Miller/Tupper: Zoology 11e

Instructor’s Manual

26
Chapter Summary

An internal transport system is useful because it reduces the distances over which
substances must diffuse—substances can be brought close to a target point via the blood
and then diffuse outward from that point. Internal transport systems of invertebrates
range from simple fluid filled cavities to closed systems, analogous to vertebrate systems.
Some invertebrate systems are outlined below:
● Sponges circulate water throughout their bodies as they filter feed
● Cnidarians and acoelomate worms have gastrovascular cavities that accomplish
transport as well as feeding functions
○ internal branching within the cavity ensures short diffusion distances
○ body movements help distribute materials within the gastrovascular cavity
● Pseudocoelomates rely on coelomic fluid to accomplish respiratory functions, as
do echinoderms, ectoprocts and sipunculids
● Molluscs have a circulatory system with a unidirectional heart and blood vessels
○ the molluscan system is an open circulatory system, where the heart
pumps hemolymph out to open areas (sinuses) where the cells are bathed
in blood
○ in cephalopods there is a closed circulatory system where blood remains
within a system of vessels throughout the body
● Annelids have a closed circulatory system, while arthropods have an open system.

The circulatory fluid in open systems is known as hemolymph. It has a high volume and
a slow circulation. Hemolymph may also assist in ecdysis and locomotion by providing
hydraulic pressure.

Hemocytes are blood cells carrying a respiratory pigment (also called erythrocytes, as in
red blood cells).
Other cells in circulatory fluid function in phagocytosis and clotting:
● insects have cells used in a nonspecific defense mechanism called encapsulation
● molluscs have hemocytes that function in nacrezation (pearl formation)

All vertebrates have closed circulatory systems; blood is always in arteries, arterioles,
capillaries, venules, veins, or within the chambers of the heart. Exchange between blood
and extracellular fluids occurs only across the walls of capillaries in vertebrates.
Vertebrate blood shares many functions with invertebrate blood, but it may function in
thermoregulation, as well.

Blood is a connective tissue made up of a fluid matrix and cellular elements.

1. Plasma is primarily composed of water, and also contains proteins, ions,
hormones, nutrients, and wastes
a. Serum is plasma without the clotting proteins
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 2. The cellular elements of blood are called formed elements (red blood cells, white
blood cells, and platelets).
a. Erythrocytes (red blood cells).
i. vary among the vertebrate classes; most are nucleated, but there
are anucleate biconcave discs in most mammals
ii. contain hemoglobin, which binds to oxygen for transport from
lungs to other tissues
iii. in mammals, red blood cells are the most numerous formed
elements in the blood, and usually number several million cells per
cubic centimeter
b. Leukocytes (white blood cells). These cells function in immunity, and are
all produced from stem cells in the bone marrow by hematopoiesis
i. Neutrophils are a type of phagocytic granulocyte and are the most
common leukocytes. Neutrophils are attracted to an inflammation
site by histamines.
ii. Basophiles are the least numerous leukocytes. Basophiles react
with foreign substances (antigens) and produce histamines and
heparin to increase circulation in an area and allow for easier
invasion of the site by Neutrophils and Macrophages.
iii. Eosinophiles are leukocytes that ingest foreign proteins and
immune complexes. They help counteract the effects of certain
chemicals during allergic reactions. They are phagocytic cells.
iv. Lymphocytes are agranulocytes; they may be distinguished as B
and T cells, both of which are active in the immune response.
1. B cells differentiate into plasma cells, which produce
antibodies
2. Differentiated T cells are called cytotoxic cells and are
programmed to actively destroy cells that are displaying
specific proteins
3. B and T cells may also remain in the circulatory system as
memory cells, and in this way, they confer long-term
immunity to an organism
c. Thrombocytes (platelets) function in blood clotting. Thrombocytes begin
their development as megakaryocyte cells and, when mature, break apart
into cellular fragments that travel through the animal's circulatory system
as thrombocytes.

Arteries and veins carry blood.

● Arteries are characterized as elastic vessels that carry blood away from the heart
○ transport blood under high pressure
● Veins are larger, less elastic vessels that carry blood to the heart
○ veins carry blood that is very low in pressure
● Capillaries are the connections between arterioles and venules and are the site of
exchange of materials with adjacent tissues. Capillaries are the most ubiquitous
of the blood vessels and provide a huge surface area for gas exchange.

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written consent of McGraw-Hill Education.
There is a trend toward greater efficiency visible if one compares the hearts of a fish,
amphibian, reptile, bird and mammal. The basic features of each system are outlined
below.

● Bony fish have 2 pumping chambers of the heart: one atrium and one ventricle
and two collecting chambers: the sinus venosus and conus arteriosus. Blood
flows from the sinus venosus to the atrium, then to the ventricle, and then from
the conus arteriosus to the gills in a single circuit. The disadvantage of this
system is the low pressure flow through the gills supports only a low metabolic
rate.
● Amphibians and most reptiles have a double circuit—blood flows to the lungs and
then returns to the heart before systemic circulation. This overcomes the slow-
flow problem, but the heart is incompletely divided to allow the shunt to the lungs
to close during diving. However, the heart of crocodilians has completely
separated ventricles and atria, as do birds and mammals—this facilitates a
complete separation of the two circuits, maintains high blood pressure and rapid
oxygen supply for an increased metabolic rate. A double circuit occurs when
blood from the lungs is sent directly to the heart, and blood from the tissues also
goes to the heart—these systems are the pulmonary and systemic circulations,
respectively. The ventricles of the turtle heart are incompletely separated, thus
allowing blood to be shunted away from the inactive lungs when the turtle is
diving or withdrawn into its shell and toward more vital systematic organs.
● The human heart pumps the entire blood volume (5 liters) every minute. The
heart is composed of cardiac muscle (the myocardium) and a fibrous outer
epicardium.
The beating of the heart is controlled intrinsically in a multi-step process:
1. the heartbeat is initiated by the pacemaker cells of the sinoatrial node (SA node)
which lies in the right atrium
2. the SA node signals the AV node
a. electrical activity at the SA and AV nodes elicits atrial contraction
3. the AV node signals the bundle of His which carries a nervous impulse through
the interventricular septum to the apex of the heart
4. the bundle of His signals the Purkinje fibers
5. Purkinje fibers promotes contraction of the ventricles and send blood upward and
out of the heart through the aorta and pulmonary trunk

Blood pressure reflects ventricular contraction, the force the blood exerts against the
vessels, measured in mmHg; it is typically 120/80 for systolic pressure/diastolic pressure.
Valves within the heart prevent the backflow of blood during ventricular systole and
diastole and are the cause of the characteristic "heart sounds" that may be heard with a
stethoscope.
The lymphatic system is primarily involved in draining extracellular fluid and in
immunity. The lymphatic system is composed of vessels, lymph (the fluid within the
vessels), as well as lymphatic organs such as the lymph nodes, spleen, and the tonsils.
Modified lymphatic vessels called lacteals transport lipids that have been absorbed by the
small intestine. The thymus gland is often considered a part of the lymphatic system in

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written consent of McGraw-Hill Education.
mammals as it is the site of antibody production in newborns and T cell differentiation in
adults. The bursa of fabricius is the site of B cell maturation in birds and acts as a part of
their lymphatic system.

There are 5 respiratory mechanisms of invertebrates:

1. diffusion - integumentary exchange via capillary networks under the skin works
with relatively small animals (such as flatworms)
2. tracheae - tracheae are an internal system of branching tubes that brings air
directly to each cell in the body. The system, formed by an invagination of the
arthropod cuticle, is efficient in terrestrial organisms (insects and some other
arthropods).
3. cutaneous exchange
4. gills - gills are very efficient in aquatic ecosystems—gills evolved from large or
small skin projections with blood vessels inside of the projections
5. lungs - book lungs are sheets of internal lamellae that are exposed to air on the
outside and filled with blood on the inside. Pulmonate snails respire via a lung
(internal cavity) derived from the mantle cavity. Some of the more primitive,
pulmonate, aquatic snails surface and take in air through their pneumostome, then
close the pneumostome and submerge.

Vertebrates respire by some combination of body surface, gills and lungs.

1. bimodal breathing occurs when an animal can respire both on land and in the
water (examples include salamanders and some fishes)
a. facilitated the evolutionary transition to land
2. trimodal breathing involves the gills, the lungs, and the skin
a. skin
i. frogs and toads have a vascular network below the skin and keep
mucus on top of the skin to keep it moist, facilitating exchange
ii. cutaneous respiration is very efficient in amphibians
b. gills may be internal or external and gas exchange (oxygen in and carbon
dioxide out) is very efficient due to countercurrent flow
c. lungs are internal structures
i. shared evolutionary history with the swim bladder of fishes
ii. lungs of primitive fishes are dorsal to the digestive tract
iii. lungs of modern teleost fishes and other vertebrates are ventral to
the digestive tract
iv. lungs of modern vertebrates vary in structure as indicated below

Vertebrate lungs vary in structure:

● birds are a flow-through system wherein air is conducted through the lungs in a
single direction, and avian lungs are particularly efficient (90% of oxygen in air is
removed). The system of air sacs prevents the mixing of incoming air with ‘old’
air. Countercurrent flow also aids in efficiency. Exchanges in lungs involve the

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written consent of McGraw-Hill Education.
 uptake of oxygen by diffusion in the alveoli, and the loss of carbon dioxide also
via diffusion. Passageways for air flow through bird lungs are called parabronchi.
● Amphibians and some reptiles employ a positive pressure pumping mechanism to
push air into lungs. Most reptiles, and all birds and mammals, utilize a negative
pressure system to bring air into lungs—they inhale by suction. Negative
pressure for intake of air is accomplished by contraction of muscles surrounding
the ribs and contraction of abdominal muscles in reptiles and birds. Mammals
utilize a muscular diaphragm to create a negative pressure within the thoracic
cavity. This respiratory system is not as efficient as in birds—only 25% of the
oxygen in air is removed.
● The human respiratory system includes both portions of the nasal cavity and the
oral cavity, in communication with the pharynx, larynx, and trachea. The alveoli
(the tremendously large surface area of the lungs) are the sites of gas exchange in
the lungs of mammals and birds. The mammalian diaphragm and intercostal
muscles facilitate inspiration and expiration.

Some vertebrates (reptiles, birds, and mammals, for example) have high concentrations of
oxygen-binding myoglobin proteins associated with their skeletal muscles. This protein
allows the animals to carry out extensive muscular activity for extended periods of time
under conditions such as diving or flying. Some vertebrates also possess a higher volume
of blood within their bodies, and this allows them to carry out activities for extended
periods of time under anaerobic conditions.

As animal size increases diffusion cannot accommodate the metabolic demand for
oxygen. Therefore, respiratory pigments are used to transport gases (O2 and CO2) to and
from the areas they are needed and produced, respectively. Circulatory systems are used
to carry the pigments and gases. These pigments, organic compounds that have either
metallic copper or iron that binds oxygen, are found from the Mollusca through the
Chordata. Examples of respiratory pigments include:
● iron-containing hemoglobin
● iron-containing hemerythrin (Sipuncula, Priapulida, Annelida)
● iron-containing chlorocruorin (a green-colored pigment found in some annelids)
● copper-containing hemocyanin (molluscs and crustaceans, gives blood a bluish
color)

Respiratory pigments reversibly bind oxygen. For example, in mammals:

● blood in alveolar capillaries interacts with air having a partial pressure of oxygen
(PO2) of about 100 mmHg
● hemoglobin is saturated with oxygen while in the lungs
● blood circulates into tissue capillaries, where tissue PO2 levels are 40 mmHg
● the affinity of hemoglobin for oxygen decreases at these lower PO2 levels:
○ hemoglobin releases 25% of the oxygen
○ oxygen diffuses from the red blood cells into tissues where O2 supports
cellular metabolism
● blood is transported back to the lungs where it picks up fresh O2

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Loading and unloading relationships are shown in oxyhemoglobin dissociation curves
(below). Note that hemoglobin is responsive to metabolic conditions at tissues. More
oxygen will be released under the following conditions:
● increased metabolism
● decrease in pH
● increase in body temperature

Respiratory pigments also reversibly bind carbon dioxide. For example, in mammals:
● CO2 is at higher concentrations at tissues and diffuses into the blood
● some CO2 is transported by combining with hemoglobin
● most CO2 is transported in the form of bicarbonate ions
● the conversion of CO2 to bicarbonate (below) is reversed at the lungs

CO2 + H2O ⇋ H2CO3 ⇋ H+ + HCO3 −

Respiratory pigments arose early in animal evolution as animal size began to increase.
Hemoglobin can be traced back 800 million years. Hemoglobin also gave rise to
myoglobin (the oxygen-storing pigment in muscle) through gene duplication. Other
respiratory pigments may have evolved independently in different animal groups.

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written consent of McGraw-Hill Education.