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 Intertidal Zone
- area that is covered & uncovered by the tides each day teeming
with life
- most challenging habitat on Earth
o Cycle of Tides
o Hot Days - temperature can double (Desiccation)
o Cold Winter Days - temp can drop almost to freezing point
o Day - high oxygen due to photosynthesizing algae
o Night - low oxygen due to respiration
o Change in H2O pH
- due to photosynthesis & respiration
- can be slightly acidic to very alkaline
o Salinity
- can increase as water evaporate and decreases on rainy day
- Common Inhabitants include the Porcelain Crabs
o Genus Petrolisthes (over 100 species)
o can live both in high intertidal & nearby subtidal zones
o strong correlation between the maximum temp. of the habit
in which a species is found & the highest temp. the which
the heart can beat
- Species found in the high intertidal in the hot tropics
= able to maintain cardiac function at substantially higher
temp than species at cooler or constant temp  shows that
physiology of cardiovascular system may play a role in
setting the geographical distribution of the Porcelain crabs
= despite high tolerance for habitat with high temp, species
found in it have high risk of fatality than species at cooler
temp  shows the responses to environmental changes
 Animal Physiology
Atom  Molecules  Cells  Tissues  Organs  Organ
- Kurt Schmidt: the study of how animals work
System  Organisms  Populations  Communities 
- the study of structure and function of various parts of an animal,
Ecosystems  Biosphere
and how these parts work together to allow animals to perform their
normal behaviors and to respond to their environments
- a central discipline in biology linking the underlying molecular &
cellular mechanisms to characteristics of whole organisms such as
performance & fitness
 Phenotype
- physiological properties of an animal
- includes all of the observable traits of an organism at all levels of
biological organization
- to produce it both the genotype of an organism and its environment
interact through development are needed
- influences the animal’s reproductive success
 Genotype
- the genes that determines a large part of an animal’s characteristics
- an individual genotype have the capacity to produce more than one
phenotype
- controls the way animals can alter their phenotype in response to
physiological & environmental conditions
 Identical twins raised in diff places, one may be taller due to
difference in external conditions
 Physiological Process
- product of the activities of complex tissues, organs & systems that
can arise through complex patterns of genetic regulation of countless
cells
 Evolutionary Change - ultimate cause of the enormous
diversity of animal species
UNIFYING THEMES IN ANIMAL PHYSIOLOGY
A. Integration in Physiology
- biologists often organize the living world by dividing it into what
are termed levels of biological organization
- processes at each level of organization interact to produce the
processes at the next level of organization
- Reductionism: assumes that we can learn about a system by
studying the function of its parts

- Emergence: feature of complex systems shows that the whole is
often more than the sum of its parts
- Emergent Properties: properties that can be observed at one level
of biological organization & that are due to the interactions of the
component parts of the system
- Basic Research: provides profound insights into how animals work
& the evolutionary cause & consequences of variation in
physiological processes
- Physiology can be applied in conservation biology, ecology,
agriculture, human health, disease & veterinary medicine
 RA 8485 - “Animal Welfare Act of 1998”
- laboratory animals are protected under this act, registered and
prescribed proper care, handling & testing of animals are also
included
- Animal Welfare: humane treatment of animals with respect to their
comfort & well-being
- Animal Rights: idea that animals have intrinsic & unassailable
rights
- Model Organisms:
= species that are chosen because they have features that make them
particularly suitable for specific experiments
= used because (1) they have features that are conductive to
experimentation & (2) understanding a process in the model provides
insight into how the process works in other species of interest
- August Krogh Principle: “For every biological problem there is an
organism on which it can be most conveniently studied.”
o Squid: have some specialized neurons that are large enough to
be easily seen & readily manipulated; use in understanding of
how neurons work in all animals
o D. melanogaster: giant salivary glands for locating polytene
chromosomes
o African Clawed Frog: oocytes have numerous nuclei for
transciption
o Sea Raven: possess a large heart
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B. Physics & Chemistry: The Basics of Physiology
- Animals are constructed from natural materials & thus obey the
same physical and chemical laws that apply to everything
- Physiology is a quantitative science
 Adolf Fick’s Law of diffusion
st
= 1 Law: substances diffuse from areas of high concentration
(potential energy) to areas of low concentration (potential energy)
~ movement is a consequence of the 2nd law of thermodynamic =
isolated systems spontaneously move toward a state of maximum
entropy
 over time, differences in concentration, charge, temperature,
or pressure will tend to equalize within the system, unless energy is
added to maintain this difference
o Concentration Gradient
- source of potential energy use to drive diffusion
o Voltage Gradient
- source of electrical potential gradient which can drive movement of
charged molecules
- used in (1) signaling in neurons & (2) active transport of material
into cells, involve the movement of charged molecules like sodium
across membrane
- together with concentration gradient, both are used to determine the
extent and rate of diffusion
o Conduction of Heat
- use the same principle that apply to diffusion od substances
- heat flows form areas of high temperature to areas of low
temperature
- form and function are shaped by the need to regulate heat loss or
gain
o Pressure Gradient
- source of potential energy from areas of high pressure to areas of
low pressure
- used in understanding the function of (1) circulatory and (2)
respiratory systems

= 2nd Law: amount of substance that diffuses across a surface is
proportional to the area of that surface & inversely proportional to
the distances across which the substance must diffuse
~ critical for understanding the form & function of epithelia such
as the lungs and the gut that are involved in the exchange of
substances by diffusion (must have as large surface are as possible
and be as thin as possible to maximize material exchange)
~ this explains why gas exchange surfaces such as lungs & gills
are extremely thin and why animals that are larger than a few
millimeters in diameter must have circulatory systems to move
substances around their bodies
o Brownian Motion
- random movement of particles in a solution, that the time needed
for a particle to diffuse across a given distance is proportional to the
square of the distances
 Molecular Properties: determine the physiochemical characteristics
of biomaterials
1. Collagen
- high level of this structural protein helps the aorta withstand the
high pressure generated by the heart
- low level of it allows small blood vessels like capillaries to be
thin to maximize the material exchange by diffusion
2. Keratin
- network of proteins that can be made more rigid by the addition
of bonds that cross-link multiple keratin proteins together
- heavily cross-linked keratins helps fingernails to be stronger and
less likely to bend
- fewer cross-linked keratins allows hair to be more flexible
 ELECTRICAL LAWS: Membrane Potential
- potential difference that drives processes needed for survival such
as movement of essential molecules across membranes
- Electrical Potential: send signals within & between cells, helping
to coordinate the complex processes of the body like how (1) neuron
and (2) muscle cells work
 CHEMICAL LAWS: Temperature
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- rate of most chemical reactions increases as temperature increases
 increasing temperature increases the energy of molecules & causes
an increases in the number of collisions between molecules in a
closed system
- high temperature that stabilize proteins begin to break down down
& protein function declines
- Catalysts: when these break down, the rate of reaction falls
 PHYSICAL LAWS: Scaling Relationship
- the relationship between anatomical or physiological traits and
body size
1. Isometric: morphology or physiology change in direct proportion
to body mass
2. Allometric: body shape or physiology changes disproportionately
as body size increases
- Galileo Galilei (1638), disapproves the isometric concept through
his description on how the bones of larger animals are proportionately
thicker than the bones of small animals  relationship between area
& volume
 volume of a cube is equal to the length of the side of the cube to
the third power (L3)
 surface area of each side of the cube is equal to the length of the
side squared (L2)
 cubes have six sides (6L2)
 surface area increases by a power of 2 while volume by a power of
3
 the ratio of surface area to volume decreases as length of the
side increases
 surface: exchange of materials with the environment
 volume: processing & use of exchanged materials
 volume increases more than does surface area as radius
increases, the surface available for exchange of materials quickly
becomes limiting
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y = aMb where y = surface area show that SA-to-Volume ratios cannot explain metabolic scaling
M = body mass (or volume) relationships
a = constant - suggested the value of scaling coefficient was closer to 0.75 (3/4) rather
b = scaling coefficient than the value of 0.67 (2/3)

Scalling Coefficient (b) C. Form, Function & Evolution

- 2/3: for cube or a sphere  surface area increases as the square of Coyote (Canis latrans) Koala (Phascolarctos cinereus)
the radius, while volume increases as the cube of the radius same mammals & similar size
1. b = 0 omnivores obligate herbivores
- the process is unaffected by body size strong preference for meat strong preference for
2. b = 1 & Eucalyptus leaves - toxic,
- the physiological variable increases proportionately with the body can’t eat tough or fibrous extremely fibrous, low
mass  process scales isometrically (e.g. mass of the heart) leaves like leaves and grass nutritional quality
 Negative (b) longer intestinal tract - fibrous
shorter intestinal tract diet takes much longer to digest
~ the variable decreases with body size (e.g. mammal’s heart rate)
 Positive (b) larger cecum - contains
smaller cecum
cellulose-digesting bacteria
~ the variable increases with body size (e.g. fiddler crab’s claws)

 Metabolic Rate: A Positive Scaling Coefficient
 Max Rubner
- reported that metabolic rate of dogs of various sizes was constant when
body surface area was taken into account
- suggested that metabolism has scaling coefficient of 0.67
 metabolic rate of one gram of tissue from a large dog is lower than
the metabolic rate of one gram of tissue from a small dog
- he developed a hypothesis about Heat Dissipation to explain his
observation on dogs’ metabolic rate
- Endotherms: mammals like dogs maintain relatively constant body
temperature by generating metabolic rate
- If metabolism scaled isometrically with body size, he suggested that large
dogs would generate too much heat to dissipate across the relatively small
surface area, and thus the must have lower metabolic rate
 Max Kleiber
- disapproved Rubner and assembled a much larger date set relating body
size and metabolic rate in a variety of species of birds and mammals to
o Proximate Cause: immediate physiological/biochemical basis of traits
o Ultimate Cause: evolution as an advantage to the ancestors
 Species that are closely related to each other are likely to share more
features and species that are distantly related are likely to share fewer
features
 Adaptation
- can be described as:
(1) synonym for “remodeling”
(2) evolutionary adaptation
- product or process of evolution by natural selection
- change in a population or group of organism over evolutionary time
- trait that arose via process such as NS that conveys an increase in
reproductive success
- Evolution of Insecticide Resistance
= excellent example of the principle of adaptive evolution
= Organophosphates

~ kills insects inhibiting acetylcholinesterase, an enzyme vital for
neuronal transmission
= over time insecticides with organophosphates showed a decreased on
its affectivity resulting to insects with beneficial mutations to survive &
reproduce like mosquito (Culex pipiens)
~ have mutations in the acetylcholinesterase genes
~ extra copies of the esterase gene - which encodes an enzyme that
converts the organophosphates into a less toxic form
= however, this mutation is only vital in the presence of insecticides
~ overproduction of esterase protein use energy that serve other
functions
- for adaptive evolution to occur:
1. the must be variation among individuals in the trait under
consideration
2. the trait must be heritable - genetically determined & passed on to
offspring
3. the trait must increase fitness (reproductive success) if the
individual that have the trait
4. the relative fitness of the different genotypes depend on the
environment  if the envi changes, the trait may not be beneficial
 Non-adaptive Evolution: Genetic Drift
- random changes in the frequency of particular genotypes in a
population over time
- result in substantial differences in the phenotypes of 2 populations,
independent of any adaptive evolution
- mostly likely to occur in small populations & result to happenstance,
not of differences in fitness
- can act in opposition with selection
- includes the following concepts:
a. Founder Effect
= the derived populations may differ from the ancestral
populations, but not for any reason related to natural selection or  Deep Homology
fitness
b. Neutral Adaption
= changes in populations that are not due to differences in fitness
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Llamas & Vicunas Dromedary (1-humped camel)
member of the family Camelidae
both have hemoglobin with very high oxygen affinity 
preadaptation or exaptation and not adaptation
high altitude habitants lowland habitants
 Phenotypic traits can be similar among organisms due to:
1. Homology
- inheritance from a shared ancestor
- e.g. limbs of vertebrates  different shapes but have similar
bones due to an ancestral similarity
2. Analogy
- share a similar function but do not share a common
evolutionary history
- e.g. wings of birds & bats  for flight but one is a mammal
the other not
- falls within the broader concept of Homoplasy
= independent evolution of similar traits
= due to either:
a. Convergent Evolution
~ unrelated taxa toward similar phenotypes can occur due to
particular shapes or functions are favored in a specific environment
~ e.g. wings of birds & bats; eyes of cephalopods & vertebrates
b. Parallel Evolution
~ shared underlying trait evolves in the same way independently
~ e.g. threespine stickleback fish (Gasterosteus aculeatus)
- Freshwater species
= lost most of the armor & have smaller spines
- Marine species
= heavily armored & larger spines
= perform better in terms of ability to swim aerobically for long
periods  larger swimming muscles & it contains more aerobic fibers
- arises from shared molecular traits among all animals & may
underlie many examples of convergent evolution like:

1. Eye Development
- uses the Pax6 transcription factor
= controls eye development for all animals
= gene associated with photoreceptors
= homologous & shared among all organisms but the camera type
eyes of vertebarates and cephalopods are independently derived from
the non-camera type eyes of their ancestors & are thus homoplastic
2. Heart Development
= In Drosophila, Tinman controls this development
= a gene similar to Tinman is also used by Cnidarians to expressed
around the base of the gastrovascular cavity, a region involved in
pumping fluids throughout the body
= In Mice, Nkx is needed for this development
= both genes have very similar proteins & share the same
evolutionary origin
= Similarly, these genes have been discovered in Lancelet, an
invertebrate chordate expressed in the developing tubelike heart
D. Regulation & Homeostasis
 Confomers
- allow internal conditions to change when faced with variation in
external conditions
- e.g. body temperature of fish  low in cold & high in warm water
- less expensive in terms of demand of metabolic energy for processes
 Regulators
- maintain relatively constant internal conditions regardless of the
conditions in the external environment
- e.g. human body temperature  likely to be approximately 37°C
- provides a more stable internal environment due to environmental
changes’ deleterious effects
 Some may be regulators with respect to one internal parameter, but
conformers with respect to another parameter
 Lizards conform to external temperature but regulate their
internal salt concentrations within a narrow range
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 Homeostasis
- coined by Walter Cannon
- importance was explained by Claude Bernard
- maintenance of internal conditions in the face of environmental
perturbations
- animal initiates specific responses to control and or regulate a
particular vital variable
- e.g. Human Body Temperature
= Shivering to produce heat when you stand in cold air
= Sweating to release heat
- Physiological responses can solve either:
Situation  dogs response to temperature
a. Short Term Challenges: when hot, move to a cooler location or
pant to shed heat in its breath
b. Long Term Challenges: in seasonal cycles, grow fur in autumn &
shed it in spring
 Allostasis
- the process of achieving homeostasis through change
- e.g. (1) Norther Mammals shed their brown summer fur & grow a
thicker white winter fur (2) Female Mammal Lactation
 Principles that govern physiological changes:
I. Some physiological strategies are effective in the short term but less
useful for the long term.
 Holding your breath: effective in swimming but not on high altitude
location with low levels of oxygen
II. Some strategies require a significant investment in resources and need
long to take effect.
 Hair Growth: relatively slow process that require metabolic energy
III. Some stressor are sufficiently predictable that animals remodel
physiology in anticipation of the stress & often in predictable cycles.
 Daily: Circadian rhythm
 Seasonal: Growth & Shedding of Fur
 Lunar Cycle: Coral Reef Spawning
 Specific Environmental Cues: Temperature or Photoperiod

 Feedback Loops
- used by the Endocrine & Nervous System
- to maintain homeostasis, animals must:
(1) detect external conditions, (2) if necessary initiate compensatory
responses and (3) keep vital areas buffered against unfavorable change
- most often animals use a Reflex Control Pathway  a change in the
internal & external environment provides a stimulus which causes a
response
- Antagonistic Controls
= independent regulators that exert opposite effects on a step or
pathway which fine-tune physiological responses
= e.g. Regulating Body Temperature through Heat Production &
Dissipation
- Set Point
= preferred physiological state defended through feedback loops
= e.g. 37°C Body Temperature  may vary between individuals and
changes throughout the day
 Negative Feedback Loop
- response send a signal back to the stimulus, reducing the intensity
of the stimulus
- e.g. Early Digestion
eat  stomach swelling  change in stomach volume and early
digestion trigger NFL  sends signals to brain  reduce appetite
 Positive Feedback Loop
- unlike NFL which minimizes changes in the regulated variable,
PFL maximizes changes
- e.g. Vomiting
= When toxin is detected in Stomach, a PFL is triggered to
induce forceful contractions that propel the food back up the
esophagus to induce vomiting
 Phenotypic Plasticity: ability of an organism to alter its phenotype in
response to environmental conditions and its phenotypic changes
could be:
o REVERSIBLE
 Acclimation - process of change in response to a controlled
environmental variable (usually in laboratory setting)
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- e.g. take a fish from water at 15°C and transfer it to water at 5°C
 Acclimatization - process of change in response to natural
environmental variation
- may be the result of temperature change, changes in day length,
food availability, any other environmental parameters that vary
between summer and winter
- e.g. take a fish in the summer (15°C) vs. a fish in winter (5°C)
o IRREVERSIBLE
 Reaction Norm
- the change when the phenotypes are part of a continuum
 Polyphenism
- the change when a particular phenotype exists in 2 or more
discrete forms
- the most common type of polyphenism is due to developmental
plasticity  development under different conditions results in
alternative phenotypes in the adult organism that can’t be reversed
by subsequent changes in the environment
- e.g. Water Fleas (Daphnia pulex)
= Presence of Predators: develop large armored, helmet shaped
heads & an elongated spiny tail
= Absence of Predators: develop small heads & a shorter and less
spiky tail
 retain these morphologies even if the predator extracts are
subseqently removed from or added to the water
 Epigenetic Inheritance
- some acquire traits can even be passed on to subsequent
generations through this mechanism
- e.g. Killifish (Fundulus heteroclitus)
= when offsprings of killifish (which live in highly polluted sites
& have high tolerance of pollutants such as PCBs) are reared in the
lab, it also have high tolerance.
= however, an F2 generation reared in lab, showed low tolerance
 suggest that the parent killfish is the result of E.I. that can
be inherited (F1) across one generation, but are then reset (F2)