The Animal Circulatory, Excretory and Respiratory

Systems

Circulatory System
o Bulk flow of nutrients, gas and wastes is the only
o An internal transport system that provides a mass method by which the entire body of larger more
flow of fluid (called convection) and frees complex organisms is accessed.
metazoans from the body size and shape o System wherein a network of vessels transport
limitations imposed by simple diffusion nutrients, gas and waste emanate from a pump.
o Multicellular animals’ mechanism for transporting
nutrients and removing wastes throughout their
body
o Simple animals (consists of a single cell layer) do
not need a CS because diffusion allows adequate
exchange of dissolved gasses, nutrient, and
wastes
e.g. Sponges and Jellyfish
o Diffusion in simple animals
o Complex system of pump and vessels: a network
of cylindrical vessels emanating from a pump
o Trend: to create an efficient and energy efficient
internal transport of nutrients, gasses, and waste
o As animals become more complex and
multicellular, simple diffusion becomes insufficient
to supply all the cells with nutrients • →
• →
→ CS in primitive metazoans:

→ Types of CS:

1. Open CS
o Pumps blood into a cavity called hemocoel
o Blood vessel pumps blood through
o Diffusion-limited metazoans are tiny, below 1 mm peristalsis to large sinuses where organs are
diameter bathed in blood
o Exists in invertebrates (e.g. insects, mollusks,
some crustaceans)
o Only works in small organisms where cells
are close to environment

2. Closed CS
o circulates blood unidirectionally from the
heart, around the body and back to the
heart
o Blood pumped through a closed system of
arteries, veins, and capillaries
o Found in some invertebrates and higher
order organisms
o Blood pumped directly into organs 1. Circulating Body Fluid
Aquatic Terrestrial
Advantages:
1. Allows for regulation of blood flow
2. Increased pressure and speed of blood
flow
3. Allows for larger organisms with cells
farther from the environment

A review of the evolutionary trends in hemostatic

mechanisms indicates the victory of complexity over
simplicity (Srimal 1996)

Open CS

o Cnidarians and Echinoderm

• Cell agglutination followed by the
formation of a syncytial network and the
shrinking and contraction of a plugof
aggregated cells has been observed.
o Mollusks
• Cell aggregation and contaction of
musculature with some increase in viscosity
of the hemolymph
 o Annelids
• Conversion of fluid into thick gum
mucilages upon contact with air; plasma
clotting
o Arthropods
• Cellular aggregation and hemolymph
coagulation involving enzymatic cleavage;
Antibacterial peptides
• Limulus (horse shoe crab)- clotting
cascade is activated by gram
negative bacteria
the placenta all introduce more challenges for
maintaining the balance between liquidity and
gelation. Having more components in the
amplification system may also make it possible to
deliver a maximum response with a smaller input
stimulus, and this in turn would necessitate having
a faster turn-off or lytic response. (Doolittle, 2009)

Close CS

o Vertebrate blood clotting evolved in parallel with

the development of a pressurized and closed
circulatory system with its all-important red cell
component
• exposure of tissue molecules triggers a
cascade of response wherein a large
number of fibrinogen molecules are 2. Increasing complexity of Cardiovascular system
converted into a gelatinous clot (Doolittle
2009) → 2 chambered heart

WHY DID THE CLOTTING SYSTEM BECOME MORE COMPLEX o In fish

(in mammals)? o 1 atria and 1 ventricle
o Blood pumped to gills to pick up O2 and
o The quality and character of the fibrin clots then travels to organs and then back to
generated in fish and mammals do not appear to heart
be significantly different. The answers must have to o Low blood pressure
do with regulating the response under a wider o Single circulation
spectrum of environments. Higher blood pressure,
more complicated cardiovascular systems, higher
metabolic rates, and new organs such as lungs or
→ 3 chambered heart o Oxygenated and deoxygenated are kept
separate
o In reptiles and amphibians o Allows for increased blood pressure,
o 2 atria and 1 ventricle thermal regulation, and sustained muscle
o 2 atria: one for deoxygenated blood from movement
body, one for oxygenated blood from lungs
o Single ventricle pumps blood out to both
lungs and body
o Oxygenated and deoxygenated blood
mixed in ventricle causing diluted O2 blood
o Amphibians are unique: they have a third
circuit that bring deoxygenated blood to
the skin for gas exchange to occur
(pulmocutaneous circulation)

→ 4 chambered heart

o Birds and mammals

o 2 atria and 2 ventricles
Excretory System

o Maintains body homeostasis by eliminating excess

substances that enter the body
o Eliminates excess water, salts, and byproducts of
cellular metabolism
o Nephridia – ciliated tubules that pump water out
of the body. The outflow is also used to discard
metabolic wastes, excess ions, spent hormones
and perhaps toxins ingested with food (i.e. urine)

Basic process that takes place in the renal tubules

o (ultrafiltration):

1. Filtration
2. Secretion
3. Reabsorption
4. Osmoconcentration
o Protonephridial excretory system usually occur in o Metanephridial excretory system usually occur in
bilaterians that lack a hemal system, a coelomic large bilaterians with both coelomic and hemal
system, or both. compartments
o Ultrafiltration is driven by fluid pressure- requires a
circulatory system
 → The Mammalian Kidneys

→ Stages of Development of Vertebrate Kidneys o Glomerular Filtration

Glomerulus- specialized fenestrated capillary; allows

1. Pronephros
o Arise from anterior part of nephric ridge
o Its posterior pronephric duct empties into the
cloaca
o First type to arise during embryonic development
o Found in adult fishes (hagfishes)
2. Mesonephros
o Arise in the middle of nephric ridge
o Often embryonic and transient
solute, water, and ions to pass through while retaining
blood cells and plasma proteins
Factors that influence the filtration of solutes and large
molecules:
• Size (molecular weight)
• Electric charge
• Glomerular capillary hydrostatic pressure

3. Metanephros
o Sprouts from the mesonephric duct, the ureteric
diverticulum grows at the posterior section of the
nephric ridge
o In males, mesonephric duct becomes the vas
deferens. In females, it degenerates.
o Found in amniotes

4. Opisthonephros
o Tubules from middle and posterior nephric ridge
form an extended kidney
o Develop into functional adult kidney in fishes and
amphibians
Respiratory System

o Gas will move from an

area where its partial
pressure is higher to an
area where its partial
pressure is lower, and the
greater the difference in
pressure, the more rapid
the gases will move.

→ Common Features of Gas Exchangers:

• Thing blood-gas barrier → Oxygen Concentration and Major Evolutionary
• Large interface Events
• Ventilatory regulation
• Low energy cost of ventilation

→ Systems of Gas Exchange: From Direct Diffusion to

Complex Respiratory Systems

o Animals independently evolved diverse respiratory

systems for acquiring oxygen
o Mechanisms, processes, and structures used for
respiration depend on the:
1. Habitat
2. Size
3. Complexity
o Habitat expansion compels the use of different
gas exchangers.
→ Oxygen Cascade: Elaborate gate keeping
mechanism to balance cellular oxygen delivery
against oxidative damage
o Supports the anaerobic origin of eukaryotes
and their preference for hypoxia
o Suggests that the ancestral bacteria that
became the mitochondria were likely
facultative anaerobes
→ Respiration in Aquatic Animals: Direct Diffusion Across
Surface Membranes
→ Respiratory Surfaces of Animals
1. Mantle (mollusks)
o air flows into the mantle cavity where
the gills (ctenidia) or lungs are present
2. Parapodia (annelids)
o are fleshy appendages used for gas
exchange and movement
3. Tracheae (arthropods)
o highly-branched chitin lined tube that
opens externally through a spiracle and
further divides into smaller tracheoles
that terminate very close to body cells
4. Book lung (arthropods)
o Folded invaginations in dorsal body wall
that forms series of lamellae where air
circulates; opens externally with a
spiracle
5. Dermal branchiae (echinoderm)
o Or skin gills are soft delicate projections
of the coelomic cavity
6. Parapodia or tube feet
o Project on the ventral side of the
echinoderm
7. Skin
8. Gills
9. Lungs
→ Invertebrate Respiration: Respiratory Organs
o Molluscs and Arthropods exhibit the greatest
variety and highest performance respiratory
systems
o These are the only taxa that evolved
dedicated structures for aerial
respiration
o Made possible powered flight in insects
o Invertebrates conquered the land and
evolved the capacity for aerial
respiration long before vertebrates did
o Parapodia in ragworm (Nereis spp.)
o Arthropoda:
- Size of the lungs and type of
embedding into the body is reduced
from dry to humid environments
o In some decapods, the gills can be used
for air breathing
1. Gill surface reduced
2. Stiffer with spacers between
lamellae to prevent collapse in
air
o Evolution of Respiratory Organs
→ Respiration in Aquatic, Semi-aquatic Animals:
Cutaneous Respiration
o Highly permeable skin of amphibians:
major site of gas exchange in terrestrial,
semi-aquatic, and aquatic species
o Gas exchange occurs by diffusion
facilitated by a thin layer of keratin and
a rich supply of capillaries in the skin
o Animals are limited to cool, oxygenated
habitats and to non-vigorous activity
o Water is dense thus ventilation is not
possible
o Oxygen is also limited in the aquatic
environment
o Requires a respiratory organ with large
surface area and extremely efficient
gas exchange
→ Respiration in Aquatic, Semi-aquatic Animals:
Vertebrate Gills
o Gills are:
a. Filamentous
b. Highly folded
c. Large surface area for gas exchange
 o Gas exchange is maximized by opposing flows of
water and blood

o The Rate of Oxygen Uptake depends on:

• Surface area of lamella
• Thickness of the epithelia across which
the oxygen moves → Gas Exchange between Air and Blood
• Oxygen tension gradients across
membranes (maintained by gill
ventilation)
o Evolutionary options to increase gill surface area:
• Increasing the number of lamellae
by spacing them closely together
• Increasing length of lamellae

→ Respiration in Basal Amniotes: The Lungs

o Primary organ for gas exchange
o Anatomically diverse
o Synthesize surfactant to prevent collapse
o Internal septation of the coelomic cavity
influenced ventilatory performance
o Helped amniotes achieve terrestriality
FOOD PROCUREMENT AND ASSIMILATION
The General Nature of Aquatic Life
Animals in the aquatic environment exist in a
1. Dense, circulating, interconnected water medium
2. Excellent solvent
3. Low dissolved oxygen
Water medium makes/allows
1. Filter feeding possible
2. Organisms with limited ability to digest organic
matter
3. Animals to increase in size as it can support the
immense height and weight
Suspension feeding group is present and common, these
animals feed on bacteria and other organic matter
suspended on water by filtering them.
• Sponge
→ Spicules, pinacocyte, and spongin fiber aids in
support and covering
→ Mesohyl is the gelatinous matrix under the
pinacoderm
→ Amoebocytes aids in food transportation to
other cells from the choanocytes which
phagocytose food particles form the filtered
water
• Cnidarians
→ Mesoglea, epitheliomuscular cells, nerve net
aids in covering and movement
→ Allows for the sessile radial animals to procure
food from all directions using their tentacles
and cnidocytes
→ They use epitheliomuscular tissues for
movement, cnidocytes and tentacles allows
food capture for sessile cnidarians.
→ Two distinct cnidarian body forms:
o POLYP
- tubular shaped body
- usually sessile, with the bottom attached
to a solid surface and the mouth opening
at the top - attachment region at the base
of the animal is called the basal plate.
o MEDUSA
- more of an umbrella or bell-shape, with
the mouth facing down
- body is often called the bell
- usually free-swimming and either propel • Echinoderms
themselves using muscle contractions or float → Uses pinnules or feathery arms to catch prey
along water currents like plankton and tube feet for movement
→ Has sensory papillae that stimulates mucus
→ Both polyp and medusa form consist of a strands which then traps organic matter
digestive sac, the coelenteron, surrounded by → Complex feeding structures; tube feet with
two layers of tissue, the endoderm and the sensory papillae and mucus to trap food
ectoderm.
o Coelenteron
- Cnidarian digestive cavity
- Has a single mouth opening through
which food or prey enters and
waste is expelled
- Considered a gastrovascular cavity
because it is where both digestion
and gas exchange between the
organism’s cells and water in the
cavity take place
- Can either be one large chamber,
several smaller chambers, or a
branched network of canals • Lophophorate
- Mouth opening is usually surrounded → Has ciliated feeding tentacle called
by tentacles lophophore
→ Tentacles are hollow and are extensions of the
o Gastrodermis mesocoel: ciliated
- Endodermal lining of the → The lateral cilia draw water bearing food
coelenteron particles into the tentacular ring, frontal cilia
- Absorbs nutrient derived from the convey it down in each tentacle and it goes
digestive process out to the base where food is trapped.
→ Mouth is located inside the lophophore ring of
tentacles

• Megalodicopia
→ Tunicates have tunic which are leathery body
 covering and hydrostatic pressure is the one
that aids in the movement of the organism
→ Has vanadium and niobium ions that are toxic
and aids in protection against predators
→ Retained gill slits which function as filter feeding
structure
→ Draw in food through the incurrent siphon then
out to excurrent siphon
֎ MOLLUSCA
• Radula - helps to graze upon microscopic
filamentous algae from a surface and feed
directly on plants. Like a chitinous ribbon and is
used for scrapping and cutting food before it
enters the esophagus
• Nephridium - invertebrate organ which occurs in
pairs and performs a function similar to the
vertebrate kidney. Remove metabolic wastes from
an animal’s body
• Siphuncle – used primarily in emptying water from
new chambers as the shell grows
֎ VERTEBRATES
• Diversification of TEETH and JAW
→ Relative jaw length = out lever length to SL
o Low level ratio- fast to close, low force
transmission, wide gape; a range of prey
items can be accessed (predatory)
o High level ratio- velocity constrained, high
force of transmission, small relative jaw
length, small frequent bites (herbivory)
→ Fish herbivory is a relatively recent event
→ In the Cenozoic, we see for the first time, fishes
characterized by small jaws
→ From Eocene to present, more stable trophic
structure
• Jawless Fish
→ First evolved
→ Include lampreys and hagfish
→ NO fins, stomachs, scales, and jaws
→ Includes a notochord, paired gill pouches, a
pineal eye, and a two-chambered heart
֎ The Vertebrate Appendicular Structure
→ Serially iterated structures (fins in fish and limbs
in tetrapods)
→ Trend towards their morphological and functional
diversification both within and between taxa.
1. For locomotion
2. Stability
3. Food procurement
→ Facilitate the feeding and locomotion in Ordovician
seas
→ Undulatory motion – limited movement like bivalves
Paired appendages- acquire faster moving prey

• Zeugopods
→ Evolved distal and radial bones
→ Digits
o For terrestrial tetrapods
o Most important component: thumb
appendages for food procurement
• Jaws
→ jaw musculature became more developed and
more force after prey is seized
→ not for chewing
o To restrain prey
o Catch prey
o Make it easier for predator to conserve
energy

• Diapsid Reptiles
→ have two pairs of temporal opening
o Evolutionary feature of tetrapods
o Allows for attachment of larger, stronger
jaw muscles
o Enables the jaw to open more widely for
stronger bite force
→ Temporal fenestrae
o Holes where strong musculature is
attached.
DIGESTIVE SYSTEM
o Digestion in primitive animals must have
been intracellular as it remains in the
Protozoa and Porifera.
o 2 groups:
1. Primitive in structure
-→ e.g. Coelenterata, Ctenophora,
most Turbellaria, and Limulus
2. More highly evolved but have retained
intracellular digestion in correlation with
their mode of feeding
→ Feed on finely divided food
(collected by ciliary mechanisms or
scraped by a radula) or on fluid or
semi-fluid food which is sucked in.
→ e.g. Brachiopoda, Rotifera,
Tardigrada, Pyncogonida,
Arachnida (other than Limulus), and
majority of Mollusca excluding
Cephalopoda
o Extracellular digestion
→ originally developed with the increased
size of available food as an aid to
intracellular digestion
→ replaced the primitive form of digestion
in certain rhabdocoel Turbellaria,
Polyzoa, Annelida, Myriapoda,
Crustacea, Insecta, Cephalopoda and
Chordata
→ results in the reduction of the ingestive
region of the gut and enables digestion,
and the removal of indigestible material,
to be hastened
→ the resultant increase in the rate of
metabolism has had profound effects on
the evolution of the Metazoa
→ appearance of extracellular digestion
has been accompanied by changes in
the structure and physiology of the gut
-→ Distinct regions have been specialized for
(1) the reception of food
(2) its conduction and storage
(3) digestion and internal trituration
(4) absorption, and
(5) conduction and formation of feces
o There is definite correlation between the
food and the nature and relative strengths
of the animal’s digestive enzymes.
o Certain animals have acquired specific
enzymes to allow them to exploit additional
sources of food, most important of which
being cellulase and chitinase.
o Adaptations acquired to increase food
diversity:
1. Periodicity of secretion in the
digestive glands
2. Evolved mechanism for the
continuous liberation of small
quantities of enzyme
3. Control of pH gut. In ciliary feeding
animals, this may be of importance
 not only in securing the optimum
conditions for the action of
extracellular enzymes but also by its
influence on the viscosity of the
mucus with which the food is
entangled.
• Sponges
→ Digestion of larger cells (>50 micrometers) are
phagocytosed by cells in the exopinacoderm.
→ Food is transferred from Choanocytes to
Archaeocytes.
o Archaeocytes- site of food digestion
→ Family Cladorhizidae (carnivorous sponge)
uses sticky cellular threads that captures small
fish and crustaceans which are digested
slowly by archaeocytes
• Cnidaria
→ mouth leasing to a blind sac called the
coelenteron
→ In Hydra, mouth opens when injured and
captured prey releases glutathione. Food is
then carried to gastrodermis which has gland
cells and releases proteases.
→ Digestion starts when nematocysts hit the prey
and Cnidae release protolytic enzymes.
→ Nutrients are absorbed by epitheliomuscular
cells, germ cells, and gastrodermal cells.
→ Large materials are phagocytosed and
digested intracellularly; process breaks down
lipids and carbohydrates as well as
completing the digestion of proteins.
→ Some cnidarians have zooxanthellae, a
symbiotic algae.
• Ctenophores
→ Have muscular pharynx that churns food.
→ With enzyme release, the processes reduces
the prey to a slurry that stream rapidly to
stomach and then to canals of the
coelenteron.
→ Wastes are eliminated through mouth and
some in anal pore
→ Large undigested food particles are removed
immediate through the mouth, not passing
the stomach.
• Flat worms
→ evolved a ciliated buccal cavity that creates
currents that draw bacteria and protozoans.
→ Intracellular digestion occurs in vacuoles.
→ During the process of digestion, pH is lowered
(approx.. 4.5 to 5) when digesting food.
→ Have a branched gut that increases surface
area for digestion and absorption .
→ Has a protrusible pharynx (simple to complex)
→ The bulbous pharynx (muscular sucking bulb)
used by free living flatworms to suck food
predisposed these animals to parasitism.
→ The pharynx is used to penetrate prey and
digest the internal content of bodies of
arthropods using endopeptidase. It also
functions to swallow prey whole.
 → Hydrolysis of ingested food is initiated by
pharyngeal enzymes. Gland cells in the gut
secrete endopeptidase; vesicles become
alkaline which marks the appearance of
exopeptidases, lipases and carbohydrases
necessary to complete digestion.
• Mollusks
→ microphagous browsers that scrape
microalgae, detritus and other organisms on
hard substrate; some are suspension feeders
→ Scraping mollusks have the radula, a ribbon of
tiny chitinous teeth that were lost in filter
feeding bivalves
→ Gut of mollusks is adapted for separating and
processing a mixture of fine organic particles
and fine mineral particles
o Foregut – mouth, buccal cavity, pharynx
o Midgut – esophagus, stomach, intestine
o Hindgut – rectum and anus
→ Absorption in mollusks occurs in midgut,
particularly in digestive caeca
→ Presence of a gastric shield, which bears the
sorting shield used to separate mineral
particles from nutritious organic particles,
increases digestive efficiency
• Arthropoda
→ Have highly specialized alimentary canal.
o Foregut - for ingestion, storage, and
initial processing of food prior to
chemical digestion
o Midgut – for enzyme secretion,
hydrolysis, and absorption of the
product of hydrolysis; with digestive
caeca to increase the area for
absorption and intracellular digestion
o Hindgut – for feces formation and
storage and reclamation of valuable
materials.
→ A valve separates foregut from midgut.
→ Hexapod gut (similar to mammalian colon)
is capable of transferring water from its
lumen back to the blood—osmoregulatory
adaptation of the gut contributed to the
success of insects in the terrestrial habitats
→ Some arthropods also have a proventriculus
a. Crop – used for storage
b. Muscular gizzard – grind food and
where indigestible particles are
regurgitated
→ For arachnids, only liquid food enters the
mouth. Digestion begins externally. The
digestive enzyme moves from the midgut to
the foregut.
• Echinoderms
→ In some, intracellular digestion is assisted or
exclusively carried out by wandering
phagocytic blood cells.
→ Sea stars have the ability to evert their
stomach to digest their prey in situ.
→ Sea urchin and sand dollars have a
complex chewing mechanism called the
“Aristotle’s lantern”
• Tetrapods
→ have specialized mouthparts and large
digestive glands (i.e. liver, pancreas) to
acquire and digest prey.