in opposite direction of water passing over gills
○ At each point in its travel, blood is less saturated w/ O2 than the water it meets
○ As blood enters gill capillary, it encounters water that is completing its passage through the gill
(water has higher Partial Pressure of O2 than blood)
○ O2 transfers from water to blood (diffusion) along capillaries
3.) Tracheal Systems​ in Insects (branching system of tubes)
● (most terrestrial animals) respiratory surfaces are enclosed within the body, exposed to the
atmosphere only through narrow tubes
● Tracheal system​ is a network of air tubes that branch ​throughout the body
● Largest tubes - Tracheae
○ Open to the outside
● Tips of finest branches - moist epithelial lining enables gas exchange by diffusion
● No need for open circulatory system b/c tracheal system brings air w/in a very short distance
● When insect is in flight, more O2 is needed
○ Cycles of flight muscle contraction and relaxation pump air rapidly through the tracheal
system
○ Pumping improves ventilation → brings ample O2 to densely packed mitochondria that
support the high metabollic rate of flight muscle
 4.) Lungs
● Localized respiratory organs​ (do not branch throughout body)
● Infolding of body surface, typically divided into numerous pocket
● Not in direct contact with all other parts of the body, so ia ​circulatory system​ transports gases
between the lungs and the rest of the body
● Present in spiders, land snails, vertebrates
● Amphibians rely more on ​diffusion across external body surfaces​ (skin) to carry out gas
exchange, and only have a relatively small lungs
● Most reptiles, birds, and mammals rely entirely on lungs for gas exchange
● Turtles
○ Supplement long breathing with gas exchange across moist epithelial surfaces continuous
with their mouth or anus

Mammalian Respiratory System (lungs)

● Branching ducts convey air to lungs located in ​thoracic cavity​ enclosed by ​ribs​ and ​diaphragm
● Air enters through
nostrils and then
nasal cavity
○ Air filtered
by hairs,
warmed,
humidified
and
sampled
for odors
● Nasal cavity leads
to ​pharynx​ -
intersection where
paths for food and
air cross
● Larynx
○ Moves
upward
and tips
the
epiglottis over glottis to cover trachea when food is ​present
○ Upper part of respiratory tract
○ Vocal folds/cords are present here as a pair of elastic bands of muscle by which air passes
● From larynx, air passes into ​Trachea
○ Like the trunk of an inverted tree
● Trachea branches into 2 ​Bronchi​, one in each lunch
● Bronchi branch further into finer tubes called ​bronchioles
● Epithelium lining major branches of this system is covered by ​cilia​ and ​thin film of mucus
● Mucus
○ Traps dust, pollen, and other particulate contaminants
○ Cilia​ move mucus up to the pharynx (mucus escalator) for cleaning of system
● Alveoli
○ Air sacs clustered at the tips of the tiniest bronchioles
○ Where gas exchange occurs
 ○ Oxygen in air entering alveoli dissolves in the moist film lining their inner surfaces and rapidly
diffuses across the epithelium into a web of capillaries that surrounds each alveolus
○ Net diffusion of CO2 occurs in opposite direction: cappiliaries → epithelium of alveolus → air space
○ Highly susceptible to contamination b/c it has no cilia
■ It has WBC instead that engulf foreign particles
○ Surfactant​ (surface-active agent) and phospholipids
■ Covers the alveoli to reduce surface tension
■ Produced by air sacs

Breathing​ or ​Ventilation of the Lungs

● Ventilation maintains high O2 and low CO2 concentrations
at gas exchange surface
● Breathing
○ Process that ventilates lungs
○ Alternate inhalation and exhalation of air

1.) Amphibians​ (frogs)

● Ventilates its lungs by ​positive pressure breathing

○ Inflates lungs with ​forced airflow

● Inhalation begins when
1.) Muscles lower floor of amphibian’s oral cavity
2.) This draws air through nostrils
3.) Nostrils and mouth close; floor of oral cavity rises
4.) Air is forced (​pushed​) down the trachea
5.) Exhalation follows as air is expelled by elastic recoil
of lungs and by compression of muscular body wall
● Male courting puff themselves up by taking in air
several times without any release

2.) Birds
● Air is passed over gas exchange surface in ​on
direction​ only
○ Incoming fresh air does not mix with air that
has already carried out gas exchange
● Air sacs on either side of lungs act as bellows that
direct air flow through lungs
● Parabronchi​ (tiny channels within lungs)
○ Serve as sites of gas exchange
● Passage of air through entire system of bird requires
two cycles of inhalation and exhalation

3.) Mammals
● Breathing is like a syringe’s plunger being pulled
back
● Negative pressure breathing​ - pulling air into lungs
● Breathing:
○ Uses muscle contraction to actively expand
thoracic cavity
 ○ This lowers pressure in lungs vs. air outside body
○ Air flows/rushes through nostrils/mouth to the alveoli (b/c air goes from high to low pressure
region)
● Diaphragm
○ Sheet of skeletal muscle that forms the bottom wall of the cavity
○ Contracting rib muscles pull ribs upwards and sternum outward, but diaphragm contracts
(moves downward) and expands thoracic cavity
■ This descending movement of diaphragm is like a plunger drawn out of a syringe
● Inhalation is always
active and requires
work; exhalation is
usually passive
○ Muscles
controlling
thoracic cavity
relax
○ Volume of cavity
is reduced
○ Increased
pressure in
alveoli forces air
up the breathing
tubes and out of
the body
● Thoracic Cavity
○ Surrounds the lungs
○ Double-membrane in between the lungs and thoracic cavity
○ Inner layer adheres to outside of lungs; outer layer adheres to inside wall of thoracic cavity
■ Thin space filled with fluid separates the two layers
■ Surface tension within fluid causes two layers to stick together
● Tidal volume
○ Volume of air inhaled and exhaled with each breath
○ Averages about 500 mL in resting humans
● Vital capacity
○ Tidal volume during maximal inhalation and exhalation
○ 3.4 L (college women) and 4.8 L (college men)
● Residual volume
○ Air that remains after a forced exhalation
○ Increases as people age, but vital capacity decreases
● NOTE​:
○ Inhalation occurs through the same airway as exhalation
○ Inhalation mixes fresh air with oxygen-depleted residual air
○ Thus, max partial pressure of O2 is always less in the alveoli than in the atmosphere
○ Also less in mammals than for birds b/c birds have a unidirectional flow of air
■ Reason why mammals function less well than birds in high altitude

Control of Breathing in Humans

● Breathing is regulated by involuntary mechanisms
○ Ensure that gas exchange is coordinated with blood circulation and metabolic demand
● Medulla oblongata
○ Near the base of the brain
○ Neurons mainly responsible for regulating breathing are in here
○ Neural circuits in medulla form a pair of ​breathing control centers​ that establish the breathing
rhythm
○ During inhalation, sensors that detect stretching of he lung tissue send nerve impulses to the
control circuits in the medulla
■ Inhibits further inhalation/lung overexpansion
○ Regulates breathing by:
■ Using pH of the fluid in which it is bathed as an indicator of blood ​CO2 concentration
■ Blood CO2 is the ​main determinant of the pH of cerebrospinal fluid​ (fluid surrounding
brain and spinal cord)
■ CO2 diffuses from blood to cerebrospinal fluid ​where it ​reacts with water​ and forms
carbonic acid (H2CO3)
■ H2CO3 quickly dissociates into ​bicarbonate ion (HCO3-) and Hydrogen Ion (H+)
○ Increased metabolism ​raises CO2 concentration in the blood and cerebrospinal fluid​ = lower
pH (more acidic)
■ Sensors in medulla and major blood vessels detect this pH change
■ Medulla’s control circuit increase the depth and rate of breathing until excess CO2 is
eliminated and pH returns to normal
○ Blood O2 level - little effect on breathing control centers unless O2 Level drops very low (e.g. in
high altitudes)
■ O2 sensors in aorta and carotid arteries in the neck send signals to breathing control
centers
■ Increase in breathing rate as response
■ Regulation of breathing is modulated by additional neural circuits - pons (next to medulla)
○ During high metabolic activity
■ Enhancement of O2 uptake and CO2 removal
■ (More air needed)
Respiratory Pigments (bind and transport gases)

Coordination of Circulation and Gas Exchange

1.) Inhalation: fresh air mixes with air remaining in lungs
2.) Mixture in alveoli has higher PO2 than blood flowing through alveolar capillaries
Net diffusion of O2 down its partial pressure gradients from air → alveoli → blood
Presence of a PCO2 in alveoli (higher in capillaries vs. in air) drives diffusion of CO2 from blood →
air
3.) After blood leaves lungs in pulmonary veins
PO2 and PCO2 match values for air in alveoli
Blood pumped through systemic circuit after returning to the heart
4.) In systemic capillaries​, ​gradients of partial pressure favor net diffusion of O2 out of blood and CO2
into blood
(gradient exists b/c of cellular respiration in mitochondria of cells near capillaries)
5.) O2 unloads and CO2 loaded; blood is returned to the heart and pumped into lungs again
6.) In lungs, ​exchange occurs across the alveolar capillaries resulting in exhaled air enriched in CO2
and partially depleted O2

Respiratory Pigments
● Low solubility of O2 in water (and thus in blood) poses a problem for animals that rely on circulatory
system to deliver O2
● Animals transport most of their O2 bound to proteins - ​respiratory pigments
○ Circulate w/ blood or ​hemolymph
○ Often contained within specialized cells
○ Pigments greatly increase the amt of O2 that can be carried in circulatory fluid
○ Presence of respiratory pigment reduces cardiac output necessary for O2 to transport to a
manageable 12.5 L of blood per minute (instead of 555 L)
○ Pigment has a distinctive color and consists of a metal bound to a protein
○ Two types:
1.) Hemocyanin
● Found in arthropods and molluscs
● Copper as oxygen-binding component
2.) Hemoglobin
● In invertebrates and all vertebrates
● Contained in erthrocytes
● Iron atom bound to heme group
● Each iron atom binds one molecule of O2, and hemoglobin can carry 4 O2
molecules
● Hemoglobin binds O2
in lungs or gills and
unload it elsewhere in
the body
 ● O2 diffuses into
pulmonary capillaries
● Most O2 combines w/
hemoglobin in RBC to form
oxyhemoglobin
● CO2 diffuses out of
pulmonary capillaries
● Most CO2 transported in
form of bicarbonate ion
● Some CO2 combine with
hemoglobin to form
carbaminohemoglobin

Dissociation Curve for Hemoglobin

● Cooperative oxygen binding and release is evident in the ​dissociation curve​ for hemoglobin
● Change in Partial pressure of O2 causes hemoglobin to load or unload a substantial amount of O2
● When cells/tissues are working harder (e.g. exercise), PO2 dips in their vicinity as more O2 is consumed in
cellular respiration
 ○ Slight drop in PO2 causes a relatively large increase in amt of O2 the blood unloads

Bohr Shift
● Hemoglobin is efficient at delivering O2 to tissues actively consuming O2
● Increased efficiency results from CO2
production, not O2 consumption
● Bohr shift
○ Effect when low pH decreases the
affinity of hemoglobin for O2
○ As tissues consume O2 in cell
respiration, they also produce CO2
which forms carbonic acid when reacts
with water
○ Where CO2 production is greater,
hemoglobin releases more O2 which
can be used to support more cellular
respiration
● Hemoglobin also assists in buffering the blood
(preventing harmful changes in pH)

Carbon Dioxide Transport

● Only 7% of CO2 released by respiring cells is
transported in sol’n in blood plasma
● Rest diffuses from plasma into erthrocytes and
reacts with water = H2CO3
● H2CO3 dissociates into H+ and HCO3=
● H+ binds to hemoglobin and other proteins
(minimizing change in blood pH)
● Most HCO3- diffuses out of erthrocytes;
transported to lungs in plasma
● Remaining HCO3- (5% of CO2) binds to
hemoglobin and transported in erthrocytes
● Blood flows through lungs, relative partial
pressure of CO2 favor net diffusion of CO2 out
of blood
● As CO2 diffuses into alveoli, amt of CO2 in
blood decreases
● Decrease shifts chemical equilibrium in favor of
conversion of HCO3- to CO2, enabling net
diffusion of CO2 into alveoli
● Overall, PCO2 gradient is sufficient to drive
about 15% reduction in PCO2 during passage
of blood through lungs