Lecture Notes - Animal Physiology (Melbourne)

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Animal physiology - Lecture notes All lectures
Comparative Animal Physiology (University of Melbourne)
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Comparative Animal Phisiologi

SEMESTER 2
Homeostasis 2
What is animal physiology?
Homeostasis 3
Hormones, receptors, pheromones 5
Stress response 8
Osmoregulation and the environment 11
Ecophysiology 18
Thermal relations
Hot & cold climates 21
Metabolism 25
Nutrition and conservation 29
Phisiological sistems 33
Cardiovascular system
Respiratory system 44
Nerves, CNS and synapses 55
Reproduction 66
Gonads and behaviour
Sex 70
Reproductive strategies 75
Environmental influences 79
Marsupials 86
KVITI2016 1

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Homeostasis
What is Animal Phisiologi?

 Animal physiologists study process that operate in living organisms at all levels including
functions of tissues and organ systems
o Also study the regulation of processes within groups of cells and how the combined
activity of cells affects the function of the animal
 Cell and molecular physiology: study at cellular level e.g. molecular genetics, biochemistry,
signal transduction
 Systems physiology: how cells and tissues interact to carry out specific functions e.g.
pacemaker cells in triggering heart muscle to contract
 Organismal physiology: the way an animal undertakes a specific process or a behaviour e.g.
metabolic rate in response to temperature
 Ecological physiology: how the physiology of an animal influences the distribution of a species
or population
 Integrative physiology: attempts to understand physiological processes at a variety of levels of
biological organisation and across multiple systems
Comparative animao physiooogy

 Enables you to identify if a particular function is carried out by related and unrelated species
in different environments
 Way of assessing outcomes of evolution
e.g. internalised breathing is an adaptation for animals than live on land
Adaptation, accoimatisation and accoimation

ACCLIMATISATION
 Physiological change that can occur within an individual animal that results from chronic
exposure to a new but naturally occurring environmental condition
 Time frame = days
 Long-term
 Reversible
ACCLIMATION
 Occurs when an animal is placed in an artificially controlled environment to mimic the high
altitude conditions of mountain slopes
 Long-term
 Reversible
e.g. hypobaric chamber
ADAPTATION
 Animals that have made adaptations to their physiological systems which enable them to
survive these conditions
o Change in a population over evolutionary time i.e. many generations
o Synonym for acclimatisation/acclimation (many argue this is an incorrect usage of the
term)
 Permanent
 Not reversible
 A physiological process is said to be adaptive if it is present at a high frequency in the
population because it results in a greater probability of survival
e.g. Llama blood has an unusually high affinity for oxygen (speculate that this was an adaptation
by llamas to live in a low oxygen environment). Camel blood also has an unusually high affinity for
oxygen. So high affinity for oxygen has more to do with the fact that these animals belong to the
camel family and less to do with the environment.
 If a similar physiological process occurs in several distantly-related animal species, living in
similar environments, then you would suggest this process or structure to be adaptive e.g.
ability of desert animals to concentrate their urine due to the extended loop of Henle in the
kidney

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 If a particular physiological process is shared by closely-related animal species living in vastly

different environments then it is not adaptive e.g. camel and llama blood sharing the unusual
high affinity for oxygen
Changes in physiooogy in response to changes in environment

 Acute changes
 Chronic changes
 Evolutionary changes
 Developmental changes
 Changes controlled by periodic biological clocks
Homeostasis
Environment
 Environment: the chemical, physical and biotic components of an organisms’ surroundings
o Internal environment e.g. body fluids, body temperature
o External environment e.g. temperature, oxygen, water (aquatic animals)
e.g. The number of species of swallow tail butterflies decreases at high latitudes. But this may not be
due simply to an effect of low temperature on the butterfly – it may be due to the effect of temperature
on food supply. But species can make changes to their physiological systems so that they can survive
in adverse environments.
Long-term changes
e.g. Rock cod in the sea around Antarctica hatch, grow, feed and mate at body temperatures near -1.9
degrees Celsius. Metabolically synthesized antifreeze compounds keep them from freezing.
e.g. Thermophilic (heat-loving) desert iguanas can survive body temperatures of over 45 degrees
Celsius, one of the highest tolerated by any vertebrate animal.
Animaos can modify their own environments

 If you live 1m below the surface (in a burrow), temperatures remain between 15-32°C
 Rather than living 2m above and having much greater and more frequent fluctuations in body
temperature
Homeostasis
 Ability of animals to maintain a relatively stable internal environment
o Physiological regulatory systems maintain internal conditions within a relatively
narrow range
 Many animals seem to live comfortably in their environments but most environments are
actually quite hostile to animal cells
 Large variations in external environment are offset by responses of control systems so that
the internal environment remains constant
HOW DOES IT COME ABOUT?

 The regulatory processes that maintain homeostasis in cells and organisms depend on
feedback systems
o Sensor e.g. thermostat
o Effector e.g. central heating/gas heater
o Output feedback
o Input feedback
Feedback systems
 Sensors in animals are called receptors
 Circuits can be composed of nerves and hormones
 Effectors have to be able to alter the internal environment and return it to the normal range
despite the wide variations in the external environment
NEGATIVE FEEDBACK
 Sensor sends an error signal to switch off the system – inverting amplifier
 Shuts off the effector
 Negative feedback is opposite to the change e.g. once heated, heating is turned off
 Sensor: pituitary

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o Detects high levels of stress

 Adrenal gland releases cortisol
o Acts on muscles, cells, liver to generate energy
o Decreases stress levels
POSITIVE FEEDBACK
 Enhances physiological response to effector
 Makes system operate on higher level rather than shutting it down
e.g. birth, vomiting (contractions get stronger)
 Sensor: hypothalamus
 Effector: releases oxytocin to contract
o Baby’s head presses on cervix
o Hypothalamus detects pressure of head and increases production of oxytocin
Conformers
 When an animal is confronted with changes in its environment, it can respond in two ways: it
can either conform or regulate
 Internal variable is always equal to the external environment
o Conformers adjust their internal conditions to reflect external environment
 Allow internal environment to change along with external environment, yet are able to
compensate for these changes and function in spite of them
o Generally, show a wide range of internal conditions that allow survival
o However, function may not be optimal over the entire range that is tolerated
e.g. Sea Star

 Their internal body fluids quickly come to equilibrium with the sea water that surrounds them
o So they experience an increase in body fluid salinity when placed in high salt water
 But they can also re-adjust if placed in low salt water and will show a decrease in body fluid
salinity
e.g. Crab-eating frog

 Live in mangroves – mangroves can change their salinity as they live in brackish water (on
the border of sea water and fresh water)
 Body fluids in 80% seawater – 205mM Na+, 227mM Cl- and 350mM urea, osmotic
concentration approx. 800mM (similar to seawater)
 Osmolar stability is achieved by increasing the concentration of intracellular organic
substances (osmolytes e.g. urea) that act to increase intracellular osmolarity
REGULATORS
 Internal variable is kept constant and therefore differs from the external environment
 Regulators use physiological, biochemical and behavioural mechanisms to regulate their
internal environment over a broad range of external environmental changes (homeostasis)
Thermoregulators
 Animals that actively maintain their body temperature within a narrow range, despite large
changes in their external environment
 Endotherms need to “manage” their heat budget so that rates of heat gain are equal to rates
of heat loss
o If the heat budget gets out of balance, the animal with either become warmer or
colder
e.g. Arctic wolves
 Remain active even when environmental temperatures drop as low as -50°C
 Thick fur coats keep their bodies warm
 By adjusting blood flow through counter current exchangers and other vessels in the legs,
wolves can keep their foot temperatures just above 0°C – cool enough to reduce heat loss but
warm enough to prevent frostbite
Conformers Regulators
Disadvantage – cells within the body are subject Disadvantage – costs energy to maintain a

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to changes in the conditions surrounding them constant internal environment
Advantage – avoids energy costs of keeping Advantages – cells function in steady internal
internal environment different from external conditions despite changes to external
environment environment
Energetically cheap
Tooerance/resistance
 The range of any specific environmental variable that an animal can survive is called its range
of tolerance
 Extremes of the ranges of tolerance are ranges of resistance, in which the animal does not
die immediately but will eventually do so if it stays exposed to these conditions
TOLERANCE
 The amount of change in the internal environment (brought about by changes in the external
environment) that an organism can withstand
Extreme temperatures and thermal tolerance

 All organisms have a range of tolerable body temperatures
o Homeothermic endotherms – narrow range
o Poikilothermic ectotherms – broad range
 Exceeding limit of thermal tolerance results in death
How does this affect populations?

 The distribution of a species will be controlled by that environmental factor for which the
organism has the narrowest range of tolerance
 Organisms that have a wide range of tolerance for limiting factors are likely to be widely
distributed
RESISTANCE
 After tolerance levels are exceeded, an animal can resist changes in the internal environment
for a certain period, depending on how greatly the tolerance levels are exceeded but will
eventually succumb leading to death
Hormones, Receptors, Pheromones

Types of communications between ceoos
 Neural
 Neuroendocrine or neurosecretory
o Only really seen in the hypothalamus in the pituitary
 Endocrine
o Situations where an endocrine cell produces a chemical to act locally on the same
cell (autocrine)
LOCAL AUTOCRINE AND PARACRINE

1. Autocrine secretions affect the cell they were secreted from e.g. cancer cells produce growth
factors
2. Paracrine secretions affect neighbouring cells e.g. cytokines which control the process of
angiogenesis (formation of new blood vessels)
 No release into blood stream
Comparison of systems for cell-to-cell-communication
Feature Autocrine/Paracrine Nervous Endocrine
Secretory cell Various Neuron Endocrine
Target cell Most cells in the body Neuron, muscle, Most cells in the body
endocrine
Signal type Chemical Electrical and Chemical
chemical
Maximum signalling Short Long i.e. muscle Long
distance length

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Transport Extracellular fluid Synapse Circulatory system
Speed Rapid Rapid Slower
Duration of response Short Short Longer
Exocrine goands
 Releases secretions into a duct system
 Usually aqueous mixtures (watery), consisting of water-based primary fluid and added
components rather than a single substance
e.g. sweat glands, salivary glands, prostate gland, cells that produce pheromones
Exocrine secretions: pheromones

 Specific chemicals produced (by exocrine glands) by one animal to communicate with another
 Released into the environment and act as a chemical language
 Initiate a range of physiological responses (attracting/selecting mates, controlling behaviour,
signalling the presence of a predator)
e.g. Clams – spawning of eggs and sperm is triggered by pheromones that are released at the same
time as the gametes
e.g. Silkworm moths – females release a pheromone called bombykol which acts as a potent sex
attractant
Endocrine goands
 Organs that secrete substances into blood stream directly – no ducts
 Consist of specialised cells which all secrete similar substances = secretory cells
o Secrete into blood stream
Chemicao signaooing in endocrine system

 Two main types of chemical messengers:
1. Hydrophilic messengers dissolve in aqueous solutions like
extracellular fluid and blood e.g. peptide hormones
2. Hydrophobic messengers bind to carrier proteins in the blood carrier
protein – help hydrophobic messengers dissolve in aqueous
solutions e.g. steroid hormones
Main groups of chemicao messengers

 Amines: catecholmines, thyroid hormones, small molecules derived
on amino acids e.g. noradrenalin, adrenalin
 Peptide and protein hormones: make up the largest group of
hormones, most widely distributed throughout the body e.g. vasopressin, insulin
 Steroid hormones: cyclic hydrocarbon derived from cholesterol e.g. testosterone, oestrogen
Synthesis of peptide hormones

 Peptides are made and stored and then released when
required
o mRNA read by ribosome and string of amino acids
are formed
o String is packaged in granules and move into golgi to
be stored in vesicles
 A signal will trigger the endocrine gland resulting in vesicles
exocytosing at the cell membrane
 Require machinery to move across membrane due to
hydrophilic nature
Synthesis of steroid hormones

 Synthesized from cholesterol
 Cells take in cholesterol and enzymes within the cell convert it into steroid hormones

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 Different steroidogenic cells have different sets of enzymes so each produces a different end
product
 Steroid hormones are not collected in vesicles before secretion – they are made on demand
when the cell is stimulated and are immediately secreted (not stored)
 Secretion occurs via diffusion across the cell membrane due to hydrophobic nature
Ligand-receptor interactions
 Only the correctly shaped natural ligand can bind to the receptor
 Ligand mimics e.g. drugs and poisons
o Agonists – activate receptors
o Antagonists – block receptors
e.g. cone snail – natural antagonist example
 In a normal fish, when a nerve cell releases acetylcholine (ACh),
the muscle cell receptors bind the Ach
o This activates the receptors and the muscle contracts
 In a poisoned fish, the alpha-conotoxin “blocks” the muscle cell
receptor and prevents the ACh from binding
o The ACh receptors cannot function, and the fish is
paralyzed
o Antagonist effect
Hormone action on target ceoos: neurotransmitters

 Ligand-gated channels: receptor and ion channel
o Function mainly in the transmission of nerve impulses across synapses
o Signal molecules are neurotransmitters
 NT ligand binds to the receptor in the cell membrane
o This causes the ion channel to open to permit ions (sodium, potassium) to pass
through the membrane
o The flux of ions changes the electrical charge across the membrane
Hormone action on target

ceoos: peptides and
catechooamines
1. Lipid-soluble (hydrophilic) hormones cannot penetrate the plasma membrane and therefore
bind to cell-surface receptors.
 This binding often leads to the stimulation or one or more second messenger systems which
combine with other molecules to produce a metabolically active complex
 One of the most important second messenger systems involves cAMP
2. These second messenger systems mediate rapid, short-term responses
Hormone action on target ceoos: steroids and thyroid hormone

1. Lipid-soluble (hydrophobic) hormones bind to
cytoplasmic receptors and are transported to the
nucleus

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 Hormone receptor complexes in the nucleus and cytoplasm act directly on the DNA of the cell
to cause long-term changes lasting hours or days
2. Once the steroid dissociates from its carrier protein, it can readily enter the cell by diffusing
across the plasma membrane
 Within the cell, steroids being specific receptor proteins in the nucleus (i) or cytoplasm (ii)
o The resulting hormone-receptor complex then binds to regulatory elements in the
DNA and causes gene transcription
o Because the cellular response to steroids involves the production of proteins, their
effects often last for hours or days
Hormone action reguoation by receptors

e.g. Prairie vole and Montane vole
 Prairie voles are monogamous and montane are promiscuous
 In prairie voles, vasopressin facilitates parental bonding – more likely to stay together
 In montane voles, vasopressin had no effect on pair bonding
 Prairie voles have many more receptors for vasopressin whereas montane were not affected
due to lack of receptors
DIFFERENCES IN DISTRIBUTION OF VASOPRESSIN (V1a) RECEPTORS IN BRAIN

 Expressed at higher levels in the ventral pallidum of monogamous than promiscuous vole
species
o Key brain region involved in reward, motivation and addiction
 Receptor location and density can affect the function of the chemical messenger (peptide or
steroid)
Stress Response
Short term (Fight or flight)
 Increased HR, contractility and systemic BP, dilation of
bronchioles and blood vessels in skeletal muscles
Reguoatory mechanisms
 Chromaffin cells – adrenal medulla
 Corticotrophin released hormone (CRH) neurosecretory cells – hypothalamus
 Anterior pituitary and adrenal cortex
Adrenao goand
 Developmental origins: ectodermal tissue and endodermal tissue
 Adrenal medulla:
o Chromaffin cells
o Derived from ectoderm
o Produces catecholamines (noradrenalin and
adrenalin)
 Adrenal cortex:
o Derived from mesodermal tissue
o Produces steroid hormones (GCs and aldosterone)
o Blood glucose regulation and osmoregulation
Adrenao meduooa
STRUCTURE/FUNCTION OF CHROMAFFIN TISSUE
 Cell body is in the spinal cord
 Axon runs to the adrenal medulla tissue
 Synapses onto chromaffin cell
 Action potential results in influx of calcium and NTs are
released via exocytosis (granular vesicles of NA and A)
 Endocrine cell? Neuro? Neuroendocrine?
 Neurally-derived tissue – acts as post-synaptic ganglion cell
Receptors for catecholamines

 Adrenoreceptors – in plasma membrane of target cells

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 ß1 receptors are located in the heart, they cause an increase in HR and strength and are
sensitive to norepinephrine and epinephrine
 ß2 also found on smooth muscle blood vessels of muscle
Feedback mechanisms – catecholamines

 ATP is stored in the granules of chromaffin cells and is released along with the catecholamine
o Inhibits further release of catecholamines by reducing calcium influx, thereby
providing –ve feedback control
 A second –ve feedback loop involves 2 receptors which are located in presynaptic cells at
noradrenergic synapses
o They also cause inhibition of NA release via an inhibitory effect on adenylate cyclase
o These receptors are part of the –ve feedback loop in which the release of NA inhibits
further release of NA
o This is also called auto-inhibition
 Catecholamine action can be modified by changing the density of the adrenoreceptors in the
plasma membrane of the target tissue
o Continual exposure to catecholamines can lead to
down-regulation in receptor concentration, and
therefore to a decreased responsiveness to the
hormone
Note: rapid inactivation of NA/A by catechol-o-methyl transferase
(COMT) in the liver and kidney to reduce effect
Long term
e.g. teleost responses to a net encounter
 Immediate response – fight or flight response – catecholamine mediated
 Swimming switches from aerobic (red muscle) to anaerobic – lactate increases (O2 debt) and
blood pH increases
 Continued swimming and contact with net induces additional stress – cortisol levels increase
 This involves the adrenal cortex
Adrenao cortex
 Zona reticularis – glucocorticoids, but also precursors to sex steroids
 Zona fasciculate – glucocorticoids
 Zona glomerulosa – mineralocorticoids e.g. aldosterone
 Neuroendocrine cells in hypothalamus which lead down to

anterior pituitary
 CRH is released into anterior pit. which then releases ACTH
 ACTH is released into blood stream and acts on adrenal gland
o Releases glucocorticoids
 Negative feedback acts to prevent production of CRH and ACTH
GCs increase the availability of quick energy to muscle and nervous

tissue
 Muscle – cause muscle cells to release amino acids into the circulation
o This increases amino acids availability for conversion into glucose in the liver
o Net result is maintenance of adequate blood glucose levels to sustain energy
production in critical tissues
 Fat cells – stimulate the mobilisation of fatty acids from
stores in adipose tissue
 Liver – promote synthesis of glucose from amino acids and
fats

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o Some newly synthesized glucose is stored as glycogen in the liver and muscle but
most is released into the circulation, causing a rise in blood glucose levels
Environmentao effects of stress

(Noise)
 Fish are often exposed to environmental sounds such as those
associated with shipping, seismic experiments, sonar and/or aquaculture
pump systems
 Goldfish were reared in quiet (110-125 dB) or noisy (white noise, 160-170
dB) conditions and cortisol/glucose levels were measured
 Regardless of rearing noise exposure a transient spike in plasma cortisol
within 10 min of noise onset, which reduced if noise persisted for 60 min
Long term effects of stress

e.g. Cheetahs in zoos reproduce poorly and have high prevalence of unusual diseases that cause
morbidity and mortality.
 These diseases are rarely observed in free-ranging cheetahs but have been documented in
cheetahs that have been captured and held in captive settings either
temporarily or permanently
 Therefore, captivity may be stressful for this species and contributes to
poor health and reproduction
 This study shows that chronic stress caused a significant reduction in
testosterone metabolites (correlated with increased cortisol levels)
o Decreased male fertility
SUMMARY OF RESPONSE TO STRESS BY ADRENAL GLAND

 A predator has been detected in the external environment, so relay information is sent to the
brain which then activates the autonomic nervous system to send neural information to the
chromaffin cells in the adrenal medulla (effector)
o The effector sends output information (catecholamines) to all the tissues and organs
in the body which need to respond to the detection of a predator e.g. increased HR,
increased blood flow
 A second system is also activated
o Information is sent to the brain to send neuroendocrine and endocrine information to
activate the adrenal cortex (effector) to release glucocorticoids
o They induce gluconeogenesis, while reducing glucose uptake by peripheral tissues
o They ensure that tissues requiring glucose for metabolism have a steady supply of
fuels
o Glucocorticoids cause mobilisation and synthesis of glucose = metabolic stress
response
 Both systems are regulated by negative feedback
Osmoregulation
 Regulation of immediate “internal” cellular environment made up of water and ions
Osmosis
 Osmotic pressure – force associated with the movement of
water across a biological membrane
 Osmolarity – ability of solute to induce water to cross a
membrane
OSMOLARITY
 Net movement of water across a cell membrane can affect its shape
(tonicity)
 Equal concentration of solutes = isotonic
 A solution which has a higher concentration of solutes = hypertonic
o Water will move out of the cell
 Osmotic regulation = control of tissue osmotic pressure which
determines movement of water across biological membranes
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 Ionic regulation = control of ionic compositions of body fluids

 Solutes and water are exchanged between internal body fluids and between external fluids
COPING WITH IONIC/OSMOTIC CHALLENGES

 Ionoconformer – exert little control over the solute profile
within the extracellular space; exclusively marine
o Cannot adapt to changes in environment
therefore require relatively stable external
environment
 Ionoreuglator – control the ion profile of the extracellular
space
 Osmoconformer – internal and external osmolarity are similar;
marine invertebrates
 Osmoregulator – osmolarity is constant regardless of the
external environment
 Stenohaline – can tolerate a narrow range of salinity
 Euryhaline – can tolerate a wide range of salinity
Osmotic proboem for terrestriao animaos

 Water conservation is essential, and diet determines whether salts must be conserved or
excreted
 Evaporation from skin
o Permeability of integument varies among groups; can be manipulated in some spp.
 Integument = outer layer
 In urine produced from the kidney
o Necessary to excrete nitrogenous waste
 From the respiratory tract when air is expired e.g. exhalation, panting
 Need to excrete salts
o Especially terrestrial animals who live in a marine environment e.g. gulls, crocodiles
Osmoreguoatory organs
(Kidneys, gills, glands, skin)
KIDNEY
 Preventing water loss in urine
 Cortex is above and consists of convoluted tubules and
glomeruli
 Medulla contains loops of Henle and collecting duct
 Solute reabsorption in different sections of the nephron – results
in water being drawn out
Proximal convoluted tubule (PCT)

 PCT cells are cuboidal with microvilli:
o Increased surface area
 Large numbers of mitochondria near the apical surface – PCT cells are highly metabolically
active
o Pump most of the solutes out (reabsorption)
 The PCT cells actively transport Na+, glucose and amino acids out of filtrate, back into tissue
fluid
 This causes water to follow by osmosis (specifically descending limb)
o Water and solutes in tissue fluid are taken up by peri-tubular capillaries (vasa recta)
Concentration of urine: Loop of Henle

 Countercurrent multiplier: creates osmotic gradient in surrounding medullary
tissue that facilitates transport processes
 Gradients are maintained by vasa recta capillaries which transport
water/ions away
 Causes urine to become concentrated; hypertonic to blood
 Found in birds and mammals
o Essential for “adaptation” for terrestrial life
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 Animals that produce more concentrated urine have longer loops of Henle and thicker
medullas
o Bigger loop = desert animal
Countercurrent multiplication
 Thick ascending limb: actively pumps out Na+ (K+ and Cl- follow
passively)
o This part of the tubule is impermeable to water
 Reabsorption of ions into the surrounding medulla generates a
concentration gradient
 Thin descending limb: impermeable to solutes but permeable to water
o Osmotic pressure causes water to leave the tubule and enter
the hyperosmotic medullary tissue
Reabsorbed solutes and water are taken away

 by the vasa recta
 Na-K-2Cl cotransporter in apical membrane
(active transport of Cl-, K+, inward movement of
Na+ due to electrochemical gradient – passive)
 Active transport of Na+ driven by N-K-ATPase in
basolateral membrane (energy from ATP)
o Passive movement of K+ and Cl- in
direction of electrochemical gradient
Distal convoluted tubule (DCT) & Collecting duct

 Active transport of Na+ out of the tubule fluid
 Elective water permeability (in later sections of DCT)
 Water is drawn into surrounding tissues by osmosis
 Collecting duct: hyperosmotic medullary tissue draws water out of the collecting duct by
osmosis – reabsorption
Aquaporins
 The osmotic concentration of the urine depends on the permeability of the nephron (tubule)
and collecting duct to water which can be regulated
o Impermeable = dilute urine
o Permeable = concentrated urine
 Water specific channel molecules in plasma membrane
 Speed up rate at which water moves through a membrane
 Does not involve metabolic energy
 Alter permeability of membranes to water
Note: Aquaporins are present in high amounts in the inner medulla/papilla = collecting ducts
Water ooss – amphibians

 Aquatic: no difficulties with water loss (excrete large quantities of dilute urine; all water loss is
via kidneys. Also need to extract salts from the environment)
 Terrestrial: big problems, water loss (evaporation) from the skin; must maintain a moist
surface for gas exchange
1. Limit skin water loss (mucous glands in the dermis)
2. Kidney modified to retain salts (refer to freshwater fish) – no loop of Henle
3. Water storage in the bladder, lymph sacs
4. Ventral pelvic region identified as the area responsible for water uptake, hyper-vascularised
o Flatten body over wet surfaces, spreading skin over moisture
AQUAPORINS
 Kidney
 Bladder
 Skin (ventral pelvic region)
e.g. water gain through the skin of a Bufo Bufo
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VASOTOCIN
 Analogous to vasopressin in mammals
o Increases permeability to water in collecting duct = more concentrated urine
 Receptors for VT in: collecting ducts of kidney, bladder, skin (ventral pelvic region)
 Activation of VT receptors increases permeability to water (increased water uptake)
 Increased expression of aquaporins
 Vasotocin receptors = GPCRs
WIPING BEHAVIOUR
 Lipid secretions secreted from skin glands (exocrine secretion)
 Wiped over skin by legs
 Reduce evaporative water loss to 5-10%
USING UREA
 Stop urine production and accumulate nitrogenous waste as guanine
 Guanine crystallises: saves water and increases skin reflectance (reduces body heating)
Body fouids v environment composition

Osmolarity = osmotic concentration
 Some animals in salt water and some in
fresh water
 Osmoconformer = elasmobranch  lower
concentration of salts x
 You cannot be an osmoconformer in fresh
water
o You would have no concentration
of salts
TELEOSTS
Teleost’s = bony fish
Elasmobranchs = sharks and rays (cartilaginous fish)
Blood concentration relative to Osmoregulatory mechanism

environment
Freshwater teleost Hyperosmotic No drinking; absorbs salt with
gills
Marine teleost Hypo-osmotic Drinks seawater; secretes salt
from gills
Marine elasmobranch Iso-osmotic (but hypotonic) Does not drink seawater;
osmolytes; excretes NaCl from
salt glands
e.g. Freshwater teleost

 The problem
o Movement of water into the body
o Loss of salts to surrounding water – by diffusion
 The solution:
o Prevent the net gain of water
o Remove excess water
o Retain sodium chloride – active process
 Internal environment is hyper-osmotic to the external
environment
 Integument with a relatively low permeability to water and salts
 Don’t drink; get all their water from diffusion across the gills
o No active drinking
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 Kidneys have more and larger glomeruli (improve capability of

filtration and Na+ reabsorption)
 Useful salts are reabsorbed from the filtrate in the proximal and
distal tubules which are much larger to assist in salt recovery
o High amount of filtrate as they want to remove the
water they are taking in from the surroundings
 No loop of Henle (don’t need to conserve water): produce large
amounts of dilute urine
1. Active transport of salts from the external dilute environment into the interstitial fluid and blood
by gill epithelial cells
 Gills have lamellae to increase surface area and thus take in more water
2. Gill epithelium has “ion pumping” cells which mediate uptake of Na + and Cl- ions from the
water (active – costs energy)
o These cells have transporters on the apical membrane and high levels of proton
pumps within the cell
Gills
 Epithelial pavement cells (PNA-, top)
have Na+ channels and a proton pump in
the apical membrane
o Take up Na+ from dilute media
using the proton pump to
generate a gradient which draws
Na+ into the cell
o Proton pumps sets up the electrochemical gradient strong enough to allow Na to
come in via its channel
o Na/K pump allows Na to be pumped into the blood plasma
 K is transported back into the blood from the pavement cell via a protein
channel
 Epithelial chloride cells (PNA+, bottom) have apical Cl-/HCO3-
exchangers
o These cells import Cl- from dilute media using the
Cl-/HCO3- exchanger, which requires production of
bicarbonate ions (HCO3-) and H+
o Bicarbonate leaves setting up an electrochemical
gradient strong enough to allow Cl to come in via its
channel
o Side cost of importing Cl = acidity
e.g. Marine teleost

 The problem
o Loss of body water to environment
o Net gain of salts
 The solution
o Prevent the net loss of water
o Remove excess salts
 Internal environment is hypo-osmotic to external environment
 Lose water continuously across gills – exposed, not covered by
scales
 Drink seawater to replace the water lost to the environment
 Don’t produce much urine (kidney has a reduced role in ion and water excretion)
 Nephrons may have small or absent glomeruli (aglomerular) or
lack Bowman’s capsules: don’t need to filter out salts
o Need to get rid of salts through gill epithelium
 Proximal and distal tubules are also reduced in size
 Eliminate ingested salts by active transport mechanisms in the gill
epithelium – pumping against concentration gradient (active)
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1. Active transport of salts out of the body and into the external environment
 Involves chloride cells; they actively transport salts from the blood into the environment
across gill epithelium
Gills
 Chloride cells with high levels of Na+/K+ pumps (driven
by Na+/K+/ATPase) associated with Na+/K+/2Cl-
cotransporter in the basolateral membrane
 Activation of this pump results in the net movement of
Cl- from the blood into the cell
o Chloride then diffuses from the cell into the
external environment via Cl- channels in the
apical membrane
o The trans-membrane potential created by this
movement increases the Na+ electrochemical
gradient so that Na+ can diffuse out of the cell
(through) paracellular channels even against high Na + gradient
 Na is being pumped out of cell into blood (energy requiring), K is being pumped passively and
as Cl is pumped into cell from blood via Na cotransporter
o A transmembrane electrochemical gradient has been set up and Cl is pumped out of
cell into surroundings
ELASMOBRANCHS
 Plasma osmolarity similar to seawater = iso-osmotic but
they are hypotonic
o Much less salts than surrounding environments
 Keep body fluid osmolarity (but not ion content) similar to
environment: don’t have large osmotic gradient (there is no
net gain or net loss of water)
 They don’t drink seawater – osmolarity is the same
 Osmotic gap is filled by an organic osmolytes (e.g. urea or
trimethylamine oxide (TMAO))
o Ensures isosmotic water balance is maintained between body fluids and environment
 Excrete less salts via the kidneys and a specialised organ called the rectal gland
 Advantages – energy saving (no need to drink) and don’t need to spend energy pumping ions
 Disadvantages – necessity for urea tolerance and the energy cost of producing urea
Are they osmoconformers or osmoregulators?
e.g. Bull Shark, Carcharhinus oeucas

 The bull shark lives many different areas and travels long distances
 It is common in coastal areas of warm oceans, in rivers and lakes, BUT it is also able to
inhabit fresh water
 Bull sharks living in fresh water reduce the concentration of solutes in their blood as they
move upstream by up to 50%
 How do they alter their blood plasma osmolarity in different environments?
o In brackish/dilute waters, they lower their blood urea concentrations by decreasing
urea synthesis and retention
Rectal gland (exocrine gland)

 Consists of many tubules surrounded by capillaries
 Epithelial cells line the tubules
 Transfer NaCl from blood to the tubule lumen
 Work like chloride cells in the gills
 Central canal lined with an epithelium
How does the rectal gland excrete salts?

 Tubular epithelial cells actively transport Cl- from the blood via
transcellular transport
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o Occurs through Na+/K+/2Cl- cotransporter in basolateral membrane

o Cl- leaves cell via Cl- channels in apical membrane (driven by Na+/K+/ATPase)
 Na+ moves between tubular epithelial cells via paracellular transport
 Same process as in gill epithelium
Excreting excess saots

 Reptiles and birds can drink seawater but have to excrete the excess salt: salt glands (nasal
glands)
 Glands consist of many lobes, each of which contains many branching secretory tubules that
release secretions into a central canal
 Located above the eye; ducts carry salt glands secretions to the nasal passages and the
secretion drip out of the external nares (nostrils)
Nasal gland (exocrine gland)

 Active secretion of salt takes place across the
epithelium of salt-secreting cells in the tubular
epithelium of the nasal gland
 High concentrations of a Na+/K+ pump are
found in these tubular cells (work like chloride
cells)
Migrating fish
 Adult salmon migrate between seawater and freshwater to breed: anadromous fish
 Young are born in freshwater but migrate to the sea
o Prior to migration, their gills undergo a dramatic change as the ion-pumping
properties of the gill prepare for the new environment
 This physiological acclimation takes a few days to occur once the fish changes environments
o Freshwater to seawater (smolting)
o Change colour to silver
1-3 years in freshwater (increasing salinity):

Smolts have increased chloride cells in the Smolts have increased Na+/K+/2Cl- in chloride
primary filaments of gills cells (active transport of salts out of the body)
 Seawater tolerance of juvenile Atlantic salmon was determined by

measuring their ability to regulate plasma ions in 24hr seawater
challenge tests.
 Acclimation (making changes, not developing new adaptation) to
increased salinity occurs within 24hr:
o Increased number of chloride cells
o Increased Na+/K+/2Cl-
o Increased Na+ATPase activity
Endocrine regulation of smolting

 Cortisol plays a major role in the physiological acclimation
associated with migration to seawater by causing changes in the
gill structure
 Increased cortisol and glucocorticoid receptors in the gill
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epithelium when fish smolt

 Cortisol induces an increase in the number of chloride cells
Smolting from freshwater – seawater

 Rising external Na+ concentrations causes a flux of Na+ into the body; raising plasma Na+ and
stimulating an increase in plasma cortisol
 Cortisol causes an increase in the number of chloride cells (increasing Na+/K+/2Cl- co
transporters and Na+/K+ pump activity) in the gill epithelium
 Increased Na+/K+/2Cl- cotransport pumping of ions out of the blood and into the chloride cell
 Increased Na+ATPase activity (increased Na+/K+ pump activity) pumps ions out of the chloride
cells into the environment
Smolting from seawater – freshwater (Smolting is reversible)

 A decrease in external Na+ levels causes the paracellular gaps between chloride cells and
accessory cells to close, preventing loss of Na+ and Cl-
 Plasma prolactin levels increase
 Prolactin functions either by reducing Na+/K+ activated ATPase activity or reducing the density
of chloride cells
 As a result, the activity (and/or the density) of the Na +/K+ pump decreases to prevent loss of
salts from the body fluids
Ecophisiologi
Ecophysiology = animal behavior + ecology + physiology
Founded by:
 Georgie Bartholomew (1923-2006)
 Knut Schmidt-Neilson (1915-2007)
Thermao physiooogy
 Temperature = measure of intensity of molecular movement
o Lowest air temperature on earth? -89.2ºC at Vostok Station, Antarctica
o Highest air temperature on earth? 56.7C at El Azizia, Libya
 Heat
o Thermal energy: increased thermal energy leads to increased movement of
molecules
o Object receiving heat: temperature increases
o Object losing heat: temperature decreases
o Objects at same temperature: equilibrium
Thermal Relations
Thermao energy and physiooogy
 Only proteins and lipids are substantially affected by temperature over normal ranges
encountered by animals
 Enzymatic reactions
o High temperatures – increase rate of activity (up to a point)
o Low temperatures – decrease rate of activity
 Protein conformation – van der Waal forces
o Increased temperatures – disrupts protein conformation
o Decreased temperatures – stabilizes protein conformation
 Lipid bilayer fluidity – van der Waal forces and hydrogen bonds (must be in very close contact
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for van der Waal forces to take effect

o High temperatures – increase membrane fluidity
o Low temperatures – decrease membrane fluidity
o Membrane fluidity affects protein movement
THE THERMAL NICHE

 Can survive outside optimal zone but risk death
Body temperature survivao oimits

e.g. cane toads (Rhineooa marina, Bufo marinus)
 Survival: 5-40ºC
 Breed: water temperature at 25-30ºC
 Disturbances of cell membrane function such as loss
of selective permeability and transport processes
 Na+/K+ pump action is 0.8% optimum at 5ºC
Body temperature and performance

e.g. cane toads
 Barely any hopping below 15ºC - if in the wild, this would mean death as it
cannot escape predators
 Measured because it is ecologically relevant
 Warmth makes the toads faster
Thermao niches and survivao
THERMAL ENVIRONMENTS
 Animals occupy varied thermal environments
requiring them to utilize different and varied
thermal strategies to maintain homeostasis
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Relations of nutrition, feeding, digestion and absorption in

vertebrates and most invertebrates:
 Break down using its own enzymes or through help of
bacteria (fermentation)
o Mutualism between microbes and animals
 Nutrients are distributed through networks such as blood
vessels to tissues and cells
Composition of human body

 Ranking of importance
 Humans are predominantly made up of protein whereas plants
are predominantly made up of carbohydrates
PROTEIN
 50% of animal organic matter (non-mineral dry mass)
 Roles:
o Enzymes – speed and regulate biochemical reactions
o Tissue proteins – muscle (locomotor), collagen and keratin (structural), eye lenses
(crystalline), oncotic (plasma proteins)
o Receptor, channel and transporter proteins in cell membranes e.g. haemoglobin and
myoglobin
o Oxygen transporting/storing proteins – haemoglobin/myoglobin
o Eggs and other reproductive materials
o Antibodies, hormones and materials
 Unlike fats or carbohydrates, protein is not stored long-term so it must be acquired regularly
 Proteins are made of strings of amino acids
LIPIDS
 Mostly comprised of fatty acids
 Readily stored as adipose tissue
 Roles:
o Primary component of membranes
o High density energy storage
o Integument (skin) waterproofing
o Steroid hormones
o Triacylglycerol’s – dolphin echolocation
 Some essential lipids e.g. omega 3 and 6
CARBOHYDRATES
 Most abundant nutritional compounds
 Simplest: monosaccharides
 Roles:
o Structural – cellulose and glycogen
 Difficult to access
o Energy storage – starch and glycogen
 Liver (6-8%) and muscle (1-2%)
o Transport – plasma and milk
 Highly soluble
 Energy for brain and muscles
 There are no essential carbohydrates
VITAMINS & MINERALS

 Trace amounts required
 Almost all are essential and cannot be synthesized
 Vitamins:
o Organic molecules
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o Incorporated into key molecular subsystems e.g. coenzymes

 Minerals:
o Often needed for synthesis of complex molecules e.g. Iron in haemoglobin, calcium in
bones
Feeding and digestion

 Once an organism has found the food it requires, a key process is digestion
o Breaking down more complex molecules into simpler molecules to be absorbed
o Egestion – faecal material which consists of some of the
unused nutrients and smaller molecules which have
been digested
 Physical and chemical processes required with some help from
microbes
DIGESTION
 Breaks course food items into smaller parts to be absorbed and
distributed – physical and (bio)chemical reactions
 Achieved by complex gut morphology
1. Batch reactor
o One meal goes in and is digested and
excreted then another one does the same
2. Continuous-flow reactor without mixing
o Pass material through and meals follow one
another
3. Continuous-flow reactor with mixing
o Mixture of all things being eaten coming out
Insectivores and carnivores

 High protein/fat diets
 Caecum – stores food for bacterial breakdown of cellulose
 Absent or vestigial in insectivores, omnivores and
carnivores
Non-ruminant herbivores
 Hindgut fermenters e.g. rhinos, apes, koalas, some rodents, ground
dwelling herbivorous birds
o Digestive system longer
o Fermentation can occur in caecum
o Coprophagia – eating your own faeces
o Fermentation occur in expanded colon
 Midgut fermenters e.g. herbivorous fish (carp, catfish and tilapia)
Ruminants and other foregut fermenters

 Ruminants – 4 chambered stomach e.g. sheep, deer, cattle, giraffes, camelids, etc.
o (Non-acidified) rumen
 Produces enzymes for cellulose
 Synthesises B vitamins
 Recycles waste N into new protein
 Food is consistently regurgitated to digest more than
once – repeated chewing and enzymes allow particles to
filter through to the abomasum
o (Acidified) abomasum
 Physiochemical (acid) digestion
 Biochemical digestion (enzymes) e.g. amylases
o Small intestine
 Biochemical digestion and absorption
o Colon
 Further breakdown and/or water absorption
 Non-ruminant foregut fermenters e.g. kangaroos, hippos, sloths, colobus monkey
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Nutritionao ecooogy: Geometric framework

 Nutrient space: gain in nutrients of an organism who
hasn’t eaten
o Somewhere is the space is the right balance
of nutrients for the organism (target intake)
o Animal can choose certain animals/plants to
eat to get to the target
o A particular food you can put in nutrient space in terms of its
composition – if it is on the same line as the target (nutritional rail),
the animal can survive on the food
1. Amount of nutrients
2. Balance of nutrients
Imbalanced Complementary
 Eating a food too high in one  Eat too much carbohydrates so try to
macronutrient causing you to miss compensate by eating too much protein in
target order to weave your way to target
Conservation physiooogy
 Must be able to provide ideal diet to animals in captivity for conservation purposes
 2013 the Conservation Physiology journal was created
 New area in the field of animal physiology
e.g. Koalas and climate change

 Koalas have a Tcore of 35.7ºC
 The environment of the koala changes but the body
temperature stays the same
o Increase metabolic rate when they’re cold and
increase water loss when hot
o Will have a certain extra cost of energy when
they are in a hot or cold environment
 Calculated what the koalas’ profit is of water and energy
at many locations across Australia to work out if they
would have enough energy to reproduce
o Results showed that they could reproduce only if they were consuming highly watery
leaves and increased their diet/seek shade
e.g. Saving the Western Swamp Turtle

 Only lays 2-5 eggs every couple of years
 Live in wetlands that dry out every summer and fill up every winter
o In winter they are very productive and grow quickly due to access to lots of food
 Winter
o Tortoises active if daylight, water available, Tb > 14ºC and < 28ºC (in water or basking
on surface)
 Late spring
o Tortoises enter terrestrial habitat if swamp dries, or water temperatures > 28ºC
 Early summer
o Turtles and their eggs spend the entire summer in burrow
o Produce clutch, burrow to Tp or die if Tb exceeds 42.5ºC
 Late autumn
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o Exit aestivation site/nest and enter water when water depth > 5cm
o When the rain comes, the baby turtles hatch
o If the water is not around for long enough, the babies can die
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Physiological Systems
Cardiovascular Sistem
Functions
To maintain homeostasis…
 Rapid convective transport of nutrients and O2
 Rapid removal of waste products
 Distribution of H2O, electrolytes and hormones
 Tissue fluid production
 Immune system infrastructure
 Temperature regulation
 Reproduction (feed-forward not homeostatic)
Oxygen transport
 Gas exchange = O2 ↔ CO2
 Oxygen cascade: air to mitochondria
 Diffusion of oxygen and carbon dioxide through gas
exchange surfaces
 Continue falling of gas concentration as you move
from air to cells (cascade)
Boood & haemooymph

 Closed cardiovascular system: blood
 Open cardiovascular system: haemolymph (everything mixed together)
 80% water
 Solutes and ions ≈ interstitial fluid
 Proteins  interstitial fluid
o Decapod crustaceans = 10-90g/L
o Vertebrates = 30-80g/L
o Cephalopods & molluscs = 110g/L
 Invertebrates – many proteins are respiratory pigments
 Vertebrates – albumin, globulins and clotting factors, not respiratory pigments
Respiratory pigments
 Involved in gas transport
 Metalloproteins that bind reversibly with O2 at specific O2 – binding sites associated with metal
atoms
 Haemoglobins (Hb): All vertebrates (>9 phyla of animals)
o Fe (heme group): binds 4 O2 molecules
 Haemocyanins: Found in arthropods and molluscs
o Cu: binds 1 O2 molecule
o Turn blue when carrying oxygen
 Haemoerythrins: some families of worms and brachiopods
o Fe (no heme group): binds 1 O2 molecule
 Chlorocruorins: 4 families of marine annelid worms
o Contain a unique iron porphyrin in a weak heme group
Red boood ceoos (RBCs) – erythrocytes

 Vertebrate Hb is always contained in RBCs
 Vary in size shape and other properties
o Mammals: no organelles, nucleus, mitochondria or ribosomes
o RBCs: in all other vertebrates and even other types of pigment containing cells in
invertebrates are nucleated
Vascuoature
 Intima: internal elastic lamina and endothelial cells line in the lumen
 Media: external elastic lamina and vascular smooth muscle
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 Adventitia: collagen and proteoglycan rich exterior provides support/stability

 Structure related to function
 Different vessels have different layers or different layer thickness depending on function
 Each part of the systemic vasculature has a unique anatomy
and function
o Capillaries very thin with no media (smooth muscle)
 Pulmonary circulation = throughout lungs
 Systemic circulation = throughout body
Lymphatic circuoation
 Collects fluid (lymph) ‘filtered’ from blood and returns it to the
circulation
 Lymph fluid = clear – no RBCs
 Runs in parallel with the cardiovascular system
 Vital part of the immune system; prevents oedema
 Lymph nodes = areas with high concentrations of
lymphocytes – a reserve for when the immune
system needs backup
Efficient circuoatory systems

With regard to efficiency of gas exchange, the most efficient have:
 Different pressures in different circulations
 No mixing of oxygenated and deoxygenated blood – optimum gas exchange
o Usually consequence of divided heart and/or separate pulmonary and systemic
circulation
 But not all animals need the most efficient circulation or experience situations where they
need an alternate (less efficient) circulatory configuration
o If you have a low metabolic rate, you have low need for oxygen
Circuoatory systems
 No circulatory system
o Sponges and cnidarians
o Move external fluid via bulk flow
through an internal cavity
 Open circulatory system
o Most invertebrates
o Heart/ostia and vessels
o Haemolymph leaves the open vessels
o Haemolymph = blood, lymph and interstitial fluid
 Closed circulatory system
o Virtually all vertebrates
o Some invertebrates (cephalopod molluscs and some annelid
worms)
o Heart(s) and entirely enclosed blood vessels
OPEN
 Haemolymph pumped by ostia
 Vessels (open) distribute blood around the body
 Haemolymph leaves vessels, bathes tissues and drains back to ostia
 Ostia create pressure to cause flow:
 Suspensory ligaments hold ostia in place
 Systole: ostia contracts, which closes the valves and forces fluid into vessel
o Suspensory ligaments contract to close ostia and recoil to open up
ostia and allow influx of blood
 Diastole: haemolymph drains into ostia via valves
 Movement and muscle contraction helps return haemolymph from the tissue to the circulatory
system
 Insects – blood doesn’t carry respiratory gases
o They have invaginations on the outside of their body coming in
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CLOSED
 Development of a closed circulatory system
correlates to increased metabolic requirements
 Generally, consists of systemic and respiratory
circulation
 Increasing complexity moving diagonally up the
chart
Mammals and Birds

 Most complex
 4 chamber heart
o Compact myocardium with cardiac vessels – myocardium is dense
o 2 atria and 2 ventricles – completely separate at any time
 2 separate circuits run in a series
o Different pressures
 Low pressure for pulmonary system
 High pressure for systemic system
o No mixing of oxy- and deoxygenated blood
 If you send deoxygenated blood to lungs, you will get more efficient exchange of oxygen
 If you send fully oxygenated blood to tissues, tissues will be better oxygenated
Fish (water breathing)

 Single circuit: heart  gills  tissues
o Gills receive high pressure blood, the body receives
low pressure blood and the heart receives
(relatively) deoxygenated blood
o Capillaries receive low pressure blood
 2 chambered heart
o Spongy myocardium; no cardiac vessels
o Fish heart gets blood that is passing through the ventricle –
it is low oxygenated and therefore limits the work that the
heart can do
 Not entirely effective as blood leaves the heart at low pressure –
therefore slow delivery to tissues
 Teleosts: 2 extra structures (sinus venosus and bulbus arteriosus) – increase cardiac output
Fish (air breathing)

 Electric eel and certain fish
o Air breathing organs (ABO) in their
mouth cavity (B)
 Certain catfish and other fish
o ABO in the gut
 ABO usually mixes with venous blood
o ABO circulation = parallel with
systemic circuit
 This O2 transport efficiency but  myocardium oxygenation
Amphibians
 2 gas exchange organs: skin and lungs
o 2 atria
o Ventricle – spongy myocardium; no cardiac
vessels; lack septum
 Heart receives both oxy- and deoxygenated blood;
lung and skin receives mixed blood; body receives
fully oxygenated blood
o Restrict cutaneous circulation and bump all
blood to lungs
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 Selective distribution of blood

o Diving – redistribute blood from cutaneous to pulmonary circulation
o Heart – most oxygenated blood directed to systemic not pulmonary circulation
e.g. bullfrogs – 91% of pulmonary venous blood is channelled to the systemic circuit
 Not a proper septum in hard
Reptiles (non-crocodilian)
o Compact myocardium with cardiac vessels
and some spongy myocardium
o Ventricle incompletely divided into 3 by
muscular ridges and partial septa
o Mixed blood sent to tissue
 Reptiles also distribute oxy- and deoxygenated
blood selectively
 Shunting: ability to bypass either the pulmonary or
systemic circulation
Shunting in amphibians and reptiles

 Non-shunting (non-crocodilian)  Shunting
 When we hold our breath, there is more pressure in the lungs

 If the heart is hungry for oxygen (L – R shunt)
Shunting in crocodiles
 4 chambers with 3 major arteries leaving the heart
 1 pulmonary artery: 2 aortae (one from each ventricle) – joined at the foramen of Panizza
o Left aorta comes out of the right ventricle
 Separation of ventricles means crocodiles only shunt R-L
Cog valve in crocodiles

 Crocodile hearts have a cog valve which has teeth nodules that mesh together to form a tight
seal; under active control of NA
o Left systemic artery coming out of the right ventricle and right systemic artery coming
out of the left ventricle
 Breathing:
o RV pressure remains lower than that of the left systemic aorta and the passive flap
valve at the left systemic aorta stays closed
 During breath-holding/diving:
o Increased pulmonary resistance and closing of cog valve raises RV pressure and
opens the passive flap valve at left aorta
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Suppoying the myocardium with O2

 Myocardial density
 Coronary vessels/lumen
 Spongy myocardium – not dense
o Can’t generate enough pressure
o Oxygen easily flows out – limiting factor
particularly if it isn’t well-oxygenated blood
Chambers of the heart

 Atria
o Thin-walled
o Pacemaker (SAN)
o Pump function (≈30% volume)
 Ventricles
o Generate different pressures
o Aorta (oxygenated), pulmonary arteries (deoxygenated)
 Valves
o Atrioventricular (AV) valves
o Semilunar valves (pulmonary and aortic)
Generation and conduction system

 Myogenic hearts – impulse begins in a muscle or modified muscle cell (all vertebrate hearts,
they are also innervated)
o Receive nervous input
 Neurogenic hearts – impulse generation occurs in nervous tissue e.g. lobsters; needs an
external electrical impulse to contract
e.g. Neurogenic heart of the lobster

 Lobster, crabs, shrimp, crayfish, spiders and scorpions
 Sectioning just after the 6 abolishes contraction
MYOGENIC HEART
 Sinoatrial node (SAN) – modified myocytes
 Autorhythmicity
o An automatic, rhythmic beat
 Spontaneous fluctuations of membrane potential which reach threshold for AP
o Increases in membrane
potential until it reaches a
certain point, and kicks
off an action potential
 Dark = wave of depolarisation
 Starts SA node, action potential
fires
o Depolarisation is sent out in a wave to contract atrial muscles
Electrophysiology of the SAN

 Movement of ions through channels changes the potential of the
membrane
 At threshold, an action potential is triggered
 ‘Unstable’ membrane potential – pacemaker potential
o Because it is leaky, the membrane potential is always
moving up slightly until it reaches threshold
 When you get opening of various channels, ions move through the
membrane which has an effect on membrane potential
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o If movement causes a higher membrane potential, an action potential may be fired

 The rate of decay determines how fast the heart beats (HR)
Electrophysiology of ventricular myocytes

 Rapid onset of AP followed by a ‘plateau’ phase in ventricular
myocytes
 Flat membrane potential (not leaky) until it is told by SA to fire
Cardiac myocyte contraction

 Muscle tension follows AP
 One twitch per AP
 Not a large refractory period (absolute and relative)
o This allows us to maintain the lowest heart rate possible
 Means no tetany – would hole ventricular muscle
o Would push blood through and then stop
Cardiac cycoe
 Atrial systole = atrial contraction
o High pressure
o Blood is pushed into ventricles
o Signal passes down
 Pressure rises within the ventricle
o Blood tries to move back into
atria but valves are shut
 Pressure becomes higher than that of
atrium and pulmonary artery/aorta and
blood is pushed out of the heart
o Through valves – PASSIVELY
CARDIAC OUTPUT
Cardiac output = heart rate x stroke volume
CO (L/min) = HR (bpm) x SV (L/beat)
 In a resting adult (70 kg) ≈ 5 L/min
 In an exercising adult ≈ 20 L/min
 In an exercising Olympian ≈ 30 L/min
 Stroke volume (SV)
o Preload is central venous pressure (CVP)
o Afterload is total peripheral resistance (TPR)
o Contractility is factors which change contractile force
 Heart rate (HR)
o Intrinsic rate
o Hormones
o Nerve activity
DISTRIBUTION OF BLOOD
Haemodynamics – blood flow
 Pressure (P): force per unit area
 Resistance (R): how hard for flow (Q) to occur through
vessels
Q= ∆ P/R
 Q must match tissue metabolic demands (can vary ≈5x)
o Q = CO or flow
 P (and so BP or MAP) needs to stay relatively constant
o ∆ P = (arterial P – venous P)
 A major control of Q must be via changes in R
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o R = TPR
CO = (MAP – CVP) / TPR
CO = MAP / TPR
Circulation – in series
 Q must be equal in each part i.e. aorta, arteries, arterioles, capillaries, venules, veins, vena
cavae
o Or blood would back up
 Pressure drops progressively
 Resistance = sum of all resistances
 Concept of TPR
Rtotal = Rarteries + *Rarterioles + Rcapillaries + Rvenules + Rveins
*Largest contribution to Rtotal is from the arterioles
Circulation – in parallel
 Flow through a particular bed and branch within the bed depends on the
individual resistance
 Blood flow in one organ can be altered fairly independently
 Each organ system gets its own blood supply
o Pressure and flow is set by the arterioles in that section
o Therefore, blood flow to one organ can be modified without affecting
another organs blood flow
e.g. if you start exercising you increase blood flow to muscles and decrease
blood flow to gut
Vascuoature: vooume, veoocity and pressure

 Cross-sectional area increases
o Each individual blood vessel gets smaller but as a
group (e.g. capillary bed), the CS area increases
 Velocity
o Inversely proportional to CS area
o Velocity is lowest in capillaries
 Pressure
o Decreases as CS area increases
o ‘Downstream’ pressure fall
o Low venous P – capacitance vessels
o Velocity picks up again in veins but pressure doesn’t pick up as veins have less
smooth muscle
LARGE ELASTIC ARTERIES

 Aorta and pulmonary arteries
 Tunica media (2) contains large amounts of collagen and elastin
 Windkessel effect: ‘elastic reservoir’
o Distend when P rises and recoil when P falls over a cardiac cycle
o Ensures our blood pressure doesn’t drop down to zero
 Pulsatile flow wave propels blood in ‘bursts’
 Elasticity allows aorta to ‘bulge’ – stores energy
 The ‘rebound’ of aortic wall helps propel blood onward
 Reduces pressure variation over a cardiac cycle
 Maintains pressure (diastolic) between beats
MICROCIRCULATION
 Flow controlled by:
o Local metabolic factors
o Sympathetic nervous system
 Capillary density and/or flow rate varies depending on tissue and/or demand
 Capillary diameter ≈6µm
≈6µm x 10 billion capillaries = 500-700m2
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Capillary structure and function

 Three main types of capillaries:
1. Continuous
o Incomplete tight junctions allow the
passage of small molecules
o Found in skeletal muscle, brain,
skin, etc…
2. Fenestrated
o Numerous pores allowing easy passage of small molecules
o Found in specialised areas: kidney, endocrine glands and intestines
3. Discontinuous or sinusoidal
o Large intercellular gaps/cleft allow protein and even rbc movement
o Found in liver and bone marrow
Capillaries: fluid exchange

 Starling’s Forces: 2 types of forces dictate the capillary fluid exchange: oncotic (osmotic)
pressure and hydrostatic pressure
o Hydrostatic pressure depends on the physical pressure exerted by the blood pressure
(Pc) or the interstitial fluid (Pif)
o Osmotic/oncotic pressure depends on the osmotic pressure exerted by solutes (and
proteins) in the blood (c) or the interstitial fluid (if)
 Capillary hydrostatic pressure: tends to drive water out of
vasculature
 Colloid osmotic (oncotic) pressure of blood: tends to draw water in
from tissues
 Rate of tissue fluid production depends on the balance of Starling’s
forces – generally, ultrafiltration dominates
RESISTANCE AND FLOW

 Arterioles set resistance
 Resistance to flow depends on vessel radius4
Poiseulle’s Law R = 8vL/r4
 R = resistance of the tube
 V = viscosity
 L = length of the tube
 r = radius of the tube
So, flow (Q) = ∆ P / (8vL/r4)
Reguoating the cardiovascuoar system

 What is getting regulated?
o Pressure (index of flow/volume)
o Osmolarity (index of volume/pH/gas content)
 How is this regulation achieved?
o Heart rate (HR)
o Vascular resistance
o Circulatory volume
 What mechanisms are involved?
o Short term (e.g. reflex) and long term (e.g. renal control of blood volume)
o Local (e.g. CO2 vasodilation) and global (e.g. ANS)
VASCULAR RESISTANCE
 Resistance set by vascular smooth muscle (VSM)
o Thick (myosin) and thin (actin) filaments
o Gap junctions between VSM cells: functional syncytium
 VSM altered by: sympathetic nervous system (vasomotor tone)
 Intramural pressure (myogenic response)
 Vasodilator/vasoconstrictor agents e.g. NO, adrenaline (local factors)
VASCULAR SMOOTH MUSCLES
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 Resistance set by vascular smooth muscle (VSM)

o Contraction (tone) controlled by [Ca2+]
o Latch state of prolonged crossbridge
 Formation (energy efficient tone)
MYOGENIC RESPONSE
 Increasing BP transiently distends arterioles which respond by
contracting
 Stretch opens stretch-activating ion channels  Ca2+ enters  VSM
contraction
 Minimises capillary flow fluctuations and irregular tissue fluid exchange
 Decreased P evokes VSM relaxation (vasodilation) by the opposite
mechanism
LOCAL FACTORS
e.g. Metabolic hyperaemia
 Matches local vasodilation to local metabolic rate
 Dependent on metabolite substrate/product concentration e.g. CO 2, acidosis, lactate,
adenosine, K+, hyperosmolarity and hypoxia
  Metabolic activity of tissue  O2, metabolites  arteriolar dilation   blood flow
 Essential in exercising muscle and overcomes other vasoactive actions
 Dilution of metabolite concentration evokes VSM vasoactive actions
 Dilution of metabolite concentration evokes VSM contraction (vasoconstriction) by the
opposite mechanism
VASOMOTOR TONE (ANS & Blood vessels)

Vasculature:
 Sympathetic nerves
o NA  increases VSM [Ca2+]
o 1 adrenoreceptors
o Ongoing vasomotor tone
ANS & the heart

 Parasympathetic (vagal) nerves
o Sinoatrial node
o ACh  decrease HR
o Muscarinic receptors (M2)
 Sympathetic nerves
o Sinoatrial node and myocardium
o NA  increase HR and contractility
o 1 adrenoreceptors
ALTERING HEART RATE

 Autonomic nervous system (ANS)
o PNS
o SNS
 Temperature
 Chronotropic agents
o Positive e.g. adrenaline, nicotine, dopamine, caffeine
o Negative e.g. ACh, -blockers
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Sympathetic effects on HR
Parasympathetic effects on HR
DETERMINANTS OF MAP
‘Normal’ arterial blood pressure
 Systolic: pressure of blood in the vessels when the heart
is contracted
 Diastolic: pressure between beats when the heart is
relaxed
 Measured in millimetres of mercury
 MAP is average arterial pressure over a cardiac cycle; an
indication of tissue perfusion pressure
Pulse pressure = systolic pressure – diastolic pressure
MAP = 1/3 pulse pressure + diastolic pressure
 MAP is the controlled variable in reflex regulation of BP
Baroreflex – short term regulation of MAP

 Beat-to-beat, rapid onset, negative feedback loop
Baroreceptor responses
 Baroreflex has a ‘setpoint’ – can change with
physiological state e.g. exercise or chronic
hypertension
Detecting a sudden fall in BP
Baroreceptor reflex and posture
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 The baroreflex is essential for normal, moment-to-moment stability of blood pressure

 Dogs in which baroreflexes are eliminated exhibit much larger decreases in blood pressure in
response to postural changes than do dogs with intact baroreflexes
Diving reflex
 Coordinated series of CVS changes elicited when diving animals and birds undergo forced
submersion
 Stimulated by breath-hold and cold (water) activation of trigeminal (ganglia) receptors
 Diving bradycardia
o Rapid onset
o Vagally-mediated (parasympathetic – ACh causes fall in HR)
 Regional vasocontriction (particular vascular beds)
o Sympathetically-mediated
o Reduced flow to viscera, body wall, limbs and skin
o Maintained flow to brain, lung and myocardium
o Creating a smaller arterial system
 Matches CO to reduced circulatory dimensions
o ‘Smaller’ arterial system
o Reduce CO (by reducing HR)
o Prevents increased MAP
o Reduces metabolic load on the heart
 Graded effect on HR during voluntary submersion
 Vasoconstriction and CO harder to record in freely behaving animals
 Similar graded effect
e.g. Freely diving grey seals, Scotland

 Basal non-diving HR 119bpm
 Fish – similar to bradycardia when exposed to air

 Response to asphyxia (hypoxia)
o Preparation for low oxygen
 Conserved, but variable – many other animal groups, including
mammals, birds, reptiles, amphibians
Respiratori Sistem
“The Fire of Life”
 Cardiovascular system and respiration system work very close together
 Respiratory gases are essential for maintaining life:
o Getting oxygen (fuel) to supply the tissue
o Remove carbon dioxide (waste) from the tissue
Oxygen cascade
Getting O2 from the environment Ventilation
  Exchange between animal and environ.
To the blood Pulmonary diffusion
 
To the tissue Circulation = distribution between
exchange sites
 
To the mitochondria Tissue diffusion = exchange between
tissues and blood

Tissue metabolism =
consumption/production
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 Essentially how oxygen from the environment gets

transported around the body via the transport medium
o Cascading = getting lower i.e. level of oxygen
(partial pressure)
o Things get lost, used, barriers
Partiao pressure of gas

Daltons law of partial pressure:
Total pressure = sum of all partial pressure
 The box is full of air
 There is a composition inside the box
o Composites of air = gas
o They bounce around in the compartment – they
exert pressure on surroundings
 Everything has to add up to 1atm
o All different partial pressures of each
component in the compartment must equal to
1atm
 Partial pressure determines what way the gas is moving
PARTIAL PRESSURE OF GASES IN SOLUTION

Henry’s law: [G] = Pgas x Sgas
[G] = concentration of dissolved gas
Pgas = partial pressure of the has above the liquid
Sgas = solubility of the gas
 At equilibrium: partial pressure of a gas dissolved in a liquid = partial
pressure of that gas in the gas phase above the liquid
 Diffusion of gases is determined by the partial pressure of the gas
above the liquid and the solubility of the gas
 Pressure different between the gas and the liquid determines which
way the gas will diffuse
 Less oxygen in water
Concentration of gases in air vs water

 The capacity of water to hold CO2 is similar to air
o From respiratory point of view we assume this is
identical
 The capacity of water to hold O2 is much lower than air
Factors affecting gas concentration:

 Solubility
o Explains difference between blue line and dotted line
 Temperature
o As temperature increases, capacity to hold oxygen
decreases
 Salinity
o Seawater has lower capacity than freshwater to hold
oxygen
e.g. In a rock pool
 Temperature will go up and oxygen level gets lower
 Will be replenished when tide comes up again and refills the rock pool
Why do animaos breathe?

Objective: Keep PaO2 high, PaCO2 low
 Solubility of respiratory gases dictates the drive to breath in air-
breather’s vs water-breathers
 Air-breathers: have high PaCO2
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o Drive to breathe is to rid plasma CO2

 Water-breathers: have low PaO2
o Drive to breathe is to load plasma with O2
Phyoogenetic controo of ventioation

 Transition from buccal pump to aspiration pump evolved with air breathing
 Mammals, fish, reptiles
Fouid movement
Boyle’s Law: P1V1 = P2V2
 Fluids flow from areas of high to low pressure
 When pistol is pulled, the air expands
 If pistol is pushed, air condenses
Tidao ventioation in mammaos = ASPIRATION

 Tidal ventilation
o Tidal = what goes in also comes out the
same way
 Quiet breathing at rest:
o Inspiration = active
o Expiration = passive (elastic recoil)
 However during exercise expiration
is active as well (engaging
abdominal muscles)
 3 phase event:
o Inspiration
o Post inspiration
o Expiration
Inspiration
 Diaphragm contracts when we breathe in and depresses (gets shorter and lower into
abdominal cavity)
o Acts like a slingshot
 Muscles contract to open upper airways
Expiration
 Smaller diameter of airway, more pressure and slower rate of flow
o This is why we have small vessels in lung and upper airways which relax to slow the
gas exchange down
 Diaphragm and intercostal muscles relax and elastic recoil occurs to push out air
Tidao ventioation in reptioes = ASPIRATION

 No diaphragm
 2 phase respiratory cycle (don’t have closure of upper airways to
slow things down)
o Inspiration and expiration
 Expiration = active
 Inspiration = active
 Glottal closure at end of inspiration
e.g. crocodilians
 Their liver is attached to a membrane and that membrane is
attached to a membrane which pulls and pushes (similar to
diaphragm)
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o Diaphragmaticus
e.g. lizards
 Use ribs and intercostal muscles to breath as pump
 However, issue arises when they move as they use the same intercostal muscles for
movement
o Have adaptation (different mechanism) to assist in breathing
Natures chaooenges
- Respiratory constraints in chelonians
How do you breathe when you are stuck in a hard shell?
 Move whatever they can e.g. abdominal muscles and limbs
 Use axillary muscles around forelimb and hindlimb to breathe
as these are soft muscles
How do you breathe if you are an air-breathing turtle stuck underwater

 The Fitzroy River turtle (Rheodytes oeukops) breathes through
its cloaca, using cloaca bursae as the site of gas exchange
 Highly vascularised tissue which allows them to take part in gas exchange in cloaca
- Respiratory constraints in snakes

How do you breathe when you have a mouth full of food?
 Some snakes can throw glottis outside their mouth while they’re eating in order to try and get
oxygen
 Normally glottis is difficult to get to, at back of the mouth
Tidao ventioation in amphibians = ASPIRATION and BUCCAL force

PUMP
 Control of valves important in amphibians
 Buccal pump
o Different places than they use as valves: nostrils,
glottis (contractible)
o Squeeze lungs to push air out into atmosphere
o Mixed air in buccal cavity
 Separate synchronized movements between buccal movement and lung movement
Muotipoe ventioation strategies in fishes

1. Buccal expansion and compression
2. Buccal and opercular expansion and compression
3. Ram ventilation
4. Tidal ventilation (feeding lamprey), and air breathing fishes
TIDAL BREATHING IN FISHES

What if you are a fish but your mouth is busy feeding?
AIR BREATHING FISHES

 Facultative and obligate air breathers
o Can evolve to breath air when there is low oxygen levels
dissolved in water
 Evolved multiple times in fishes
 Modified respiratory structures
o Reinforced gills that do not collapse in air
o Mouth or pharyngeal cavity
o Vascularised stomach
o Specialised pockets of the gut
o Lungs
 Behavioural component in breathing
 Only minimal fish take part in tidal ventilation (old fish)
e.g. Arapaima – obligate air breather
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Water breathing in Eoasmobranchs = BUCCAL PUMP

1. Buccal expansion
 Pressure in buccal cavity falls
2. Mouth and spiracles close (valves)
3. Buccal contraction
 Pressure in buccal cavity increases
 Water expelled through hill slits
e.g. Shark breathing
Breathing in teoeost fishes = BUCCAL PUMPING

1. Mouth open, opercular valve closed
 Buccal and opercular expansion
2. Close mouth
 Buccal contraction
3. Opercular valve open
 Buccal and opercular
compression
 If opercular is open and mouth is closed, water has to go
out of the opercular
4. Mouth and opercular valve open
 Opercular compression
RAM BREATHING IN FISHES

 Ram ventilation
o Force water over their gills by forward propulsion – continuous flow of water
o Extremely efficient oxygen extraction
o Highly active fish, including some sharks and tuna
Note: progression from buccal pumping and aspiration pumping throughout phylogenetic tree
Respiratory strategies
1. Circulating the external medium through the body
 Thin surface – able to exchange gases easily
e.g. insects
 Constantly circulating
2. Diffusion of gases across the body surface + circulatory

system e.g. cutaneous respiration
 Small animals with thin skin
e.g. Amphibians
e.g. Neonatal marsupials
 For example, newborn dunnarts, tamar wallabies
Strategies for gas exchange

 Non-directional ventilation: diffusion/cutaneous respiration
 Diffusion of gases across the body surface + circulatory system
e.g. cutaneous respiration
 Diffusion of gas is limited (≈1mm)
 If there is a boundary layer/thick surface, the blood and medium
doesn’t get as close
 Frogs have highly vascularised skin to assist in cutaneous
blood flow
3. Diffusion of gases across a specialized

respiratory surface + circulatory transport
 Internal surfaces: lungs, gills
o Majority of fishes have internal gills
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o Gill slits for elasmobranch or eel

 External surfaces: gills
o Axolotl – wiggles gills to stir the water
GAS MOVES BY PASSIVE DIFFUSION

Fick equation: J = K (P1-P2)/X
 J = diffusion rate
 K = permeability gas constant (Krogh diffusion coefficient)
 P1, P2 = partial pressure of the two regions
 X = membrane thickness
 Diffusion occurs in the direction of partial pressure gradient
 Bigger pressure difference will give you a bigger diffusion rate
 A smaller thickness will give you a bigger diffusion rate
Gas exchange membrane

How does the thickness and area of the membrane compare to the metabolism of the species?
 Birds/mammals vs. rainbow trout
o Trout has thicker membranes than
the birds
o Trout has smaller surface area
o Trout has lower metabolic rate –
something with a lower metabolic
rate has a thicker membrane and
smaller surface area
 Yellowfin tuna vs. Dogfish
o Dogfish has thicker membrane than
tuna
o Dogfish has smaller surface area than the tuna
o Dogfish is less metabolically active than the tuna
Gas exchange organs

GILLS
 Thick cartilage with “fingers” called filaments which are
covered by lamellae (where gas exchange happens)
 Blood and water flow in opposite directions =
countercurrent flow
o The most efficient means of exchange
 Water is unidirectional
 Always a pressure difference until blood reaches lamellae
Unidirectional ventilation
 Co-current
o Blood flows in the same direction as the medium
o Slightly less efficient
o As gradient gets smaller, P1 and P2 gets smaller and
the diffusion rate is lower
Adaptable and multipurpose organs

 Mitochondrial rich cells (MRC) in gills are also used for osmoregulation
 Gills are highly variable between species
 Gold fish
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o Can tolerate a range of environmental conditions e.g. changes in temperature,

salinity, oxygen, and pH
o When you expose a gold fish to a hypoxic environment in higher temperature, they
grow long, thin lamellae to increases surface area in order to increase the diffusion
rate (and therefore gas exchange)
 Gills can adapt to changes in just hours
Adaptation in gill morphology

 Exposure to suspended sediment during larval development:
o Excessive mucous discharge
o Increase growth of protective cell layers
o Increase gill epithelium
o Increase diffusion distance
LUNGS
 Structural complexity increasing with phylogeny
 Gas exchange surface:
o Thin membrane
o Large surface area
o Highly vascularised
 Lungs of bullfrog – transparent
 Lungs of lizard – transparent
o Very thin layer
 Alveoli are specific to mammals
Alveoli
 To improve gas exchange:
o Increase surface area
o Small diffusion distance by decreasing membrane thickness
o Highly vascularised
Tidal ventilation
 Air flows in and out the same path
 Amount of oxygen in air is decreasing as it hasn’t been
replaced yet
 The difference between the partial pressures is getting
smaller so diffusion rate drops gradually
 Not as efficient as countercurrent exchange
 This is a useful mechanism is you have short airways
Natures chaooenges
- Respiratory constraints in snakes
How do you get air to your lungs when you are long and skinny?
 Long neck, therefore a lot of dead space (not gas exchange area)
 Only anterior part of lung is gas exchange surface
o Rest of (majority) of lung just moves air
o Therefore, the first section of the airway is gas exchangeable and the bottom half is
dead space
Phyoogenetic comparison of gas exchange organs

 Transition from gills to lungs happens during transition from water to air breathing
Unidirectionao ventioation
CROSSCURRENT in birds
 Blood flows cross-wise over the flow of air
 Very efficient
 Very thin membrane
Oxygen carriers
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HAEMOGLOBIN (Hb)
 Found in erythrocytes (red blood cells), muscles and neurons
 Contains 4 subunits
 Central ion = Fe
 Increases oxygen carrying capacity of blood – oxygen
dissolves poorly in water (blood)
 Only free particles of gas in liquid contributes to partial
pressure
o O2 bound to Hb only contributes to concentration
of O2
 Blood oxygen concentration (CaO2) is a combination of both bound and dissolves oxygen
Binding curves
 P50 = measure of oxygen affinity of Hb
o = PO2 where 50% of Hb is saturated
 useful for comparing Hb affinity of different species, and conditions
 Increases haematocrit or Hb levels, decreasing oxygen carrying capacity
pH and Pco2 affect O2 affinity

 Bohr effect or shift = a decrease in pH r increase in P CO2
reduces oxygen affinity; right shift
 Root effect = a Bohr effect with a reduction in the oxygen
carrying capacity
o Changes amount of oxygen blood can carry
o Common in fish
Temperature effect on oxygen affinity

 Increase in temperature decrease oxygen affinity; right shift
In exercising muscle (metabolic hyperaemia)

 Acidification of local muscle tissue (acidosis)
o Lowers affinity of Hb, allowing muscle to take up more oxygen as it
is released from Hb
 When muscle is active, heat is produced, decreasing Hb affinity and giving more oxygen to
muscles
HAEMOCYANIN
 Blue blood
 Found in:
o Arthropods (crabs, lobsters, shrimps, crayfish)
o Molluscs (including cephalopods, gastropods)
 Contain copper not iron
 Oxygenated form = blue
 Deoxygenated form = clear
Note: tropical fishes have even narrower range of aerobic scope, thus under even greater threat from
warming oceans
 Increase in water temperature decreases ability for oxygen to dissolve in water
Adaptation to Altitude
High aotitude
Barometric pressure = 1/10 decline per km
 Dalton’s law of partial pressures:
o In a gas mixture, each gas exerts its own partial pressure
that sum to the total pressure of the mixture
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Aotitude and barometric pressure

Sea level Summit of Mt Everest
 Pressure = 101 kPa  Pressure = 33 kPa
Chaooenges of high aotitude

 Decreased barometric pressure
o 1/10 decline per km
 Decreased partial pressure of oxygen
o 1/10 decline per km
 Decreased temperature
o -20ºC
 Decreased humidity
o 30% RH (relative humidity)
OXYGEN CASCADE
 There is a minimum oxygen requirement at mitochondria
CHEMORECEPTOR REFLEX
 Objective: keep PaO2 high and PaCO2 low
Response to hypoxaemia (low PaO2)

 Increase in all parameters
 Information sent to cardiovascular control centres and central respiratory rhythm and pattern
generator to increase BOTH
o Increase in drive to respiratory muscles and increased ventilation as a result (pulling
in more air at a faster rate – don’t have to both increase at same time)
o Increase activity of heart (increased HR) and vessels (vasoconstriction) resulting in
increased circulation
 Result = increased gas exchange
o Moving more blood through lungs to pick up oxygen
Chemoreception in mammals
 Peripheral chemoreceptors
o Carotid and aortic bodies
o Sensitive to changes in PaO2
 But can also respond to changing PaCO2, pHa, glucose
(metabolic sensor)
o Activation = restore PaO2
 Central (medulla) chemoreceptors
o Sensitive to changes in PaCO2
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o Activation aims to restore PaCO2 homeostasis (and also pHa)

o Important for moment-to-moment regulation of breathing for air-breathers
Acute response to hypoxia

 Increases ventilation to get more oxygen
 Immediate response is to increase respiratory rate
Acclimation to hypoxia
 Increased haematocrit or haemoglobin levels
 Increased oxygen carrying capacity
 Polycythemia
o Increase release of erythropoietin
o Increased red blood cells
o Increased haematocrit/haemoglobin
o Increased oxygen carrying capacity
 Maladaptive response – for every adaptation there is a
potential for maladaptation (bad)
o Increased blood viscosity
o Vascular resistance
o Cardiac afterload (work load)
o Pulmonary pressure
o Oedema (pulmonary and systemic)
o Local tissue perfusion
Humans at high aotitude
 1/3 of oxygen at summit
Humans at sea level vs. high altitude

 High altitude dwellers have adapted in a way
to maintain oxygen partial pressure
 What adaptive strategies could maintain
oxygen requirement at mitochondria?
Human adaptation at high altitude
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o Interference with estrogen receptors and steroidgenesis
Eggshell thinning in British and North American birds

 In the 1950s widespread reproductive failure and severe population crashes were observed in
predatory birds from both Britain and North America
o Increases in number of broken eggshells in nests
o Reduced eggshell weight/size ratios
 Changes in eggshell thickness (calcium-dependent), began to occur 1 year after rapid
increase in organochlorine pesticide (predominantly DDT) usage started in both locations
o Calcium metabolism in birds in regulated by estrogen and parathyroid hormones
o High concentrations of DDE, PCB and other contaminant residues found in eggs
 This occurred in two different parts of the world
Diesthylstillbestrol (DES) use in pregnant women in the US

 Between the 1940s and 1970s, a synthetic estrogen, disethylstilbestrol, was prescribed to
pregnant women to (supposedly) reduce the risk of miscarriage, morning sickness etc.
o Doesn’t actually do this
 DES daughters
o Increase in obscure reproductive tract cancers and adverse reproductive outcomes
o Increased risk of developing breast cancer
 DES sons
o Increase in reproductive tract abnormalities e.g. undescended testes
 Causes alterations in reproductive hormone biosynthesis in females – altered sex steroid
ratios
 Causes epigenetic alterations in mouse-models (heritable changes in gene function)
o Increases in ovarian cancers, changes in sperm production, changes in menstruation
patterns, genetic tract differentiation
 Drug has since been banned
Pesticide exposure in pregnant women in Denmark

 Two cohorts of pregnant women, classified as occupationally exposed and occupationally
unexposed to greenhouse pesticides were examined between 1996-2000
 197 pregnancies – 203 infants (113 boys; 90 girls)
o 3-fold increase in the risk of delivering a son with cryptorchidism amongst exposed
group i.e. undescended testes
o Decreased penile length and testis volume
o Lower serum concentrations of testicular hormones (testosterone and inhibin B)
o Increased serum concentrations of SHBG and FSH
o Increase in LH: testosterone ratio
 Impacts on Leydig cell and Sertoli cell function during testicular development
 Long way to go to understand mechanism – something acting on endocrine function/hormone
biosynthesis metabolism
Difference in fertility amongst Scandinavian men

 A review published in 1992 reported an overall decrease in male fertility over a 50-year period
o Reduced sperm counts
o Increased rates of testicular cancer
o Increased rates of reproductive abnormalities i.e. undescended testis
 Happening in Scandinavia but also occurring globally
 In 2002, a study was published that compared ‘healthy’ men from four Nordic countries (West:
Denmark, Norway); (East: Finland, Estonia) – didn’t present to doctors with any fertility issues
o Lower sperm counts in the West
o Lower frequencies of sperm with normal morphology in the West
 East-West gradient in semen parameters followed a parallel gradient in incidence of testicular
cancer
 Unresolved – open questions as to why is there a geographic gradient where this is
occurring?
EDCs in Victorian Estuaries: Why we need estuarine biomonitors
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 Several studies have shown that Victorian environments are polluted with chemicals that are
known to cause endocrine disruptive effects in organisms, yet very few studies have
attempted to measure effects in resident fauna
Effects of endocrine disruptors on early development stages of fish

 Embryo and larval laboratory exposure studies
 Hatch rates, deformities, growth and survival
 Grow out to observe gonad development and sex ratios
Effects of pesticides on embryonic heart rate

 Chlorpyrifos (insecticide) causes a significant reduction in heart rate
 Will have effect on normal organ development
 Increases deformities
Vitellogenin detection using Western blot

 Vitellogenin is a protein involved in yolk synthesis in egg laying
species
 It should only be produced in sexually maturing female fishes – not
males
 Laboratory exposure to a high dose of estrogen – 17-oestrodial
 Samples from field collected black bream
Gonad histopathology
 Egg production in testes of male goldfish exposed to urban pollutants
Applied Marsupial Reproduction and Development
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Lecture Notes - Animal Physiology (Melbourne)

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Lecture Notes - Animal Physiology (Melbourne)

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Animal physiology - Lecture notes All lectures

Comparative Animal Physiology (University of Melbourne)

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Comparative Animal Phisiologi

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What is Animal Phisiologi?

Comparative animao physiooogy

Adaptation, accoimatisation and accoimation

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 If a particular physiological process is shared by closely-related animal species living in vastly

Changes in physiooogy in response to changes in environment

Animaos can modify their own environments

HOW DOES IT COME ABOUT?

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o Detects high levels of stress

e.g. Sea Star

e.g. Crab-eating frog

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to changes in the conditions surrounding them constant internal environment

Extreme temperatures and thermal tolerance

How does this affect populations?

Hormones, Receptors, Pheromones

LOCAL AUTOCRINE AND PARACRINE

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Transport Extracellular fluid Synapse Circulatory system

Speed Rapid Rapid Slower

Duration of response Short Short Longer

Exocrine secretions: pheromones

Chemicao signaooing in endocrine system

Main groups of chemicao messengers

Synthesis of peptide hormones

Synthesis of steroid hormones

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Hormone action on target ceoos: neurotransmitters

Hormone action on target

Hormone action on target ceoos: steroids and thyroid hormone

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Hormone action reguoation by receptors

DIFFERENCES IN DISTRIBUTION OF VASOPRESSIN (V1a) RECEPTORS IN BRAIN

Receptors for catecholamines

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Feedback mechanisms – catecholamines

 Neuroendocrine cells in hypothalamus which lead down to

GCs increase the availability of quick energy to muscle and nervous

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Environmentao effects of stress

Long term effects of stress

SUMMARY OF RESPONSE TO STRESS BY ADRENAL GLAND

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 Ionic regulation = control of ionic compositions of body fluids

COPING WITH IONIC/OSMOTIC CHALLENGES

Osmotic proboem for terrestriao animaos

Proximal convoluted tubule (PCT)

Concentration of urine: Loop of Henle

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Reabsorbed solutes and water are taken away

Distal convoluted tubule (DCT) & Collecting duct

Water ooss – amphibians

e.g. water gain through the skin of a Bufo Bufo

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Body fouids v environment composition

Blood concentration relative to Osmoregulatory mechanism

e.g. Freshwater teleost