Student Workbook

Comparative Animal Physiology

PCB4723
Dr. Wayne A. Bennett

COMPARATIVE ANIMAL PHYSIOLOGY

CLASS SYLLABUS
SPRING 2015
OVERVIEW
This course will provide you with a thorough overview of animal physiological processes. We
will be taking a comparative approach to understanding physiological similarities and differences
across multiple organizational levels. Animal physiology comprises a massive body of
information collected over nearly 3000 years of human study. So be prepared to cover large
amounts of material during each lecture session, and to review the material on your own carefully
and frequently.
The following suggestions will greatly enhance your in-class performance:
1)
2)
3)
4)

Attend class regularly

Ask questions
Read the book or other outside materials on the topics covered
STUDY, STUDY, and STUDY

DETAILS AND GENERAL STUFF

INSTRUCTOR:
Dr. Wayne A. Bennett, Professor of Vertebrate Physiology
Office hours: Tuesday and Thursday 08:00-10:00; or by appointment
UWF office: 58/62-H
Phone:
474-3362
E-mail:
wbennett@uwf.edu
MEETING TIME:
Tuesday and Thursday 2:30 to 3:45
TEXT BOOK:
Comparative Animal Physiology, by Philip C. Withers

STUDENT LEARNING OUTCOMES (SLO)

UPON COMPLETION OF THIS COURSE, STUDENTS SHOULD BE ABLE TO:
1) Identify similarities and differences in the functional morphology and physiology between
major animal groups.
2) Define the Fry Paradigm and explain how relationships between animals and their abiotic
environment alter physiology to determine behavior using specific references to: oxygen
uptake, temperature regulation, ion balance, sensory physiology, and circulation.
3) Describe key morphological and physiological attributes that result in differing sensory,
communication and cognitive abilities among/between vertebrate and invertebrate groups.
4) Describe the propriety, need and benefits of basic and applied comparative physiology
research.
5) Solve basic biophysical equations that define major physiological attributes of animals
including: oxygen uptake, temperature regulation, ion balance, sensory physiology, and
circulation.
Additional detailed SLOs are listed in the unit Module outlines

ATTENDANCE:
You should make every effort to attend class regularly. Students who have poor
attendance records fail this class spectacularly and without exception!
CLASSROOM MATERIALS AND HANDOUTS:
Handouts will come to you via group e-mail. The information can be opened by clicking
on the link, but in some cases you will need to copy the link and paste it into the address
line on your internet browser.
READING ASSIGNMENTS:
The text book compliments much of the lecture material but is not identical to it. I also
often include material that isnt in the book that I think you should know. Reading the
book or other outside material can aid you in grasping concepts that you did not clearly
understand during class or filling in areas that we will not have time to cover in depth.

IN-CLASS OPPORTUNITIES
OPPORTUNITIES:
In-class assessments are an opportunity to show me what you have learned. Here's how it
works.
1)
2)
3)

We will have three test opportunities.

THERE WILL BE NO MAKE-UP TESTS!
Test will consist of 40 to 50 multiple choice and 10 fill in the blank
questions.

Exam questions will be taken from lecture topics and will be approximately 60% recall
(What is the typical transmembrane potential?), 30% application (If sodium concentration
inside the cell increases, what happens to the transmembrane charge?), and 10% analysis
(How might a decrease in transmembrane charge affect animal response times?)
questions.

TEST SCHEDULE:
Thursday February 5th at 2:30-3:45
Thursday March 19th at 2:30-3:45
Thursday April 23th at 2:30-3:45
SPECIAL ACCOMMODATIONS:
If you require special in-class accommodations or test-taking arrangements due to
physical or perceptual limitations, please contact the UWF Student Disabilities Resource
Center 474-2387.
GRADING SYSTEM:
Three in-class exams (33% each)

GRADING SCALE:
Exams will be will be adjusted to the following scale:
90 - 100:
80 - 89:
70 - 79:
60 - 69:
< 60:

A
B
C
D
F

GRADES OF INCOMPLETE (I) OR WITHDRAW (W)

If you have a question regarding UWF policies for assignment of grades of 'W' or 'I', please visit:
UWF Withdrawal Policy
http://uwf.edu/registrar/withdrawal.cfm
UWF Incomplete Grade Policy
http://uwf.edu/registrar/Incomplete%20Grade%20--%20Assignment%20Report.pdf

CLASS TOPICS

CHAPTER

SYLLABUS & INTRODUCTION TO COMPARATIVE PHYSIOLOGY

1&2

CELL & MEMBRANE PHYSIOLOGY

NEURONS AND THEIR FUNCTION

NERVOUS SYSTEMS

GENERAL SENSES

CHEMORECEPTION

VISION

INTRODUCTION TO METABOLISM

BODY SIZE & METABOLISM

TEMPERATURE & METABOLISM

4&5

SKELETONS & SUPPORT

10

MOVEMENT WITHOUT MUSCLE

10

MOVEMENT WITH MUSCLE

10

DIRECT WATER BALANCE

16

ORGANS OF EXCRETION

16

DIFFUSION & RESPIRATION

12

RESPIRATORY SYSTEMS

12&13

VENTILATION SYSTEMS

12&13

INVERTEBRATE CIRCULATORY SYSTEMS

14

VERTEBRATE CIRCULATORY SYSTEMS

14

STUDY MODULE I

UNDERSTANDING ANIMAL NERVES AND NERVOUS SYSTEMS

STUDY GUIDES
Comparative Animal Physiology

Dr. Wayne A. Bennett, Professor of Animal Physiology

ON THE IMPORTANCE OF STUDY SKILLS AND HOW TO BE A STUDENT OF

BIOLOGY, INSTEAD OF JUST PLAYING AT IT
OR
THE BEST 36 MINUTES OF YOUR LIFE

Since the days when crowds gathered to listen to open-air oratories by Aristotle on the
diversity and function of animal life, students of biology have been bound by one
immutable rule:

Every Hour Spent in Instruction, Requires Two Hours of Outside Study

Ten Facts You Should Know:
1.

Historically 2 of 5 Comparative Animal Physiology students drop or fail to attain a passing grade

2.

In 1970, the average college student studied 25 to 35 hours per week. Today the average college
student studies 8 hours per week, but spends 28 hours on social media (CPTI)

3.

32% of college seniors agreed with the statement that Google has made studying obsolete

4.

The average college student doesnt start studying until 2 days before a major exam

5.

50% of college graduates had difficulty or could not interpret information on a graph

6.

Almost 85% of college students said cheating was necessary to get ahead (U.S. News and World
Report).
Only 27% of college graduates work in a field related to their major. Employers report new-hire
graduates lack core knowledge to do the jobs for which they were hired, exhibit poor or no
writing skills, and are unable to work effectively as a team

7.

8.

In 1969, US students ranked first in the world in Science, Math and Technology we now rank
th
29 among developing countries

IMPORTANT WEBSITES
How to Get the Most Out of Studying:

Video 1: Beliefs That Make You Fail...Or Succeed [7 min]

Video 2: What Students Should Know About How People Learn [7 min]
Video 3: Cognitive Principles for Optimizing Learning [6 min]
Video 4: Putting Principles for Learning into Practice [9 min]
Video 5: I Blew the Exam, Now What? [7 min]

You will get the most out of the material if you view the 5 videos in order. I

COMPARATIVE PHYSIOLOGY
THE QUEEN OF BIOLOGICAL SCIENCES
THE OBJECTIVES:

Understand the philosophy of comparative physiology, its goals, approaches and

domain
Understand strengths and weaknesses of the two major paradigms used to study
animal physiology

THE MAJOR CONCEPTS:

Interdependence of form-function, and animal-environment as units of study

The relationship of comparative physiology to other biological sub-disciplines
The resource-condition concept
The Fry paradigm an Frys five famous environmental entities

THE DETAILS:
1. A brief summary of the study of Comparative Animal Physiology
a.
b.
c.
d.

Definition-study of animal
DomainQueen of the biological sciencesWhat does the quote Structure without function is a corpse; function without
structure is a ghost mean to you?
(1) Studying anatomy can give insight to physiology vise versa

2. Why is the comparative approach used?

a. Almost everything we know with ecology, genetics and biology. Quantify the
range of variation between given traits among organisms. Limits of life. Helps us
understand structure and pattern. Mechanism of the physiological features.
3. The domain of the comparative physiology: One world, two views
a. The resource-condition concept
i. Assumptions of the Resource/Condition concept
ii. Definitions(1) Conditions; pH, Temp
(2) Resources: water ,food, space,
b. Advantages of the Resource-Condition Concept-very easily organized
c. Disadvantages- overly simplistic reductionist

4. The Fry Paradigm-animal response

a. Advantages/disadvantages-allows to ask and answer questions that A?C cannot
b. Assumes you have a large understanding of physiology
c. Frys Famous Environmental Entities
i.
ii.
iii.
iv.

Lethal Factors- animal can die

Controlling Factors-changes rate and pace of development
Limiting Factors-reduce active metabolism
Regulatory [Webb 1978] Factors (=Masking Factors)-ineractions between
factors that influence the other ie temp and oxygen
v. Directing Factors- taxis attraction or avoidance factor.

CELL MEMBRANES: A PRE-ADAPTATION TO INTEGRATED CONTROL

STUDENT LEARNING OUTCOMES:
Know the major factors & conditions effecting solute movement across cell
membranes
Understand the physical bases for these movements
Relate the import of these characteristics & conditions to cell homeostasis and
electrotonus
THE MAJOR CONCEPTS:

The Steady Ionic State

The General Diffusion Equation (GDE)
Ficks Law of Diffusion
The Nernst Equation
Electrochemical potentials
Donnan equilibrium
Molecular Pumps

THE DETAILS:
Understanding Steady Ionic State & How it is Preadaptative for Electrical
Integration
1. List 5 important cell membrane functions that make electrical events possible
a. Excellent barriers to free diffusion (semi permeable) concentration gradientsource of potential energy
b. actively picking things up and spitting things out-exo/endocytosis
c. receptive to certain types of compounds that changes the feature/state of the
membrane receptors
d. transmembrane potential/charge-voltage is a potential source of energy
e. conduct bioelectric charges-nervous systems
2. What is the steady ionic state?
a. Ionic steady state - Although cells are in osmotic equilibrium with their
environment, ion concentrations differ greatly between the cytosol and interstitial
fluid.
b. This definition leads to three important questions you should be able to answer.
i. How is this difference established?

ii. How is the difference maintained?

iii. How can this condition lead to integrated nervous control?
3. Four important characteristics that establish the ionic steady state
a. Membrane Permeability
i. Permeability (P) - the rate at which a substance passively penetrates the
membrane under a given set of conditions - is the key factor determining
physiological exchange rates between the internal and external cell
environment.
ii. Know the five physical & chemical factors affecting permeability
b. Solute concentration gradients
i. Chemical potential
ii. Diffusion
c. Electrical concentration gradients effects of charged molecules
i. Attributes
ii. Donnan Equilibrium - ion distribution in the face of undiffusible or fixed
ions
d. Active transported via molecular pumps
i. Membrane or molecular pumps have three basic features
ii. The sodium-potassium pump: A classic example.

Predicting Solute Exchange Rates

The General Diffusion Equation
A biophysics lesson
1. Rate of net solute flow (Jnet) = conductance of the solute driving force
dc z d
J net AD
c

dx RT dx

where A = area (cm2)

D = diffusion coefficient (cm2/s)
c = solute concentration (mol/cm3)
z = charge on the solute
= the Faraday (96,500 coulombs/mol)
R = gas constant (8.314 V coulomb/K mol)
T = absolute temperature (K)

 = electrical potential (V)

x = distance (cm)
2. Can we determine the rate of solute movement across a cell membrane using GDE?
Assume

z=0

What is the effect on the GED?

J ADdc / dx
This form is known as Fick's Law of Diffusion
(1) A is the surface area for diffusion
(2) D = permeability (diffusion constant) of the membrane tissue or substance
(3) dc/dx is the chemical potential (concentration or partial pressure gradient)
3. Can the GDE be used to determine movement of ions?
Assume
z0
Let Vm = electrical potential across the membrane
Flux occurs in both directions from compartments 1 & 2
Net flux = 0
Solve for Vm the GDE reduces to:

Vm

RT c1
ln
z c 2

This form is the famous Nernst Equation!

i. Shows how a diffusing ion is distributed across a membrane at equilibrium
ii. A most useful equation - accounts for both concentration & electrical gradient

NERVES AND THE EXCITING CONCEPT OF EXCITABLE MEMBRANES

HOW EXCITING!
STUDENT LEARNING OUTCOMES:
Understand excitable membranes physiology and how they differs from typical cell
membranes
Explain the bio-physical & morphological events of the resting & active excitable
membrane
Link the bio-physical & morphological characteristics of excitable membranes to
their functional attributes
THE MAJOR CONCEPTS:

The Resting Potential

The Action Potential
The Nernst Equation
the Goldman Adjustment
Gated Channels
Action Potentials Properties

THE DETAILS:
1. Four important attributes about cell membrane permeability and molecular pumps
2. Overview - The Resting Potential
a.
b.
c.
d.
e.

Resting potential and its magnitude for a typical excitable membrane

Do you remember the four factors that establish the resting potential?
List the resulting ionic characteristics we have seen these before
The electrical consequences each ions contribution to the resting potential
One KEY FACTOR leads to a negative trans-membrane potential in an excitable
membrane

3. What is the trans-membrane potential of a typical excitable membrane?

a. Defined by the Nernst equation:

RT [ K ] o
Vm
ln
z
[K ] i

Where:
Vm = trans-membrane potential (mV)
R = universal gas constant
T = Absolute Temperature (K)
z = charge carried by 1 gram equivalent of ions
[K+] o & [K+] i = outside and inside potassium concentration
b. What is the resting potential when inside and outside [K+] is equal?
4. Why doesnt the Nernst equation return a precisely accurate value?
a. The Goldman Adjustment
RT
E=
ln
z

P K [K ] o + P Na [Na ] o + P Cl [Cl ] i

P K [K ] i + P Na [Na ] i + P Cl [Cl ] o

Where:
Px is the membrane permeability for ion x
[X] o & [X ]I ion X, concentration outside and inside the membrane
5. Sodium permeability and the action potential
a. The permeability of Na+ is so low that it has a minor affect on resting potential.
b. What happens if there is a momentary increase in membrane permeability to Na+?
i. Depolarization & Action Potential: things every biologist should know!
ii.
Key Concept - membrane potential can be changed 125 mV, merely by
altering the relative permeability of sodium and potassium!
6. How is sodium permeability controlled?
a. The Na+/K+ pump & Gated Channels
b. Types of gates:
i. Ligand-gated channels
ii. Voltage-gated channels are self-closing
iii. Mechanical-gated channels
7. The Hodgkins Cycle and its key attributes

a.
b.
c.
d.

Stimulus
Local response
Threshold value
Hodgkin cycle
i.
ii.
iii.
iv.
v.

Sodium channels activated

Overshoot
Sodium channels deactivate
Rectified potassium channels activate
Hyperpolarization

8. Important properties of action potentials.

a. Regenerative or local response that is the question.
b. All or none. Conduction without decrement.
i.
ii.
iii.
iv.
v.
vi.

Absolute refractory period

Relative refractory period
Rectification.
Accommodation.
Adaptation: Is habituation a better term?
Hyperpolarization

NERVES: THE BASIS OF ANIMAL INTEGRATION

STUDENT LEARNING OUTCOMES:
Understand functional & morphological differences & similarities between the two
nervous tissue types
Explain the basic ultra-structure the typical neuronal cell
Identify explain morphological and functional attributes used to classify neuronal
cells

THE MAJOR CONCEPTS:

Excitable and non-excitable nerve tissue

Synaptic action potential transfer
Importance of neurotransmitters
Cable properties, ephaptic transmission, and saltatory conduction

THE DETAILS:
1. Two basic types of nervous cells What are they?
1. Neuroglial cells
1. Distribution among the phyla
2. Neuralgia cell types form and function
(1)
(2)
(3)
(4)
(5)

Ependymal cells
Astrocytes
Oligodendricytes
Schwann cells
Microglia cells

2. Neurons
1. Basic structure
(1) Cell body or soma
(2) Cell processes and their function
1
2

Dendrites
Axon
1
2

Collateral axons
Unmyelinated and myelinated

1) Schwann cells
2) Nodes of Ranvier
Synaptic
terminals

Motor nerve
ending

Dendrite
Schwan cell
(myelin sheath)
Node of Ranvier

Axon

Soma

2. Connecting neurons together - Axon terminals and synaptic batons

(1) Presynaptic membrane or terminal
1
2

AP reaches baton
Synaptic vessels

(2) Synaptic clefts or nexus

(3) Postsynaptic membrane and receptor (ligand) channels
3. Transmitter substances
(1)
(2)
(3)
(4)

norepinephrine (noradrenaline)
epinephrine (adrenaline)
acetylcholine
-aminobuteric acid (GABA)

2. Classification of neurons
1. Morphological
1. Multipolar neurons
2. Bipolar neuron

Some examples of multipolar and bi-polar neurons

Can you tell the difference?

3. Unipolar neuron

Some examples of unipolar neurons

2. Conduction rates Functional classification of nerves

1. Low resistance electrical pathways septal synapses or ephaptic transmission
2. Diameter - invertebrate solution (still found in anamniote vertebrates).
(1) Rate of conduction is defined by the neurons cable properties
(2) u=kd
1

Where:
u = transmission velocity
k = animal-specific fiber properties constant
d = diameter of the nerve fiber

3. Invertebrate use of giant neurons (Always large muscle groups for quick

escape)
4. Vertebrates - Lateral line nerves & Mauthner fibers in fish & amphibians
Mauthner neron

Axon

Lateral dendrite

Batons
Cell body

Ventral dendrite

3. Insulation
1. Myelinated neurons - vertebrate solution
(1) Myelinated neurons
(2) Nodes of Ranvier
(3) Saltatory conduction (from the Latin to dance or jump). The question is
how?

Saltatory Conduction

Node

Axon

Myelin sheath

Membrane under myelin

remains negative

Direction of impulse
_

- - - - Reverse potential
At active node

+
+
_

Outward current depolarizes

next node of Ranvier

-+ -+- +- +- +- +- +- -+ -+ - - - - - -+ -+- - - - - - - - - - - - - + + + + + + + + +
+ +

Current flows back

outside myelin

2. Invertebrate solution to meylinated nerves Do they differ from

vertebrates?

FROM NERVES TO NERVOUS SYSTEMS

THE OBJECTIVES:
Understand functional integration & emergent properties of nervous systems
Be able to track nervous system evolution within the animal kingdom & relate
important contributions of major animal groups
Identify key strategies of nervous integration and recognize advantages and limits of
each
THE MAJOR CONCEPTS:

Typical structure of nervous systems

Independent Effectors
The radiate approach to nervous systems
Similarities & differences in nervous systems of the bilateria
Evolution of nervous systems in the vertebrates
Measures of intelligence

THE DETAILS:
1. Nervous systems - Excitable nerve cell groups providing an interface between
sensory & motor responses.
a. Functions
b. Organization - Sensory-motor circuit:
i. Three parts (usually)
(1) Neuron receptor
(2) Motor neuron
(3) Effector cell
ii.

Interneurons may allow complex interpretation & integration.

(1) Divergence (2) Convergence (3) Feedback loops -

2. Are nervous systems necessary to support highly integrated and coordinated life?
a. Independent effector cells
b. Major advantages
c. Major disadvantage
3. Three important examples:

a. Complex movement patterns of ciliates

b. Poriferians
c. Cniderian cnidocytes
4. Radiates: Evolution of true effector/motor nervous systems
a. Radial symmetry
b. Sophisticated nerve nets
i.
Motor reflex response
ii. Semi- independent nerve nets
(1) Fast specific nerve net
(2) Slow diffuse nerve net

5. Bilateral animals: trends in nervous system evolution

a.
b.
c.
d.

Reduction in reflex motor units

Cephalization
Centralization of nervous control
Neuronal aggregation
i. Ganglia
ii. Nuclei

e. Fusion/reduction of nerve cords

6. Flatworms - show a wide diversity of nervous development
a. Primitive forms

b. Moderately advanced commissures or nerve ladders

c. Advanced forms - Brain or endom

Primitive flatworm nerve net

7. Mollusks - highly cephalized (not bivalves) with ganglia fused into few large central
masses
a.
b.
c.
d.

Circumesophageal ganglia
Ventral pedal nerve cords
Visceral nerve cords
Cephalopod circumesophageal ganglion

8. Annelids - distinctly more organized nervous systems

a. Bilobate brains
b. Ganglionic swellings

9. Arthropods refine basic annelid nervous plan to its highest integration level
a. Low sodium diet - perineurium
b. Brain sensory organs produce highly modified brain - 3 distinct regions:
i. Protocerebrum
ii. Deuterocerebrum
iii. Tritocerebrum

c. Segmental ganglia
i.
ii.

metathroacic ganglia
CO2 & hypoxia

d. Central pattern generators

10. Vertebrates differ from invertebratesa. CNS housed in bony chamber cushioned in cerebral spinal fluid to protect from

damage
b. The vertebrate brain divided into three regions
i. Rhombencephalon
(1) Pons-medulla
(2) Cerebellum - proprioreception
ii. Mesencephalon
(1) In fish and amphibians
(2) Sub-mammalian vertebrates
(3) Mammals
iii. Prosencephalon forebrain - divided into two parts
(1) Diencephalon - Thalamus, hypothalamus and posterior pituitary
(2) Telencephalon - undergone extensive changes in the course of evolution
(a) Fish & amphibians
(b) Reptiles & birds
(c) Mammals

c. How can we measure intelligence?

i. Observations of intelligence
ii. Physiological measurements of intelligence
(1)
(2)
(3)
(4)
(5)

Brain size
Cephalic index
Cephalic volume
Cortical Surface Area
Cortical Volume Ratio (SA:Thickness)

STUDY MODULE II

SENSORY SYSTEMS AND THEIR FUNCTION

STUDY GUIDE
Comparative Animal Physiology

Dr. Wayne A. Bennett, Associate Professor of Physiology

MECHANORECEPTORS
A most versatile sensory system
STUDENT LEARNING OUTCOMES:
Recognize importance, distribution and use of the mechanical receptor/sensory
systems
Identify and classify various mechanical sensory receptor types
Understand the structural functional components of general and special sensory
systems
THE MAJOR CONCEPTS:

Type and classification of senses

Sensory encoding
General mechanoreceptor types and structure
Equilibrium sensory adaptations of various animal groups
Audition in air & water
Magnetoreception

THE DETAILS:
1. Important definitions:
a. Receptors nerve/epithelial tissue responding to stimuli by developing action
potentials
i. Primary receptor cells specialized neurons (ancestral)
ii. Secondary receptor cells epithelial cells synapsing with neurons
(vertebrates only)
b. Stimulus - any environmental parameter causing a response in a nerve muscle or
gland
c. Sensation - perception or awareness of a stimulus received by sensory receptors
d. General and special senses
i. General
ii. Special senses
2. Sensory coding
a. Receptor specificity and morphological encoding

Olfactory Auditory

Muscle
stretch

Cutaneous

Examples of different receptor

types

b. Additional attributes of morphological encoding

i. Filtering
ii. Amplification
c. Innate receptor encoding
i. Pulse-code messages two types

(1) Tonic receptors

Horseshoe crab optic nerve tonic discharge rates

relative to intensity. Broken white line gives the
1-sec period during which the eye was illuminated.

(2) Phasic receptors

Stimulu
s

Nerve
fiber
Myeli
n

Node of Ranvier

3
5
Time in (msec)

Stimulation of a Pacinian corpuscle and

the resulting action potential spike.

ii. Signal filtering

(1) Low-pass
(2) High-pass
(3) Band-pass filter
3. General mechanoreceptors - response to sheer or torque forces
a. Structure
b. Function
c. Types

Free nerve
endings

Meissners Merkels
disks
corpuscles

Hair
Follicle

Ruffinis
endorgans
Pacinian
corpuscle

i.
ii.
iii.
iv.

Diagrammatic representation of mechnoreceptors

found in mammalian skin. Thick lines are
myelinated nerves, thin are unmyelinated
Light touch/pressure receptors - Merkels disks, Hair follicles
Discriminative touch receptors - Meissners corpuscles
Barrow receptors - Ruffinis end-organs
Nociceptors un-myelinated neurons two types:
(1) Fast response nociceptors
(2) Slow propagation nociceptors

v. Proprioreceptors - Pacinian corpuscles

vi. Thermoreceptors - un-myelinated neurons
vii. Infrared receptors

Pit Organ

Palmate sensory
neurons on pit
membrane

Single sensory
neuron showing
palmate design

SPECIAL MECHANORECEPTORS WITH ACCESSORY STRUCTURES

1. Morphology of secondary sense cells
i. Hair cells
ii. kinocilium & stereocilia
Sterocilia

Afferent
nerve

Kinocilium

Efferent
nerve

Typical vertebrate
hair cell or
secondary
mechanoreceptor

Hyperpolarization

+
Receptor
potential
Depolarization

Nerve
impulse

Resting
discharge

Increase impulse
frequency

Excitation

Decrease impulse
frequency

Inhibition

iii. Accessory structures

2. Types of Special Mechanoreceptors
a. Organs of equilibrium and orientation
i. statocyst & statolith
Receptor
cell

Nerve
s

Statolith
s

Typical invertebrate statocyst

ii. semicircular canals & vestibule (ampullae)

Ampullae

Ampullae

Otoliths

Cupulae
Hair tufts
Hair cells
Nerves

(1) Static equilibrium - otoliths.

(2) Kinetic equilibrium
3. Lateral line (Octaveolateralis) systems

Cupula

Hair tufts
Hair cells

Naked neuromast of
fish and amphibians
4. Audition
a. Hearing in water

i. Near-field sound

ii. Far-field sound

(1) Osterophysan fishes

(2) Hearing specialists

Swimbladder
extensions to
endorgans

Sonorific
muscle

Swimbladder

Swimbladder of the spotted sea trout

(Cynoscion nebulosus) a hearing specialist
that used sound for communication

(3) Cetaceans
b. Hearing in air:
i. Insects
Tympanum
Attachment cell
Sensory dendrite
Scolopale cell
Scolopidium
Scolopidium
sensory cell
Schwann cell

Sensory axon

Chordotonal organ of
insects are comprised of
many scolopidia

(1) chordotonal organ

(2) scolopidium
ii. The vertebrate ear
Pinna

Middle ear
Semicircular canals
Tympanic
membrane

Nerves
Cochlea

Auditory meatus

External ear

Ossicular chain

Internal ear

(1) External ear (auricle or pinna)

(2) Middle ear air- auditory ossicle: malleus, incus & stapes
(3) Inner ear - bony & membranous labyrinth, tactorial membrane,
basalar membrane, and spiral organ of Corti
iii. Hearing

Oval window

Round window
Basilar Membrane

Simplified view of mammalian cochlear

structure in the extended position

Vestibular canal
(round window)

Tactorial
membrane

Basilar
membrane

Hair
cells

Nerves
Tympanic canal
(round window)

Cross section of the cochlear

apparatus showing the organ of
corti (basilar and tactorial
membrane, and hair cells)

c. Specialized hearing adaptations in vertebrates

i. Echolocation
(1)
(2)
(3)
(4)

Bats microchiroptrans & macrochirptran

Shrews and some other insectivores
Whales, dolphins and seals
Cave swiftlets

d. Use of infrasound
i. Baleen whales
ii. Elephants
iii. Pigeons
5. Magnetoreception
i.
ii.
iii.
iv.

Electrically sensitive marine fish

Birds
Some bacteria & insects
Magnetite

CHEMOSENSORY AND VISUAL ADAPTATIONS

Gustation, Olfaction, and Vision three very important sensory modalities!

STUDENT LEARNING OUTCOMES:

Understand the physiological processes leading to chemosensory and visual
perception
Identify key integrative steps in each process
Know the range of variability among animal groups and how system variation affects
what is perceived for each modality
THE MAJOR CONCEPTS:

Type and classification of chemical senses

Sensory encoding of chemical and visual stimuli
General receptor types and structure
Evolution of visual adaptations

THE DETAILS:
1. Chemoreception - gustatory and olfactory receptors
1. Most universal sensory modality
2. Evolved independently in many groups
2. Olfaction
1. Bipolar neurons but secondary sense cells often involved
2. Exact physiological mechanisms not well known
Pore

Dendrites
Cuticle

Receptor cell
Axon

Diagram of an insect olfactory

cell

Cilia

Sustentacular cells
(non-neuronal)

Receptor cells
(differentiated dendrites)

Primary receptor cells from

human olfactory mucosa.

3. Steps in the initial process

4. But what is the exact mechanism involved?
1. Primary odor hypothesis
(1)
(2)
(3)
(4)
(5)
(6)
(7)

musky
floral
pepperminty
camphoraceous
ethereal
pungent
putrid

5. Uses
1.
2.
3.
4.

Foraging/feeding
Location or navigation
Reproduction and development
Protection

3. Gustatory (taste) responses

1. Found in all major phyla
2. Vertebrate taste buds

Taste hair

Taste pore

(microvilli)

Receptor cell

Capsule

Neurons

Vertebrate taste bud with supporting cells

omitted

1. Structure
(1) Taste hair and taste pores
(2) Process of gustation
2. Distribution
3. Four classic taste sensations plus two non-traditional
(1)
(2)
(3)
(4)
(5)
(6)

sour
salty
bitter
sweet
water
umami

3. Vomoronasal organ (Jacobsons organ) an olfactory system augmentation

Mushroom body

Lachrymal duct
Jacobson organ duct

Upper palate

Jacobsons Organ of a monitor lizard.

Dark arrows represent movement of
odiferous molecules

(1) Reptiles, amphibians & mammals

(2) Role of the forked tongue

ALL ABOUT VISION

4. Photoreceptors- more detailed information about near & distant environment than
other sense
1. Types of information
1. Intensity
2. Wavelength
3. Plane of polarization
2. Major advantage?
5. Photoreception in the animal kingdom
1. Dermal light sense or diffuse photosensitivity - nearly all phyla.
1. Exact mechanism not clear
2. Perhaps photosensitive nerve endings?
3. Type of information
2. Eye spots - flatworms, annelids & arthropods- not image forming
1. Forms:

Flat sheet

(1) Flat sheets

(2) Convex
(3) Cup-shaped

Concave

Convex

6. Image forming eyes - complete sharp image - two evolutionary lines

Laminar stacks

Rhabdomeric
tubes

Basic arrangements of light traps

within photoreceptors

1. Vesicular eyes comprised of - cornea, lens and retina

(1) Independently evolved twice
(2) Found in four different groups
2. Convex eyes - omatidia
(1) Found in annalids, mollusks & arthropods
(2) All evolved independently

7. Important anatomical features of image forming, single lens eyes

Cornea
Aqueous
humor
Iris
Ciliary
muscles
Lens
Vitreous
humor

Sclera
(outer tunic)

Choroid
(middle tunic)

Optic
disk
Fovea

Retnia
(inner tunic)

Neurons

Basic structure of the vertebrate

eye

1. Cornea
1. Accommodation and light refraction
2. Underwater vision
2. Iris and lens - anterior eye
1. Iris
2. Lens
(1) Shape - terrestrial vs. aquatic
(2) Color
(1) Affect on ultra-violet sensitivity
(2) Yellow lenses and countershading
(3) Using the lens to focuses light 3 ways to change focal length
(1) Move retina
(2) Move lens
(3) Change lens shape
3. The retina

1. Outer pigmented retina

2. Inner sensory retina
(1) Rods
(1) Rhodobsin
(2) Cones (wavelength)
(1) Iodopsins
(3) Trichromatic vision theory
Outer
segments
Photosensitive region
Cilium

(generation of AP)

Mitochondria
Inner
segment
Metabolic region
(synthesis & energy production)

Plexiform region
(synapse with nuerons)

8. Multifaceted / Convex eye differ radically from vesicular eye

Cross-section of the multifaceted

or compound eye

1. Ommatidia - photoreceptor units

Corneal lens

Crystalline cone

Primary iris cells

Rhabdom
Secondary iris cells

Retinula cells

Axon

Example of insect ommatidia

1. Fixed angle of resolution

2. Light guide, light shade & photoreceptors
9. Pineal or median eye of vertebrates
1. Best developed in lamprey and some lizards

2. Pineal organ complex

3. Photo-sensitive
4. Endocrine function
10. Eyes of the eyeless shrimp, Rimicaris exoculata
1.
2.
3.
4.
5.

Hydrothermal vents - black smokers

Lack image-forming optics
Location of rhodopsin
Black-body radiation
Sulphide bacteria

STUDY MODULE III

METABOLISM AND MOVEMENT

STUDY GUIDE
Comparative Animal Physiology

Dr. Wayne A. Bennett, Associate Professor of Physiology

ENERGY METABOLISM MEASURING THE COST OF LIVING

THE OBJECTIVES:
Understand, define, and explain what is meant by metabolism
Enumerate ways metabolism can be measured; recognize advantages & disadvantages
of each
Understand the metabolic components and interrelationships of each method
THE MAJOR CONCEPTS:

Relationship between metabolism and oxygen consumption

Four related but different metabolic measurement techniques
Various measures of metabolism
Food types and the concept of isocaloric weight

THE DETAILS:
1. What is metabolism?
2. Types of metabolic reactions
a. Anabolic reactions
b. Catabolic reactions
3. Why are metabolic measurements useful to physiologists?
4. Factors influencing metabolic rates
To be meaningful, metabolism must be measured carefully!
5. Some basic principles in the measurement of metabolic rates:
a. Basal Metabolic Rate (BMR) - metabolic rate of resting, fasting mammals and
birds under minimal physiological and environmental stress.
i. Endothermic animals
ii. Constant body temperature
b. Standard Metabolic Rate (SMR) resting and fasting metabolism of
poikilotherms under minimal physiological and environmental stress, at any given
temperature.
i. Poikilothermic animals
ii. Variable body temperature

c. Why fasting? Because of Specific Dynamic Action

6. * Key Point BMR and SMR are measured under unnaturally controlled and quite
conditions that vary greatly from an animals normal state & say nothing about
activity.
7. Other common metabolic measures
a. Routine Active Metabolic Rate - average metabolic rate of normally active
animal.
b. Maximum Sustained Metabolic Rate metabolic rate at sustained, vigorous
activity.
c. Metabolic Factorial Scope or Index of Metabolic Expansibility - ratio of
MSMR to BMR or SMR.
8. Metabolic rates can be determined in four different ways.
a. Mass balance equations
i. Ballistic bomb calorimetry - anabolic heat energy
ii. Also require moment-to-moment (catabolic) measures
iii. Advantages & disadvantages
b. Direct calorimetry Determine total heat production
i. Hesss law heat released through breakdown of a fuel to a given set of end
products is constant irrespective of the intermediate chemical steps or
pathways used.
ii. Advantages & disadvantages
c. Indirect calorimetry - Determine the rate of oxygen consumption (most often
used).
i. Respirometry
(1) Manometric respirometry

Manometer
Tube

Gas-tight
Syringe

Water
Bath

Reference
Flask

Respirometer
Flask

(2) Flow-through respirometry

Water level

Water inflow

Sample tube

Water outflow

Example of a flow-through
respirometer. From Cech 1990
(3) Sealed-jar respirometery

Water
Bath

Measurement
Flask
ii. Advantages and disadvantages
d. Nuclear magnetic resonance (NMR)
i. Seldom used
ii. Advantages/disadvantages

9. Conversion of oxygen consumption to heat Isocaloric values (Table 1)

TABLE 1. Energetics of common foods.
kcal g-1

liter O2 g-1

kcal per liter O2

Carbohydrate

4.20

0.84

5.0

Fat

9.40

2.00

4.7

Protein (urea) 1

4.30

0.96

4.5

Protein (uric acid) 1

4.25

0.97

4.4

Food

Protein energy values are higher than listed; however, proteins are incompletely burned
in vertebrates.
a. Fats, carbohydrates and proteins have different energy values by weight.
b. Amount of energy released per liter of oxygen consumed remains relatively
constant.
c. Important Shortcut - 4.8 kcal per liter of O2 (avg. kcal / L) 6% largest error
possible

FACTORS AFFECTING METABOLIC RATES OF ANIMALS

THE OBJECTIVES:

Explain why metabolism is the common biotic component of life

Understand the relationship between metabolic rate and environment
Assess how changes in the oxygen, pressure effect animal metabolic rates
Enumerate the effects of allometery on metabolism

THE MAJOR CONCEPTS:

Pressure & oxic influences on metabolism

Metabolic conformation & regulation
Anaerobic/aerobic coupling
Low volume to high volume reactions
The mouse to elephant curve
Kleibers Law and its potential cause

THE DETAILS:
OXYGEN
1. Oxygen in terrestrial environments
2. Oxygen in aquatic environments
a. Metabolic regulators - oxygen independent
b. Metabolic conformers - oxygen dependent
c. Critical oxygen tension

Metabolic response of goldfish acclimated at various temperatures

3. Adaptations of water-breathers to survive hypoxic events

a. Behavioral
i. Vertical migration
(1) Top-ward migration ASR
(2) Bottom migration
ii. Horizontal migration
b. Physiological adaptations
i. INCREASE OXYGEN UPTAKE
ii. ANAEROBIC PATHWAYS
(1) oxygen debt
(2) aerobic coupling

PRESSURE

Pressures at various points in the atmosphere and hydrosphere.

RELATIVE POSITION IN THE
ATMOSPHERE OR HYDROSPHERE

ABSOLUTE PRESSURE IN
ATMOSPHERES

PRESSURE IN PSIG*

Mt. Everest

0.25

3.5

Sea level

1.0

14.1

28.2

300 to 500

4,200 to 7,050

10 m below sea level

Average abyssal pressures
(3 to 5 km below sea level)

Deepest ocean trenches

(> 10 km below sea level)
> 1,000
* Pounds per square inch exerted at ground (sea) level

1. Four ways high pressure affects metabolism:

141,000+

i.
ii.
iii.
iv.

Disrupt pH systems
Shift velocity constants
Collapse weak chemical bonds
Alter liquid to solid phase transitions

b. How do deep-sea animals withstand pressures of up 1000 atmospheres?

i. Colloidal biophysics adapted to different dissociation and velocity constants.
ii. Enzymes show significant increases in strong disulfide bonds & salt bridges.

BODY SIZE
1. Total oxygen consumption vs. Specific oxygen consumption
TABLE OF Oxygen Consumption in Mammals of Various Body Size

Animal

Body Mass
(g)

Total O2
consumption
(ml/h)

Mouse

25

41.0

1.65

Ground squirrel

96

98.8

1.03

Dog

11,700

3,870

0.33

Human

70,000

14,760

0.21

Horse

650,000

71,100

0.11

3,833,000

268,000

0.07

Elephant

2. Take-home Message

Specific O2
consumption
(ml/g h)

3. The famous mouse-to-elephant curve Illustrated graphically

The famous mouse to elephant curve of metabolic rate on body size

a. linear relationship is represented by the equation:

VO/Mb (l/kg h) = 0.676 x Mb-0.25

Oxygen consumption
(liter O2 h-1

b. If mass (Mb) is removed from the equation the equation becomes:

10
1

Slope = 0.75

0.1
0.01
0.01

0.1

10

100 1000

Body Mass (kg)

VO2 = 0.676 x Mb0.75

c. Kleibers law.

d. Max Rubner surface area hypothesis

i. Should this hypothesis be rejected completely?
ii. Two examples: One hypothetical and one observed.
e.
f.
g.
h.

McMahon and Bonners (1983) cross-sectional area hypothesis

Swans additive scaling hypothesis
Blums four-dimensional scaling hypothesis (1977)
Sernetzs fractal scaling hypothesis (1985)
i. Can be defined by fractal dimensional analysis equation
Vo2 = aMb-f
Where:
b = scaling exponent
a = constant for that group
f = fractal exponent (which changes with mass)

TEMPERATURE AND METABOLIC RATE:

Response of animals to minimize temperature effects

THE OBJECTIVES:
Understand the importance of temperature to animal life
Enumerate the thermal physiological strategies and tactics used by ectotherms
THE MAJOR CONCEPTS:

Thermal Primacy
The Arrhenius Principal
Temperature Quotient
Homeothermy
Precht Type Curves

THE DETAILS:
1. Heat and Temperature
a. Thermal Primacy Paradigm
b. Effects of heat on chemical reactions
i. The Arrhenius principal.
ii. Consequences
c. Effects on enzyme systems
i. Jacobus van't Hoff - and the temperature quotient or Q10.
(1) vant Hoffs rule
(a) Temperature quotient
(i) Q10 = Rate (at T1 + 10C) Rate at T1
(ii) Q10 = (Rate at T2 Rate at T1) 10 (T2 - T1)
2. Terminology
a. Warm-blooded & Cold-blooded
b. Homeothermic & heretothermic or poikilothermic
c. Endothermic & ectothermic - Terms of origin
3. Metabolism and ambient temperatures
a. Endotherms.
(1) thermal neutral zone
(2) upper and lower critical temperatures
b. ectotherms
i. Relationship to vant Hoffs rule?
c. The concept of compensation in ectotherms
i. Precht type compensation curves
d. Effects of extreme temperature on animal systems
i. Cell damage
ii. Equilibrium imbalances
iii. Loss of enzymatic control

4. Endotherms - homeothermy
a. Thermogenesis in endotherms
b. General physiological control- The futile cycle
c. Specific physiological control
i. The hypothalamus
ii. Conclusions?
d. Shivering thermogenesis
i. Female Indian pythons
ii. Honeybee swarms
5. Non-shivering Thermogenesis
a. Brown adipose tissue or BAT.
i. termogenin
b. Mechanisms to conserve metabolic energy
i. Dual set-point regulators
(1) Hibernation
(2) Diel torpor
(3) Carnivorus lethargy
ii. Brumation
iii. Estivation
c. Regional homeothermy/heterothermy
i. rete mirabile
(1) Warm bodied fishes - rete mirabile
ii. Regional endothermy - Swordfish

SKELETAL SYSTEMS
THE OBJECTIVES:
Evaluate stress/strain curves and give their meaning.
Describe the types of skeletons found in the animal kingdom.
THE MAJOR CONCEPTS:

Material properties
Elastic modulus
Hydraulic skeletons
Ridged skeleton types

THE DETAILS:
1. Functions of a skeletal system
a. Major functions
b. Other functions
2. Material properties of skeletons
a. Density (g/cm3)
i. Some important ratios
(1) Body fluids
(2) Flexible biological materials
(3) Rigid skeletal materials
b. Elastic modulus
i. Elasticity
ii. Compliance
iii. elastic modulus
(1) Stress
(2) Strain
(3) Stress on Strain
c. Plasticity
i. yield point
d. Ultimate strength
i. fracture point
3. Materials
a. Elastic organic compounds
b. Inorganic compounds that resist compression
TYPES OF SKELETONS
1. Hydraulic skeletons - three elements
i. Fiber angle
2. Hydrostat types
a. Fluid & soft walls
b. Fluid & muscle cells
i. Muscular hydrostats

3.

1.

2.
3.

c. Fluid & rigid elements

Rigid skeletons
a. Exoskeleton
i. Advantages
ii. Disadvantages
(1) Compressive buckling
iii. Arthropods exoskeleton
(a) epicuticle
(b) procuticle
(i) Exocuticle
(ii) Endocuticle
iv. Mollusks
b. Endoskeleton
i. Advantages
ii. Disadvantages
(a) cancellous bone/pneumatized bone
6. Examples of endoskeletons
Poriferians
i. spongin
ii. spicules
Echinoderms
i. Ossicles - test
Vertebrate endoskeleton
a. Notochord
b. Cartilage
i. Chondrocytes
c. Bone
i. Osteocytes - hydroxyapatite
(1) Lacunae
d. Bone types
i. Long bones
ii. Short bones
iii. Flat bones
iv. Irregular bones
e. Bone structure
i. Compact bone - osteon central Haversian canal
ii. Cancellous (spongy) bone trabeculae

MOVEMENT WITHOUT MUSCLE

THE OBJECTIVES:
Understand basic principles of non-muscular movement
Be able to describe and give the function of various molecules of motion
Identify key physiological processes of ciliary, amoeboid, & flagellar movement
THE MAJOR CONCEPTS:

Basic movement types

Molecules associated with motion
Cytoplasmic streaming
Cilia & flagella structure and function

THE DETAILS:
1. Basic types of movement
i. Amoeboid
ii. Ciliary or flagellar bending
iii. Direct cell movement
iv. Muscle contraction
2. Molecules of Motion
a. Contractile proteins
i. Actin
(1) G-actin & F-actin
ii. Intermediate fibers
iii. Tubulin
(1) heterodimers
(2) microtubules.
b. Molecular motors
i. Myosin
(1) spontaneous cross-bridges
ii. Dynein
iii. Kinesin
c. Regulatory proteins
i. Tropomyosin
ii. Troponin
iii. Calmodulin
iv. Alpha-actinin
3. Amoeboid Movement
a. cytoplasmic streaming
b. sol and gel state
i. actin regulated
ii. myosin regulated.
iii. How it works
(1) endoplasm

(2) fountain zone.

(3) gel-like ectoplasm.
4. Cilia and Flagella
a. Disadvantages?
b. Differences between cilia and flagella
(a) Neuroid hypothesis
(b) Coupled oscillator hypothesis
c. Mechanism of movement
i. The doublet microtubulin structure

MUSCLE MOVEMENT
1. Muscle types
2. Muscle fibers or myofibers or muscle cell
a. sarcolemma
i. transverse tubule system
ii. T tubules
b. Sarcoplasmic reticulum
i. Terminal cisternae
(1) Primary functions
3. Myofibrils
a. sarcomeres - functional unit of muscle contraction.
b. myofilaments or muscle filaments.
i. Thick filaments
(1) myosin
ii. Thin filaments
(1) actin
(2) Z-line
iii. Tropomyosin
iv. Troponin
c. zonation and lines
i. Z line (Zwischenscheibe or between disk)
ii. I band (isotropic)
iii. A band (anisotropic)
iv. H (helle or bright) zone
MUSCLE MOVEMENT - the sliding filament theory
4. motor unit concept
5. The neuromuscular junction
a. acetylcholine & neuromuscular junction
b. motor end plate
6. Excitation-contraction coupling
a. inositol triphosphate
7. Muscle Contraction
a. Resting stage
b. Cross-bridge formation
c. Power stroke
d. Release stage
8. Regulation of muscle contraction
VARIATION AMONG MUSCLE TYPES
9. Two basic types
i. Tonic muscle fibers
ii. Slow phasic fibers
iii. Fast phasic glycolytic fibers
iv. Fast phasic oxidative fibers
b. Cardiac muscle
c. Smooth muscle

(1) (varicosities)
(2) calmodulin

WATER WATER EVERYWHERE

THE OBJECTIVES:
Gain a general understanding of water balance budgets
Explain mechanisms used in direct and indirect water conservation and loss
Evaluate water balance adaptations and explain how they complement animal life
history and environment
THE MAJOR CONCEPTS:

The Water Budget Concept

Direct control vs. indirect control of water
Resistance to evaporative water loss
Water absorption from sub-saturated air

THE DETAILS:
1. Major water balance problems in different habitat types
a. The water budget concept
i. Potential avenues of water loss
ii. Potential avenues of water gain
2. Two major strategies of maintaining a water budget
a. Direct control of water balance
b. Indirect control of water balance by controlling osmotic flux (osmoregulation)
3. The Biophysics of Water Balance
a. In terrestrial animals EWL = -D xWV / d
i. D = diffusion coeficieny
ii. xWV = difference in water vapor density
iii. d = diffusion path length
b. Not usually measured in this way but rather as resistance
i. Resistance r is substituted for D & xWV
ii. EWL xWV / r
Some Adaptations for Direct control of water
4. Contractile vacuoles

a. spongiome layer.
b. Arginine vasopressin
5. Adaptations for retarding water loss across body surfaces
a. Mollusks
b. Arthropods epicuticle
c. Fish
d. Amphibians
i. Waterproof frogs wiping behavior cutaneous lipid glands
ii. Casque-headed frogs (Trachycephalus) - co-ossified skull
iii. Estivating amphibians
e. Aminote vertebrates - Keratinized epidermal cells, stratum corneum
f. Reptiles
g. Mammals
i. Three possible adaptations for preventing water loss from mammalian
respiratory surfaces in the absence of heat stress.
ii. Do not fully saturate expired air.
iii. Higher oxygen extraction. (Greater than 5%)
iv. Exhale air at lower than body temperature.
v. Temporal counter-current heat exchanger
h. Female mammals have an additional water burden during lactation.
6. Direct water vapor absorption from sub-saturated air via hygroscopic organs.
a. agranular cells
b. eversible bladder
c. rectal sacs containing hygroscopic fluid
7. Storage of water
a. Chiroleptes,

OSMOREGULATION AND EXCRETION

Secretory Organs of Excretion
THE OBJECTIVES:
Describe the relationship between osmoregulation and excretion
Explain the two basic types of excretory organs
List and describe the basic function of various cells/organs present in various animal
groups

THE MAJOR CONCEPTS:

Ultrafiltration, reabsorption and secretion

Chloride cells
Salt glands
Malpigian tubules

THE DETAILS:
1. Basic Processes of Excretion
a. Ultrafiltration
b. Active transport
i. Active secretion
ii. Active reabsorption
2. The following are secretion (not filtration) organs and thus are specific to certain
compounds.
3. The teleost gill - Chloride cells
a. Gill filament - Secondary lamellae
(1) Lamella epithelium
(2) Chloride cells
b. Keys and Willmer
(1) (trans-cellular transport)
(2) (para-cellular transport)
4. Salt glands
a. Control
i. Hypothalamus and osmoreceptors
ii. Hormones
iii. Major advantage
b. Location
(1)
(2)
(3)
(4)

Birds and reptiles.

sea turtles and marine iguanas.
Chocadillians
sea snakes

c. Structure

i. Knut Schmidt-Nielsen
ii. Secretory tubular system, tubular capillaries and a central duct.
iii. counter current system
5. Maligian tubules
a. Structure/Function
i. mid hindgut junction
ii. passive movement chloride
WATER BALANCE ORGANS OF EXCRETION
THE OBJECTIVES:
Gain a general understanding of excretory organs
Explain distribution and basic function of ultrafiltration organs in various animal
groups
Explain how various adaptations differ for animals in different environments
THE MAJOR CONCEPTS:
Protonephredia and metanephredia
The vertebrate kidney
Counter-current magnification systems
THE DETAILS:
1. Protonephridia and metanephridia.
a. Protonephridia
i. "flame calls"
(1) Found mainly in acoelomate or psuedocoelomate animals.
b. Metanephridia
i. Found ONLY in eucoelomate animals, but the reverse is not true i.e., some
animals with a coelome have protonephridia.
c. Function
2. Vertebrate Kidneys
a. Function
b. Structure

i. The Renal Tubular system

(1) Renal Corpusle
(a) Glomerulus
(b) Bowmans capsule
(2) Nephron (all of the above plus the following)
(a) Proximal convoluted tubule
(b) Loop of Henle
(i) Counter-current magnification
(c) Distal convoluted tubule
(3) Collecting ducts
(4) Renal blood flow
(a)
(b)
(c)
(d)

Afferent arterioles
Glomerulus
Efferent arterioles
Peritubular capillaries

c. Variability of kidney morphology among marine vertebrates.

i. Excretion in fish
(1) Aglomerular kidneys
(2) Examples
(3) Elasmobranchs have a reabsorption mechanism for urea.
ii. Excretion in the crab-eating frog
(1) Crab-eating frogs
iii. Excretion in reptiles
iv. Excretion in marine birds
v. Excretion in mammals

The Oxygen Environment, Diffusion, & Respiration

Behavior of Gases in Aerial and Aquatic Environments
THE OBJECTIVES:
Understand the physical laws that affect excretory organ form and function
Explain how each law dictates basic function of respiratory organs in various animal

groups
Explain how diffusion and gas absorption differ between aerial and terrestrial
environments
THE MAJOR CONCEPTS:

Daltons Law
Henerys Law
Ficks Law
Ficks famous silo problem

THE DETAILS:
1. Importance of Oxygen
2. Gases in air - the aerial environment
a. What is the fate of oxygen in aerobic organisms?
i. Energy Production by Cellular Respiration.
ii. Electron Transport Chain
O2 + 4H+ + 4e- = 2H2O
b. Composition of Gases in the Atmosphere
i.
ii.
iii.
iv.

Nitrogen
Oxygen
Argon
Carbon Dioxide

78.09%
20.95%
0.93%
0.03%

c. Dalton's Laws of Partial Pressure (Three parts but only two are important to us)
i. Effects of altitude on gas concentration and partial pressure
3. Gases in Water
a. Gas tension
b. Defined by Henry's Law which states - Tension of a gas in water is precisely
equal to the partial pressure of that gas in the gas phase with which it is in
equilibrium
c. Solubility - solution dissolves a specific amount of any gas it comes into
equilibrium with
i. Solubility coefficient ()
ii. V g mls / L ( Pgas ) PB
d. Factors determining the Vg in water

i.
ii.
iii.
iv.

Nature of the gas

Pressure of the gas in the gas phase
Temperature
Other dissolved solutes

4. Diffusion of gases
a. Diffusion of gas between two gases or between two liquids.
b. Diffusion occurs in response to partial pressure (tension) gradients only
c. Diffusion is the only mechanism of gas exchange between environment and living
cells
5. Animal Systems and the Movement of Gases Respiratory Membranes
a. Important definitions of respiration
i. Ventilation (Breathing) - Bulk air or water movement across a respiratory
surface
ii. Respiration - exchange of O2 and CO2 in all living organisms.
iii. Cellular respiration Cellular O2 & CO2 exchange resulting in ATP
production
iv. External Respiration - O2 and CO2 exchange across a respiratory membrane
v. Internal Respiration - Exchange of O2 and CO2 at the tissue level
FICK'S FIRST LAW OF DIFFUSION
11.

J = -D A (C2 - C1) x

vi. Where:
(a) J = Total O2 or CO2 flux per unit time (moles per sec)
(b) D or K = Diffusion coefficient or Kroghs diffusion constant; a
physical constant.
(c) A = respiratory surface area (cm2).
(d) C2 - C1 = Difference in the concentration (or partial pressure whichever
is appropriate) between the medium and the organism.
(e) x = distance over which diffusion occurs (cm)
6. Fick's famous silo problem
Think like an ecological physiologist!!
7. Based on these limitations, how would you design a respiratory gas exchanger?

THE PHYSIOLOGY OF RESPIRATORY SYSTEMS

All I need is the air that I breathe
THE OBJECTIVES:

 Describe how temperature, pressure and diffusing gas attributes affect diffusion
Explain how V/Q relationships are useful in maximizing oxygen uptake
List and describe major respiratory structures in the major animal phyla
THE MAJOR CONCEPTS:

Utilization coefficient
V/Q properties
Diversity of respiratory systems
Adaptations that enhance animal respiratory systems

THE DETAILS:
12.
1. Factors affecting diffusion of oxygen and their consequences
a. Temperature
b. Nature of the biological material
i. Nature of the medium
ii. Utilization coefficient
1.
iii.

EW

CiO2 CoO2
CiO2

100

2. Engineering a respiratory structure some general physiological considerations

a. Convection - ventilation/perfusion considerations
i. Thin membranes
ii. V/Q match
(1)
(2)
(3)
(4)

V/Q ratios are highly variable

Properties of respiratory medium
Temperature
Type & concentration of respiratory pigment

iii. V/Q flow patterns

iv. Ventilated pool
v. Parallel or con-current flow arrangement
vi. Cross-current arrangement
vii. Counter current arrangement
3. Diversity of respiratory structures

a.
b.
c.
d.
e.
f.
g.
h.

Integument
Gill
Water lung
Trachea
Tracheal gill
Compressible gill
Incompressible gill
Air lung

4. Respiratory systems in the animal kingdom

a. Cutaneous respiration
i. Limiting thickness
(1)

8 [O2 ]

K
M O2

(2) Boundary layers

b. Gills & branchial gas exchange - Expansion of the body wall to form gills
i. Invertebrate gills
(1) Polychaetes
(2) Echinoderms
(a) Retractile dermal papillae skin gills
(i) Perivisceral system
(b) Tube feet podia
(c) Holothuroidians water lung or respiratory tree
(3) Molluscs
(a) Ctenidia gills of broad flattened filaments with ciliated margins
(4) Gilled arthropods
(a) Crustaceans
(i) Ventilated by scaphognathite derived from the 2nd maxilliaped
(b) Horseshoe crabs book gills
(c) Aquatic insects
(i) Gills pass oxygen into branched trachea
(ii) Diffusion gill
(iii)Plastron gill

ii. Vertebrate gills

(1) Aquatic classes branchial filaments (1 lamellae) with secondary
branches
(2) Secondary lamellae show folding micro-ridges
c. Air breathing
i. Tracheal systems
ii. Diffusion lungs and bladders gas-filled chambers within the body
(1) Book lungs many arachnids
(a) lamellae held apart by spacer-bars
(2) Diffusion lungs of pulmonate snails
(3) Branchial chamber specialization
(a) Terrestrial crabs gills greatly reduced or absent
(i) Branchiostegites (region of carapace covering the branchial
chambers)
(ii) Called a branchiostegal lung
(b) Variety of branchial chamber modifications seen in fishes
(i) Labyrinth organs
(ii) Buccal-branchial expansion Panamanian swamp eel,
Synbranchus
(4) Lungs in air-breathing fish
(5) Amphibian lungs
(a) African clawed frogs, Xenopus
(b) marine toad Bufo marinus
(6) Reptile lungs intermediate between amphibians and mammals
(a) Lung divided into anterior and caudal sections.
(b) Saccular non-vascularized caudal air sac
(c) Highly vascularized anterior section divided into faveoli.
(7) Mammal lungs
(a) alveoli
(8) Avian lungs Anatomy - the most specialized of all vertebrate lungs.

(a) Air sacs

(b) parallel cylindrical tubes called parabronchi
(c) air capillaries