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Communication I Nerves: Neurons:

Neurons: The Basic Functional Units of the Nervous System
The functional unit of the nervous system is a highly specialized cell called the neuron (from the Greek
word "nerve"). Neurons are designed to produce signals that can be sent over short or long distances,
from one part of an animal’s body to another. Neurons have two key properties:

1. Excitability: The ability to respond to stimuli.

2. Conductivity: The ability to carry a signal.

Types of Neurons

There are three main types of neurons:

1. Sensory Neurons: Act as receptors of stimuli themselves or are activated by receptors. Changes
in the internal or external environments stimulate sensory neurons, which respond by sending
signals to the major integrating centers where information is processed.

2. Interneurons: Comprise the integrating centers and receive signals from the sensory neurons
and transmit them to motor neurons.

3. Motor Neurons: Send the processed information via a signal to the body's effectors (e.g.,
muscles), causing them to contract, or to glands, causing them to secrete.

Neuron Structure: The Key to Function

Most neurons contain three main parts:

1. Cell Body: Contains a large, central nucleus.

2. Dendrites: Short, thread-like branches that are extensions of the cell body and conduct signals
towards the cell body. The motor neuron has many dendrites (from the Greek word "dendron,"
meaning tree).

3. Axon: A relatively long, cylindrical process that conducts signals (information) away from the
cell body.

Sheathing of Neurons

1. Neurons without Sheath: The neurons of hydras and sea anemones do not have a sheath
covering the axon of the neuron.

2. Sheathed Neurons: Other invertebrates and all vertebrates have sheathed neurons. When
present, the laminated lipid sheath is called myelin. In some neurons, a neuromelocyte
(formerly known as a Schwann cell) wraps the myelin sheath in layers. In these neurons, gaps
called neurofibril nodes (formerly nodes of Ranvier) segment the myelin sheath at regular
intervals. The neuromelocyte also assists in the regeneration of injured myelinated neurons.

Function of the Nervous System

The nervous system has the following functions:

1. Receiving Data (Input Stimulus): In a given neuron, the dendrites are the receptors.

2. Integration: The cell body acts as the integrator.

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3. Effecting Change (Output Response): The ends of the axon are the effectors, resulting in a
change in the animal's physiology.

Neuron Communication
Neuron Signals

The signal of a neuron is the nerve impulse or action potential. The key to this nerve impulse is the
neuron's plasma membrane and its properties. Changes in membrane permeability and the movement
of ions produce a nerve impulse that travels along the plasma membrane of the dendrites, cell body, and
axon of each neuron.

Resting Membrane Potential

Definition

A "resting" neuron is not sending a nerve impulse. The plasma membrane of a resting neuron is
polarized; the fluid on the inner side of the membrane is negatively charged compared to the positively
charged fluid outside the membrane (see figure 24.3).

Cause of Polarization

The difference in electrical charge between the inside and outside of the membrane at any given point
is due to the active movement of positive and negative ions in the fluids on either side of the membrane,
and to the permeability of the plasma membrane to these ions. The difference in charge is called the
resting membrane potential. All cells have such a resting potential, but neurons and muscle cells are
unique in that they can change and recycle it quickly.

Measurement

The resting potential is measured in millivolts (mV). A millivolt is 1/1,000 of a volt. Normally, the
resting membrane potential is about -70 mV. Due to the unequal distribution of various electrically
charged ions, sodium (Na+) ions are more highly concentrated in the fluid outside the plasma
membrane, and potassium (K+) and large negative protein ions are more highly concentrated inside.

Ion Movement

The Na+ and K+ ions constantly diffuse through ion channels in the plasma membrane, moving from
areas of higher concentrations to areas of lower concentrations.

Ion Concentrations and the Sodium-Potassium Pump

There are also larger Cl- ions and large negative protein ions, which cannot move easily from the inside
of the neuron to the outside. However, the concentrations of Na+ and K+ ions on the two sides of the
membrane remain constant due to the action of the sodium-potassium ATPase pump, which is powered
by ATP (figure 24.3). The pump actively moves Na+ ions to the outside of the cell and K+ ions to the
inside of the cell. Because it moves three Na+ ions out for each two K+ ions that it moves in, the pump
helps to establish the resting potential of about -70 mV. Both ions leak back across the membrane—
down their concentration gradients. K+ ions, however, move more easily back to the outside, adding to
the positive charge there and contributing to the membrane potential of about -70 mV.

Mechanism of Neuron Action: Changing the Resting

Membrane Potential into the Action Potential (Nerve
Impulse)
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Resting Membrane Potential:

• Resting electrical potential across the plasma membrane is crucial for nerve impulse creation
and conduction.

• A stimulus reaching a threshold level initiates an impulse, causing depolarization of the

membrane.

Depolarization:

• Threshold stimulus increases Na ion permeability, leading to an inflow of Na ions.

• Positive Na ions influx reduces membrane polarity, termed depolarization.

Action Potential Generation:

• Depolarization triggers voltage-gated Na channels to open, allowing more Na ions inside.

• Subsequently, Na gates close, and voltage-gated K channels open, enabling rapid K ion efflux.

• Outward movement of K ions restores positive charge outside, repolarizing the membrane.

Conduction of Action Potential:

• Series of membrane changes induce depolarization and repolarization, propagating a wave

down the axon.

• Refractory period follows each action potential, during which hyperpolarization occurs before
restoration of resting potential.

All-or-None Law:

• Minimum threshold stimulus initiates action potential; increased stimulus intensity doesn't
affect its strength.

• Axon fires at full power or not at all, following the all-or-none law.

Speed of Impulse Conduction:

• Axon diameter and myelin sheath influence impulse conduction speed.

• Larger diameter and myelinated axons transmit impulses faster.

• Myelination insulates axons, allowing action potential "jumping" at neurofibril nodes, known
as saltatory conduction.

Evolutionary Adaptations:

• In invertebrates like crayfish and earthworms, large-diameter axons facilitate rapid escapes.

• Myelination in vertebrates enhances conduction velocity, enabling efficient nervous system

function in compact spaces.

Transmission of the Action Potential Between Cells

After an action potential (a nerve signal) travels along an axon, it reaches the end, called the end bulb.
The synapse is the connection point between the axon of one neuron and the dendrite of another neuron
or an effector cell (like a muscle or gland cell). The space between the end bulb and the next neuron’s
dendrite is called the synaptic cleft.
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Synapse Structure

• Presynaptic Neuron: The neuron sending the signal towards the synapse.

• Postsynaptic Neuron: The neuron receiving the signal after the synapse. This can also be a
muscle or gland cell.

Types of Synapses

• Electrical Synapse: Nerve impulses transmit directly from one neuron to another through ions
that move across the synapse, allowing rapid signal transmission in both directions. These are
common in fish, helping them react quickly to predators.

• Chemical Synapse: Neurons communicate via a chemical called a neurotransmitter. The

presynaptic neuron releases the neurotransmitter, which then binds to the receptor on the
postsynaptic neuron, creating a new action potential in the postsynaptic neuron.

Chemical Synapse Mechanism

1. Nerve Impulse Arrival: When a nerve impulse reaches an end bulb, it causes vesicles
containing neurotransmitters to fuse with the plasma membrane.

2. Neurotransmitter Release: The neurotransmitter is released into the synaptic cleft by

exocytosis.

3. Neurotransmitter Action: The neurotransmitter (e.g., acetylcholine) binds to receptors on the

postsynaptic membrane, causing depolarization and continuation of the nerve impulse.

4. Neurotransmitter Inactivation: Enzymes like acetylcholinesterase break down the

neurotransmitter to prevent continuous stimulation, which can cause a diseased state.

Practical Example

Flea sprays often contain parathion, which prevents the breakdown of acetylcholine in fleas, causing
continuous stimulation of their nerve cells and eventually immobilizing them. The dose is low enough
not to harm pets or humans significantly

Invertebrate Nervous Systems

Introduction to Nervous Systems

• All cells can respond to stimuli and send information both internally and externally. Even
protozoa and sponges, which lack a true nervous system, show coordination and reaction to
stimuli. For instance, protozoan cilia beat regularly, and flagellates respond to light changes,
requiring internal coordination.

• True nervous systems are present only in animals with tissue-level organization, excluding
protozoa and sponges.

Evolutionary Trends in Nervous System Development

• Trend 1: Increased Complexity

• More complex animals have more detailed nervous systems.

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• Cnidarians (e.g., hydras, jellyfish, sea anemones) possess the simplest nervous
systems, consisting of a nerve net. This latticework of neurons conducts impulses
bidirectionally. Cnidarians lack brains and clusters of neurons, with nerve impulses
spreading throughout the body.

• Trend 2: Cephalization

• As animals evolved to move forward, sensory organs and nervous tissue became
concentrated at the front (anterior end).

• Flatworms and Roundworms exhibit this trend. Flatworms have ganglia (clusters of
nerve cells) in the head region, functioning as a primitive brain. Lateral nerve cords
carry sensory information to and from these ganglia.

• Trend 3: Bilateral Symmetry

• Bilateral symmetry involves having symmetrical right and left body halves, leading
to paired neurons, muscles, sensory structures, and brain centers.

• This pairing allows coordinated movements, such as climbing, crawling, flying, or

walking.

• Trend 4: Increased Complexity in Nervous System Structure

• Invertebrates like crustaceans, segmented worms, and arthropods show advanced

nervous system organization.

• Axons join into nerve cords, and besides a small central brain, there are smaller
peripheral ganglia coordinating different body regions. These ganglia occur in each
body segment or near the organs they regulate.

• Trend 5: Increase in Interneurons

• More complex animals have more interneurons, leading to more complex behavior
patterns.

• In echinoderms (e.g., sea stars), the nervous system is divided into parts: the ectoneural
system for sensory and motor functions, a radial nerve for each arm, a deeper
hyponeural system for motor functions, and an apical system with potential sensory
functions.

• The brain, containing the highest number of neurons, becomes more complex in
animals with more intricate behaviors, like vertebrates.

Summary

• Nervous systems evolved from simple nerve nets in cnidarians to more complex
arrangements with central brains and peripheral ganglia in advanced invertebrates and
vertebrates.

• The increasing complexity of the nervous system in animals is characterized by cephalization,

bilateral symmetry, increased interneurons, and more specialized structures, enabling
more sophisticated behaviors and coordination.

Vertebrate Nervous System

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Key Characteristics

• Bilateral Symmetry: Vertebrates have symmetrical right and left halves.

• Notochord: A rod of tissue present in embryos, later replaced by the vertebral column in most
species.

• Tubular Nerve Cord: Located above the notochord, it develops into the spinal cord and brain.

Notochord and Vertebral Column

• The notochord is mesodermally derived and provides structural support.

• Vertebral Column: Replaces the notochord during development, allowing for strong muscles
and fast movement, facilitating predatory behavior.

• Jaw Development: Bones evolve into powerful jaws, enhancing predatory abilities.

Evolution of the Nerve Cord

• Single, Tubular Nerve Cord:

Located above the notochord, it
expands and specializes into the
spinal cord and brain.

• Brain Development: The

anterior end thickens and divides
into the hindbrain, midbrain, and
forebrain.

• Sensory Receptors: Paired

structures like eyes and ears
develop for better environmental
awareness.

Divisions of the Nervous System

• Central Nervous System (CNS): Composed of the brain and spinal cord, responsible for
information processing.

• Peripheral Nervous System (PNS): Consists of all nerves outside the brain and spinal cord,
divided into sensory and motor nerves.

Sensory and Motor Nerves

• Sensory (Afferent) Nerves: Transmit information to the CNS.

• Motor (Efferent) Nerves: Carry commands away from the CNS.

Motor Nerve Subdivisions

• Voluntary (Somatic) Nervous System: Controls skeletal muscles.

• Involuntary (Autonomic) Nervous System: Stimulates smooth and cardiac muscles, and
glands.

• Sympathetic System: Prepares the body for action.

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• Parasympathetic System: Calms the body and conserves energy.

Nerve Fiber Bundles

• CNS: Bundles of nerve fibers

are called tracts.

• PNS: Bundles of nerve fibers

are called nerves.

Cell Body Clusters

• CNS: Clusters of cell bodies

are called nuclei.

• PNS: Clusters of cell bodies

are called ganglia.

The Spinal Cord

The spinal cord is essential for two main reasons: it links the brain with most of the body, and it manages
spinal reflex actions.

Functions

• Connecting Link: Connects the brain to most of the body.

• Reflex Actions: Handles predictable, involuntary responses to stimuli, aiding both voluntary
and involuntary limb movements and certain organ functions.

Structure

• Extension: Part of the central nervous system, running from the brain to near or into the tail.

• Neural Canal: Contains cerebrospinal fluid.

• Gray Matter: Made up of cell bodies and dendrites, mainly handling reflex connections at
various levels.

• Ventral and Dorsal Roots: Extend from the spinal cord, containing motor and sensory fibers
that form major spinal nerves.

• White Matter: Named for the whitish myelin covering the axons.

Protective Layers

• Meninges: Three layers of protective membranes surrounding the spinal cord and brain.

• Dura Mater: Outer, tough, fibrous membrane.

• Arachnoid: Middle, delicate layer connecting to the pia mater.

• Pia Mater: Innermost layer with small blood vessels that nourish the spinal cord.

Spinal Nerves
Number of Spinal Nerves
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• Directly Related to Segments: The number of spinal nerves corresponds to the number of
segments in the trunk and tail of a vertebrate.

Examples

• Frog: Evolved strong hind legs for swimming or jumping, a reduced trunk, and no tail in the
adult; has only 10 pairs of spinal nerves.

• Snake: Moves by lateral undulations of its long trunk and tail; has several hundred pairs of
spinal nerves.

The Brain
Anatomical Development

• Vertebrate Brain: Develops at the anterior end of the spinal cord.

• Embryonic Development: Undergoes regional expansion, forming the hindbrain, midbrain,

and forebrain.

• Ventricles: The central canal of the spinal cord extends into the brain, expanding into chambers
called ventricles filled with cerebrospinal fluid.

Hindbrain
• Components: Includes the medulla oblongata, cerebellum, and pons.

Medulla Oblongata

• Location: Enlargement where the spinal cord enters the brain.

• Functions: Contains reflex centers for breathing, swallowing, cardiovascular function, and
gastric secretion.

• Development: Well-developed in all jawed vertebrates, reflecting its role in controlling visceral
functions and screening information entering or leaving the brain.

Cerebellum

• Function: Coordinates motor activity associated with limb movement, maintaining posture,
and spatial orientation.

• Structure:

• Cartilaginous Fishes: Distinct anterior and posterior lobes.

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• Teleosts: Large in active swimmers, small in inactive fishes.

• Amphibians: Rudimentary cerebellum due to simple locomotor patterns.

• Tetrapods: Laterally expanded for locomotor control of limb muscles.

• Birds and Mammals: Much larger due to complex locomotor patterns and
evolutionary history of limb development.

Pons

• Function: Acts as a bridge of transverse nerve tracts from the cerebrum of the forebrain to both
sides of the cerebellum.

• Connectivity: Contains tracts connecting the forebrain and spinal cord in all vertebrates.

Midbrain
• Original Function: Coordinated reflex
responses to visual input.

• Evolved Functions: Added roles in tactile

(touch) and auditory (hearing) input
without changing in size.

• Optic Tectum: The thickened region of

gray matter in the roof of the midbrain that
integrates visual and auditory signals.

Forebrain
• Evolutionary Changes: The forebrain has
significantly changed during vertebrate
evolution.

• Main Parts: The forebrain consists of the diencephalon and telencephalon.

Diencephalon

• Location: Lies just in front of the midbrain.

• Components: Pineal gland, pituitary gland, hypothalamus, and thalamus.

• Thalamus: Relays all sensory information to higher brain centers.

• Hypothalamus: Regulates body temperature, sexual drive, carbohydrate metabolism, hunger,

and thirst.

• Pineal Gland: Controls some body rhythms.

• Pituitary Gland: A major endocrine gland.

Telencephalon

• Evolutionary Expansion: Rapidly increased in size and complexity.

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• Corpus Striatum: Important in reptiles and birds for complex behavior patterns.

• Cerebrum:

• Structure: Divided into right and left cerebral hemispheres by a large groove.

• Function: Related to sensory and motor integration.

• Cerebral Cortex:

• Expansion: Progressively increased in size and complexity in mammals.

• Function: Contains primary sensory and motor areas, involved in the perception of
visual or auditory signals.

• Human Specificity: Includes areas for language processing, both written and spoken.

Cranial Nerves
• Cranial Nerves in Vertebrates: The peripheral nervous system of vertebrates includes paired
cranial nerves.

• Number of Pairs: Reptiles, birds, and mammals have 12 pairs, while fishes and amphibians
have 10 pairs.

• Types of Cranial Nerves:

• Sensory Nerves: Contain only sensory axons (e.g., optic nerve) that carry signals to
the brain.

• Mixed Nerves: Contain both sensory and motor axons (e.g., vagus nerve) with sensory
axons leading to the brain and motor axons leading to the heart and smooth muscles of
visceral organs in the thorax and abdomen.

The Autonomic Nervous System

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• Function: Composed of two divisions that act in opposition to control involuntary muscles
(smooth and cardiac) and glands.

• Parasympathetic Nervous System:

• Function: Operates during relaxation.

• Nerve Origin: Arises from the brain and sacral region of the spinal cord.

• Nerve Structure: Long efferent nerve fibers synapse at ganglia near organs, and short
efferent neurons extend from ganglia to organs.

• Sympathetic Nervous System:

• Function: Responsible for the "fight-or-flight" response.

• Nerve Origin: Arises from the thoracic and lumbar regions of the spinal cord.

• Nerve Structure: Short efferent fibers extend to ganglia near the spine, and long
efferent neurons extend from ganglia to each organ.

• Dual Input to Organs:

• Parasympathetic Input: Stimulates salivary gland secretions, intestinal movements,

contracts pupillary muscles, and relaxes sphincter muscles.

• Sympathetic Input: Inhibits salivary gland secretions and intestinal movements,

relaxes pupillary muscles, and contracts sphincters.
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Communication II: Senses:

Sensory Reception
• Aristotle's Five Human Senses: About two thousand years ago, Aristotle identified five
human senses—sight, hearing, smell, taste, and touch—commonly referred to as the “five
senses.”

• Additional Animal Senses: Today, zoologists know that animals also have other senses. For
example, invertebrates possess an impressive array of sensory receptors through which they
receive information about their environment.

• Examples of Invertebrate Sensory Receptors:

• Tactile receptors that sense touch

• Georeceptors that sense the pull of gravity

• Hygroreceptors that detect the water content of air

• Proprioceptors that respond to mechanically induced changes caused by stretching,

compression, bending, and tension

• Phonoreceptors that are sensitive to sound

• Baroreceptors that respond to pressure changes

• Chemoreceptors that respond to air- and waterborne molecules

• Photoreceptors that sense light

• Thermoreceptors that are influenced by temperature changes

• Vertebrate Sensory Abilities: Most vertebrates have a sense of equilibrium (balance) and a
sense of body movement, and they are also sensitive to fine touch, touch-pressure, heat, taste,
vision, olfaction, audition, cold, pain, and various other tactile stimuli.

• Circulatory and Digestive System Receptors: Receptors in the circulatory system register
changes in blood pressure and blood levels of carbon dioxide and hydrogen ions, and receptors
in the digestive system are involved in the perception of hunger and thirst.

• Importance of Senses: Overall, an animal’s senses limit and define its impression of the
environment. In fact, all awareness depends on the reception and decoding of stimuli from the
external environment and from within an animal’s body.

• Function of Sensory Receptors: Sensory receptors consist of cells that can convert
environmental information (stimuli) into nerve impulses.

• Stimulus Definition: A stimulus (pl., stimuli) is any form of energy an animal can detect with
its receptors.

• Transducer Function: All receptors are transducers (“to change over”); that is, they convert
one form of energy into another.

• Generator Potential: Different types of receptors convert different kinds of stimuli, such as
light or heat, into a local electrical potential called the generator potential. If the generator
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potential reaches the sensory neuron’s threshold potential, it causes channels in the plasma
membrane to open and creates an action potential.

• Action Potential and Sensations: All action potentials are alike. Furthermore, an action
potential is an all-or-none phenomenon; it either occurs or it doesn’t. In those animals that have
brains, some nerve signals from specific receptors always end up in a specific part of the brain
for interpretation.

• Intensity of Stimulus: When the stimulus strength increases, the number of action potentials
per unit of time also increases. Thus, the brain can perceive the intensity and type of stimulus
from the timing of the impulses and the “wiring” of neurons.

Basic Features of Sensory Receptors

1. Sensitive Receptor Cells: They contain sensitive receptor cells or finely branched peripheral
endings of sensory neurons that respond to a stimulus by creating a generator potential.

2. Specific Structure: Their structure is designed to receive a specific stimulus.

3. Synapse with Afferent Nerve Fibers: Their receptor cells synapse with afferent nerve fibers
that travel to the central nervous system along specific neural pathways.

4. Translation into Recognizable Sensation: In the central nervous system, the nerve impulse is
translated into a recognizable sensation, such as sound.

Invertebrate Sensory Receptors

An animal's behavior is largely based on how it responds to information from its environment.
Invertebrates have many different types of sensory receptors that help them gather this information.
Let's look at a few examples of these receptors and understand their structure and function.

Baroreceptors
Function: Baroreceptors detect changes in pressure.

Details: While no specific baroreceptor structures have been identified in invertebrates, many ocean-
dwelling invertebrates respond to pressure changes. Examples include:

• Copepod crustaceans

• Ctenophores

• Jellyfish medusae

• Squids

Some intertidal crustaceans may use pressure changes from water depth to coordinate their migratory
activities with the tides.

Chemoreceptors
Function: Chemoreceptors respond to chemicals in the environment.

Details: Chemoreception is a direct sense where specific molecules stimulate a response. It is the oldest
and most universal sense among animals. Examples of chemoreception include:

• Protozoa reacting to acids, alkalis, and salts with avoidance behavior.

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• Predatory ciliates being attracted to chemicals released by their prey.

In aquatic invertebrates, chemoreceptors are often located in pits or depressions that allow water
carrying chemicals to circulate through. In arthropods, chemoreceptors are usually found on the
antennae, mouthparts, and legs in the form of hollow hairs called sensilla. These sensilla contain
chemosensory neurons.

Types of Information Gathered:

• Humidity detection

• pH assessment

• Prey tracking

• Food recognition

• Mate location

For example, male silkworm moths (Bombyx

mori) can detect a single molecule of bombykol,
a sex attractant released by female moths, among
a trillion molecules of air. This ability helps
males locate females from several miles away,
which is a significant reproductive advantage for
this species.

Georeceptors
Function: Georeceptors respond to the force of gravity, providing an animal with information about its
orientation relative to "up" and "down."

Structure: Most georeceptors are statocysts,

which consist of a fluid-filled chamber lined
with cilia-bearing sensory epithelium. Inside
the chamber is a solid granule called a statolith.

Mechanism: When an animal moves, the

position of the statolith and the fluid within the
chamber change, altering the information from
the sensory epithelium. This provides
information about the animal’s linear and
rotational acceleration.

Examples: Statocysts are found in various gastropods, cephalopods, crustaceans, nemertines,

polychaetes, and scyphozoans. These animals use statocysts to orient themselves within their
environment. For instance, burrowing invertebrates rely on georeceptors rather than photoreceptors
for orientation within the substratum, while planktonic animals use statocysts to orient in their three-
dimensional aquatic environment, especially at night and in deep water.

Special Adaptation: Some aquatic insects detect gravity using air bubbles trapped in passageways,
such as tracheal tubes. These air bubbles act like the air bubble in a carpenter’s level, moving according
to gravity and stimulating sensory bristles that line the tubes.

Hygroreceptors
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Function: Hygroreceptors detect the water content of air.

Examples: Some insects have hygroreceptors that can detect small changes in ambient relative
humidity. This sense enables them to seek environments with specific humidity or to modify their
physiology or behavior (e.g., controlling the opening or closing of spiracles).

Location: Hygrosensory structures are found on the antennae, palps, underside of the body, and near
the spiracles of insects.

Unknown Mechanism: The exact process by which a hygroreceptor transduces humidity into an action
potential is not yet known.

Phonoreceptors
Function: Phonoreceptors respond to sound.

Presence: True phonoreceptors are found in insects, arachnids, and centipedes. Other invertebrates
may respond to sound-induced vibrations of the substratum.

Examples in Insects: Crickets, grasshoppers, and cicadas possess phonoreceptors called tympanic or
tympanal organs, which have a tough, flexible tympanum covering an internal sac. When sound waves
strike the tympanum, it vibrates, stimulating sensory neurons attached to it and producing a generator
potential.

Examples in Arachnids: Most arachnids possess phonoreceptors in their cuticle called slit sense
organs that can sense sound-induced
vibrations.

Examples in Centipedes: Centipedes

have organs of Tomosvary, which may
be sensitive to sound, though the
physiology of both slit sense organs and
organs of Tomosvary is poorly
understood.

Photoreceptors
Function: Photoreceptors are sensitive to
light.

Light-Sensitive Pigments: All photoreceptors have pigments (like carotenoids and rhodopsin) that
absorb light and produce a generator potential.

Types and Examples:

• Euglena: These flagellated protozoa have photoreceptor granules called stigmata (red
pigments). The actual photoreceptor is at the base of the flagellum, helping Euglena orient
toward light for photosynthesis.

• Earthworm (Lumbricus): Simple unicellular photoreceptor cells scattered over the epidermis
or in specific areas.

• Ocelli: Small cups lined with light-sensitive receptors and backed by light-absorbing pigment.
Found in Annelida, Mollusca, and Arthropoda, ocelli provide information about light direction
and intensity but do not form images.
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• Compound Eyes: Made up of many units called ommatidia. Best developed in arthropods,
these eyes detect movement and may form images. Each ommatidium has its own nerve tract,
and the eye adapts to light changes and may provide color vision.

• Complex Camera Eyes: Found in squids and octopuses, these eyes are similar to vertebrate
eyes. They have a cornea, lens, and retina. The receptor sites face the incoming light, unlike in
vertebrates where they are inverted. Squid eyes focus by moving the lens, while vertebrate eyes
focus by changing the lens shape.

Proprioceptors
Function: Proprioceptors, also known as "stretch receptors," respond to mechanical changes like
stretching, compression, bending, or tension.

Role: They provide information about the movement and position of an animal's body parts.

Examples in Arthropods: Proprioceptors are associated with appendage joints and body extensor
muscles. Sensory neurons are attached to parts of the body that stretch, such as muscle cells, connective-
tissue fibers, or membranes spanning joints. Changes in shape of these structures distort sensory nerve
endings, initiating a generator potential.

Key Terms and Their Meanings

• Georeceptors (earth receptor): Respond to gravity.

• Statocysts (standing bladder): Fluid-filled chambers with sensory epithelium and a solid
granule (statolith) that detect movement.

• Hygroreceptors (moist receptor): Detect humidity.

• Phonoreceptors (voice receptor): Respond to sound.

• Photoreceptors (light receptor): Sensitive to light, containing pigments like carotenoids and
rhodopsin.

• Stigma (eye-spot): Mass of photoreceptor granules in Euglena.

• Ocellus (small eye): Simple light-detecting organ.

• Ommatidia (little eye): Units making up compound eyes.

• Proprioceptors (self receptor): Internal receptors for body part movement and position.
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Tactile Receptors
Function: Tactile (touch) receptors detect physical contact with objects in the environment.

Structure: These receptors are generally derived from modifications of epithelial cells associated with
sensory neurons. They often involve projections from the body surface, such as bristles, spines, setae,
and tubercles.

Mechanism: When an animal contacts an object, these receptors are mechanically deformed. This
deformation activates the receptor, which in turn activates underlying sensory neurons, initiating a
generator potential.

Examples:

• Tube-dwelling polychaetes: Have receptors that allow them to retract quickly into their tubes
in response to movements in their surroundings.

• Web-building spiders: Possess tactile receptors that can sense struggling prey through
vibrations of the web threads.

Thermoreceptors
Function: Thermoreceptors (heat receptors) respond to temperature changes.

Sensing Mechanism: Some invertebrates can directly sense differences in environmental temperatures.

Examples:

Protozoan Paramecium: Collects in areas where water temperature is moderate and avoids
temperature extremes.

Leeches and Ticks: Attracted to warm-blooded hosts through a heat-sensing mechanism.

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Certain Insects, Some Crustaceans, and Horseshoe Crab (Limulus): Can sense thermal variations.

Unknown Structure: In all these cases, the specific receptor structures responsible for thermoreception
have not been identified.

Summary

Main Points:

• Tactile Receptors: Respond to mechanical deformation and vibrations, essential for detecting
contact and movement.

• Thermoreceptors: Detect temperature changes, important for environmental adaptation.

• Environmental Adaptations: Sensory receptors evolve based on environmental demands,

highlighting the intricate relationship between sensory systems and habitats.

• Vertebrate and Invertebrate Similarities: Both groups have sensory receptors adapted to
their environments, underscoring the universality of sensory adaptation in the animal kingdom.

Vertebrate Sensory Receptors

Adaptations to Environments: Vertebrate sensory receptors reflect adaptations to the sensory stimuli
present in different external and internal environments. Each environment has specific chemical and
physical characteristics that affect the types of energy and molecules that carry sensory information.

Examples of External Environments:

• Human: The earth and air.

• Trout: Cool, clear water of a mountain stream.

• Turtle: Turbid water of a swamp.

• Salmon: Salty water of the sea.

Media Characteristics:

• Air: Transmits light well, conducts sound efficiently, carries a limited number of small
molecules detectable by smell, and passes little or no electrical energy.

• Water: Sound travels faster and farther, dissolves and carries a wide range of chemicals,
conducts electricity well (especially seawater), but absorbs many wavelengths of light.

Evolutionary Adaptations: Vertebrate sensory receptors, like invertebrate sensory receptors, have
evolved to function effectively in their environments. Despite different habitats, many underlying
similarities unite vertebrate senses. Each sense involves:

1. Environmental Information: The type of sensory information present in the environment.

2. Receptor Cell Adaptation: How receptor cells evolved to detect this information.

3. Information Processing: How the central nervous system processes this information for the
animal's use.

Lateral-Line System and Electrical Sensing

 21

Function: Specialized organs for balance, gravity detection, hearing, and magnetoreception evolved
from the lateral-line system of fishes. This system is used for electrical sensing in most fishes, some
amphibians, and the platypus.

Structure:

• Sensory Pores: Located in the skin's surface, leading to canals connected to electroreceptors
called ampullary organs.

• Ampullary Organs: These electroreceptors can sense electrical currents in the surrounding
water.

Benefits: The ability to detect weak electrical fields generated by living organisms helps fish find mates,
capture prey, and avoid predators. This sense is especially valuable in deep, turbulent, or murky water
where vision is limited. Some fishes generate electrical fields and use their electroreceptors for
electrocommunication, detecting how surrounding objects distort the field to navigate in murky or
turbulent waters.

Vertebrate Sensory Receptors

Adaptations to Environments: Vertebrate sensory receptors reflect adaptations to the sensory stimuli
present in different external and internal environments. Each environment has specific chemical and
physical characteristics that affect the types of energy and molecules that carry sensory information.

Examples of External Environments:

• Human: The earth and air.

• Trout: Cool, clear water of a mountain stream.

• Turtle: Turbid water of a swamp.

• Salmon: Salty water of the sea.

Media Characteristics:

• Air: Transmits light well, conducts sound efficiently, carries a limited number of small
molecules detectable by smell, and passes little or no electrical energy.

• Water: Sound travels faster and farther, dissolves and carries a wide range of chemicals,
conducts electricity well (especially seawater), but absorbs many wavelengths of light.

Evolutionary Adaptations: Vertebrate sensory receptors, like invertebrate sensory receptors, have
evolved to function effectively in their environments. Despite different habitats, many underlying
similarities unite vertebrate senses. Each sense involves:

1. Environmental Information: The type of sensory information present in the environment.

2. Receptor Cell Adaptation: How receptor cells evolved to detect this information.

3. Information Processing: How the central nervous system processes this information for the
animal's use.

Lateral-Line System and Mechanoreception

Mechanoreceptor: A receptor excited by mechanical pressures or distortions (e.g., sound, touch, and
muscular contractions).
 22

Lateral-Line System: This system is present in cyclostomes, sharks, some advanced fishes, and aquatic
amphibians. It includes various hair-cell mechanoreceptors called neuromasts, which are found in pits
along the body, but not in the head region.

Function of Neuromasts: These are responsive to local water displacement or disturbance. When water
moves near the lateral line, it moves the water in the pits and distorts the hair cells, causing a generator
potential in the associated sensory neurons. This allows the animal to detect the direction and force of
water currents and the movement of other animals or prey in the water.

Hearing and Equilibrium in Air

Hearing: Initially important to vertebrates as a mechanism to alert them to potentially dangerous

activity, and later became important for finding food, mates, and communication.

Ear Functions:

1. Auditory Apparatus: Concerned with hearing.

2. Vestibular Apparatus: Concerned with posture and equilibrium.

Sound Transmission: Sound results from pressure waves transmitting energy through a medium, such
as air or water. In air, sound produces less than 0.1% of the pressure it produces in water, requiring
adaptation for effective hearing.

Evolution of Tympanum: Adaptation to hearing in air resulted in the evolution of an acoustic

transformer incorporating a thin, stretched membrane (eardrum or tympanic membrane). This structure
first evolved in amphibians.

Anuran (Frog) Ear Structure:

• Tympanum: Modified integument stretched over a cartilaginous ring that vibrates in response
to sounds.

• Middle Ear: A chamber behind the tympanum with a small bone called the columella (or
stapes) that touches the oval window.

• Inner Ear: Contains fluid-filled semicircular canals where pressure waves stimulate receptor
cells.

High-Frequency Sounds: Transmitted through the middle ear via the columella, causing pressure
waves in the inner ear fluid. Low-Frequency Sounds: Substrate-borne vibrations transmitted through
front appendages and the pectoral girdle cause the operculum to vibrate, stimulating a second patch of
sensory receptor cells.
 23

Adaptations in Other Vertebrates:

• Salamanders: Lack a tympanum and middle ear, sensing sounds transmitted through the
substratum and skull.

• Reptiles: Ear structures vary, with some lacking a middle-ear cavity and tympanum. In snakes,
a jaw bone articulates with the stapes to receive vibrations of the substratum.

• Birds: Well-developed hearing with external ear openings covered by delicate feathers.

• Mammals: Adaptations include an ear flap (auricle) and auditory tube, leading to the
tympanum and middle ear with three ossicles (malleus, incus, and stapes). The cochlea in the
inner ear provides greater sensitivity to pitch and volume.

Human Ear Structure:

• Outer Ear: Auricle and external auditory canal.

• Middle Ear: Begins at the tympanic membrane and ends at the oval and round windows.

• Inner Ear: Vestibule and semicircular canals for equilibrium, and cochlea for hearing.
 24

Hearing Process:

1. Sound waves enter the outer ear, creating pressure waves that reach the tympanic membrane.

2. Tympanic membrane vibrates, moving the malleus.

3. Malleus articulates with the incus, vibrating it.

4. Incus moves the stapes against the oval window.

5. Oval window movements create pressure changes in the inner ear fluid, transmitted to the
basilar membrane.

6. Receptor hair cells in the organ of Corti bend, causing a generator potential and an action
potential that travels to the brain.

7. Vibrations dissipate through movements of the round window.

Equilibrium (Balance):

• Static Equilibrium: Sensing movement in one plane (vertical or horizontal).

• Dynamic Equilibrium: Sensing angular and/or rotational movement.

• Otoliths: Rest on hair cells in semicircular canals, displacing and bending hairs during
movement, initiating a generator potential.

Motion Sickness: Continuous fluid movement in the semicircular canals can cause motion sickness or
seasickness in humans.

Hearing and Equilibrium in Water

Inner Ear Functions in Bony Fishes: In bony fishes, the receptors for equilibrium, balance, and
hearing are located in the inner ear, similar to other vertebrates.

• Semicircular Canals: Detect rotational movements.

 25

• Sensory Patches: Assist with equilibrium and balance by detecting the direction of
gravitational pull.

Vibration Detection: Fishes lack the outer and middle ear found in other vertebrates. Therefore,
vibrations pass from the water through the bones of the skull to the inner ear.

• Swim Bladder and Ossicles: Some fishes have chains of bony ossicles (modified vertebrae)
between the swim bladder and the back of the skull. Vibrations are amplified by the swim
bladder and transmitted through the ossicles to the skull.

Skin Sensors of Damaging Stimuli

Pain Receptors (Nociceptors): Bare sensory nerve endings present throughout the body of mammals,
except for the brain and intestines. These nerve endings respond to severe heat, cold, irritating
chemicals, and strong mechanical stimuli (e.g., penetration), which the brain interprets as pain or
itching.

Skin Sensors of Heat and Cold

Thermoreceptors: Bare sensory nerve endings in either the epidermis or dermis that respond to
temperature changes.

• Cold and Warm Spots: Specific areas of the skin sensitive to either cold or warm stimuli.

• Cold Receptors: Respond to temperatures below skin temperature.

• Heat Receptors: Respond to temperatures above skin temperature.

Materials and Temperature Sensations: Materials coming into contact with the skin need not be
warm or cold to produce temperature sensations.

• Example: Metal feels colder than wood because it absorbs more heat from the skin.

Heat-Sensitive Pit Organs in Snakes: Rattlesnakes and other pit vipers have heat-sensitive pit organs
on each side of the face between the eye and nostril. These organs help snakes locate warm-blooded
prey by detecting temperature differences from the surroundings.

Auditory Sonar System in Bats

Sophisticated Imaging: Bats use an auditory sonar system to create three-dimensional imaging for
locating and catching prey. This system provides highly detailed information about the environment.

Skin Sensors of Mechanical Stimuli

Tactile Stimuli: Many animals rely on touch to get information about their environment. Mechanical
sensory receptors in vertebrate skin detect stimuli that the brain interprets as light touch, touch-
pressure, and vibration.

• Light Touch: Perceived when the skin is touched but not strongly deformed. Receptors for
light touch include bare sensory nerve endings and tactile (Meissner’s) corpuscles.

• Bare Sensory Nerve Endings: Widely distributed in the vertebrate body and involved with
pain, thermal stimuli, and light touch.

• Bulbs of Krause: Found in the dermis in certain parts of the body, these receptors respond to
some physical stimuli, such as position changes.

• Touch-Pressure Receptors: Include Pacinian corpuscles and organs of Ruffini.

 26

• Sensory Hairs (Vibrissae): Many mammals have specialized sensory hairs on their wrists,
snouts, and eyebrows. Nerves around the base of each vibrissa carry impulses from
mechanoreceptors to the brain.

Sonar
Echolocation (Sonar): Bats, shrews, certain cave-dwelling birds, whales, and dolphins can determine
distance and depth using echolocation.

• High-Frequency Sounds: These animals emit high-frequency sounds and determine how long
it takes for the sounds to return after bouncing off objects.

• Example: Bats emit clicks that last 2-3 milliseconds, repeated several hundred times per
second. The returning echo helps the bat locate and catch prey.

• Sophisticated Imaging: This auditory sonar system provides detailed three-dimensional

imaging.

Smell (Olfaction)

Olfaction: The sense of smell is due to olfactory neurons in the roof of the vertebrate nasal cavity.

• Olfactory Cells: Specialized endings of fibers that make up the olfactory nerve, lying among
supporting epithelial cells.

• High Density: For example, a dog has up to 40 million olfactory receptor cells per square
centimeter.

• Odor Perception: Odor molecules interact with protein receptors on the cell membrane,
altering membrane permeability and leading to a generator potential.

Olfaction in Fishes:

• External Nares: Openings in the snout lead to olfactory receptors.

• Salmon and Lampreys: Use their sense of smell to return to spawn in the same streams where
they hatched, guided by characteristic odors.

Olfaction in Amphibians:

• Used in mate recognition, detecting noxious chemicals, and locating food.

Olfaction in Reptiles:

• Jacobson’s (Vomeronasal) Organs: Blind-ending pouches that open into the mouth, best
developed in snakes and lizards. These organs help perceive odor molecules.

• Forked Tongues: Snakes and lizards use their tongues to sample airborne chemicals and bring
them to Jacobson’s organs.

Olfaction in Birds:

• Plays a minor role for most birds. Vultures are an exception, using smell to locate prey.

Olfaction in Mammals:

• Mammals can detect olfactory stimuli over long distances and use them to locate food,
recognize members of the same species, and avoid predators.
 27

Taste (Gustation)

Receptors for Taste: Chemoreceptors that may be on the body surface or in the mouth and throat.

• Tongue Papillae: Small protuberances covering the surface of the mammalian tongue.

• Taste Buds: Specialized receptors located in the crevices between the papillae, consisting of
clusters of chemoreceptor cells (gustatory cells) and supporting cells.

• Taste Sensations: Generally recognized as sweet (sugars), sour (acids), bitter (alkaloids), and
salty (electrolytes).

• Taste Mechanism: The exact mechanism that stimulates a chemoreceptor taste cell is not
known. One theory suggests that different gustatory stimuli change the permeability of the
receptor-cell membrane, leading to a generator potential.

Taste in Different Vertebrates:

• Reptiles and Birds: Taste buds are typically located in the pharynx rather than on the tongue.

• Fishes and Amphibians: Taste buds may also be found in the skin. For example, in sturgeons,
taste buds are abundant on the head projection called the rostrum.

• Distribution in Fishes: Taste buds are distributed in various locations, including the pharynx,
skin, and barbels (whiskers) of catfish. They help monitor the flow of water and detect potential
food sources.

Vision (Photoreception)
Overview: Vision is the primary sense used by vertebrates in light-filled environments, facilitated by
well-developed photoreceptive structures.

Eye Structure:

• Lens: Focuses light onto the retina.

• Sclera: Tough outer coat of the eyeball.

• Choroid Layer: Thin middle layer.

• Retina: Inner layer containing light-sensitive photoreceptor cells.

• Cornea: Transparent covering over the front of the eyeball.

• Iris: Contains pigments and muscles regulating light entering the pupil.

• Aqueous Humor: Fluid filling anterior and posterior chambers.

• Vitreous Body: Jellylike substance behind the lens.

• Conjunctiva: Moist mucous membrane covering the eyeball.

Accommodation:

• Definition: Adjustment of vision for light from close-up or distant objects.

• Mechanism: Coordinated stretching and relaxation of eye muscles and fibers for lens
adjustment.
 28

Adaptations Across Vertebrates:

• Fishes: Lidless eyes with a rounded lens close to the cornea, focusing by moving the lens
forward or backward.

• Amphibians: Varied eye adaptations for terrestrial environments, including binocular vision
for prey capture
and lateral eyes for
peripheral vision.

• Reptiles: Similar
eye structure to
amphibians, with
upper and lower
eyelids, a
nictitating
membrane, and a
protective
spectacle.

• Birds: Unique
double-focusing
mechanism for rapid adjustments during flight, with a nictitating membrane for eye protection.

Retinal Photoreceptors:

• Rods: Sensitive to dim light.

• Cones: Respond to high-intensity light and enable color perception.

• Rhodopsin: Pigment in rod cells absorbing light energy, triggering nerve impulses.

• Color Perception: Mediated by cones with red-, green-, or blue-absorbing pigments,

translating light absorption into nerve impulses for brain interpretation.
 29

Communication III: The Endocrine System and

Chemical Messengers:

Chemical Messengers in Animal Development

The development of most animals starts with fertilization and the subsequent division of the zygote.
Further development depends on continuous cell division, growth, and specialization. The integration
of these events, as well as the communication and coordination of physiological processes like
metabolism, breathing, waste removal, movement, and reproduction, depend on chemical messengers—
molecules that specialized cells produce and release. Chemical messengers can be categorized as
follows:

1. Local Chemical Messengers

Many cells release chemicals that change physiological conditions in the nearby area. Most of these
chemicals act on nearby cells and do not accumulate in the blood. Vertebrate examples include some of
the chemicals called lumones that the gut produces and that help regulate digestion. In a wound, mast
cells release a substance called histamine that participates in the inflammatory response.

2. Neurotransmitters

As mentioned in chapter 24, neurons release chemicals called neurotransmitters (e.g., nitric oxide and
acetylcholine) that act on immediately adjacent target cells. These chemical messengers reach high
concentrations in the synaptic cleft, act quickly, and are actively broken down and recycled.

3. Neuropeptides

Some specialized neurons (called neurosecretory cells) release neuropeptides (neurohormones). The
blood or other body fluids transport neuropeptides to distant target cells, where neuropeptides exert
their effects. In mammals, for example, certain nerve cells in the hypothalamus release a neuropeptide
that causes the pituitary gland to release the hormone oxytocin, which induces powerful uterine
contractions during childbirth.

4. Hormones

Endocrine glands or cells release hormones that the bloodstream transports to distant target cells. Many
examples are given in the rest of this chapter.

5. Pheromones

Pheromones are chemical messengers released to the exterior of one animal that affect the behavior of
another individual of the same species.

Neuroendocrine System

Scientists now recognize that the nervous and endocrine systems work together as a combined
communicative and integrative network called the neuroendocrine system. In this system, feedback
systems regulate chemical messengers in their short- and long-term coordination of animal body
functions to maintain balance (homeostasis).
 30

Hormones and Their Feedback Systems

A hormone (from the Greek word "hormaein," meaning to set in motion or to spur on) is a specialized
chemical messenger produced and released by an endocrine gland or tissue. The study of endocrine
glands and their hormones is called endocrinology. Hormones circulate through body fluids and affect
the metabolic activity of a target cell or tissue in a specific way. By definition, a target cell has receptors
that specifically bind to these chemical messengers or are affected by them.

Typically, one hormone influences, depends on, and balances another hormone within a controlled
feedback network. It is rare for a hormone to operate independently.

Biochemistry of Hormones
Most hormones fall into one of the following categories:

• Proteins (Polypeptides): For example, hormones that the vertebrate pancreas secretes.

• Derivatives of Amino Acids (Amines): For instance, hormones secreted by the thyroid gland.

• Steroids: Secreted by the ovaries, testes, and the cortex of the adrenal glands.

• Fatty Acid Derivatives: Although less common, some hormones fall into this category.

In invertebrates, neurosecretory cells often produce polypeptides called neuropeptides.

Hormone Function and Effects

Hormones are effective in extremely small amounts. Just a few molecules of a hormone can produce a
significant response in a target cell. In the target cell, hormones help control biochemical reactions in
three primary ways:

1. Regulating Substance Movement: A hormone can increase the rate at which other substances
enter or leave the cell.

2. Stimulating Synthesis: A hormone can stimulate a target cell to synthesize enzymes, proteins,
or other substances.

3. Activating or Suppressing Enzymes: A hormone can prompt a target cell to activate or

suppress existing cellular enzymes.

Like enzymes, hormones are not changed by the reactions they regulate.

Feedback Control System of Hormone Secretion

Although hormones are always present in some amount in endocrine cells or glands, they are not
secreted continuously. Instead, glands secrete the amount of hormone needed to maintain balance in the
body, known as homeostasis. A feedback control system monitors changes in the body or the external
 31

environment and sends information to a central control unit (such as the central nervous system), which
makes adjustments.

Types of Feedback Systems

• Negative Feedback System: This system produces a response that counteracts the initiating
stimulus. It helps maintain stability and homeostasis. For example, if a process slows down, the
feedback system will stimulate it to speed up and vice versa.

• Positive Feedback System: This system reinforces the initial stimulus, leading to an increase
in the response. Positive feedback systems are rare in animals because they can lead to
instability or pathological states.

Example of Negative Feedback

Suppose the metabolic rate (rate of chemical activity) in the body cells of a dog slows down:

1. Detection: The hypothalamus detects the slower metabolic rate.

2. Response by Hypothalamus: The hypothalamus releases more thyrotropin-releasing hormone

(TRH).

3. Response by Pituitary Gland: TRH causes the pituitary gland to secrete more thyrotropin,
also known as thyroid-stimulating hormone (TSH).

4. Response by Thyroid Gland: TSH stimulates the thyroid gland to secrete thyroxine.

5. Effect of Thyroxine: Thyroxine increases the metabolic rate, restoring homeostasis.

Conversely, if the metabolic rate speeds up:

1. Detection: The hypothalamus detects the faster metabolic rate.

2. Response by Hypothalamus: The hypothalamus releases less TRH.

3. Response by Pituitary Gland: The pituitary gland secretes less TSH.

4. Response by Thyroid Gland: The thyroid gland secretes less thyroxine.

5. Effect of Reduced Thyroxine: The metabolic rate decreases, restoring homeostasis.

In summary, negative feedback systems monitor hormone levels and adjust cellular activity as needed
to maintain a stable internal environment.

Mechanisms of Hormone Action

 32

Hormones modify the biochemical activity of target cells or tissues through two basic mechanisms: the
fixed-membrane-receptor mechanism and the mobile-receptor mechanism.

Fixed-Membrane-Receptor Mechanism
This mechanism applies to hormones that are proteins or amines. These hormones are water-soluble
and cannot diffuse across the plasma membrane, so they initiate their response by means of specialized
receptors on the plasma membrane of the target cell.

Steps of Fixed-Membrane-Receptor Mechanism

1. Hormone Secretion: An endocrine cell secretes a water-soluble hormone that circulates

through the bloodstream.

2. Binding to Receptor: At the cells of the target organ, the hormone acts as a "first messenger"
by binding to a specific receptor on the plasma membrane.

3. Activation of Adenylate Cyclase: The hormone-receptor complex activates the enzyme

adenylate cyclase in the membrane.

4. Formation of Cyclic AMP: The activated enzyme converts

ATP into cyclic AMP (cAMP), which acts as the "second
messenger."

5. Activation of Protein Kinase: Cyclic AMP diffuses

throughout the cytoplasm and activates an enzyme called
protein kinase.

6. Cellular Response: Protein kinase causes the cell to

respond with its specific physiological activity.

7. Inactivation: The enzyme phosphodiesterase inactivates

cyclic AMP, and the receptor on the plasma membrane
becomes available for a new reaction.

Mobile-Receptor Mechanism
This mechanism applies to steroid hormones, which are lipid-soluble and can diffuse easily into the
cytoplasm of target cells, where they initiate their response by binding to cytoplasmic receptors.

Steps of Mobile-Receptor Mechanism

1. Hormone Release: The steroid hormone is released from a carrier protein in the bloodstream.

2. Diffusion into Cell: The steroid hormone enters the target cell by diffusion.

3. Binding to Cytoplasmic Receptor: The hormone binds to a specific protein receptor in the
cytoplasm.
 33

4. Formation of Steroid-Protein Complex: The steroid-protein complex gains an affinity for

DNA and enters the nucleus.

5. Binding to DNA: The complex binds to DNA and regulates

the transcription of specific genes to form messenger RNA
(mRNA).

6. mRNA Translation: The newly transcribed mRNA leaves

the nucleus and moves to the rough endoplasmic reticulum,
where it initiates protein synthesis.

7. Protein Synthesis: The newly synthesized proteins, which

may include enzymes, affect cellular metabolism and
constitute the cellular response to the specific steroid
hormone.

These mechanisms illustrate how hormones can significantly influence cellular activities, either by
triggering enzyme actions through a fixed-membrane-receptor pathway or by regulating gene
expression via the mobile-receptor mechanism.

Hormones of Invertebrates
The survival of animals depends on their ability to grow, mature, and reproduce at times when
conditions such as climate and food supply are optimal. Invertebrates rely on various hormones to
regulate these processes. Most invertebrate hormones are neurosecretions called neuropeptides, though
some more complex invertebrates have additional types of hormones. Here's a brief overview of the
hormonal systems in some invertebrate groups:

Porifera (Sponges)
• Hormonal System: Sponges do not have classical endocrine glands or neurons, and thus, they
do not have neurosecretory cells.

Cnidarians (e.g., Hydra)

• Growth-Promoting Hormone: Cnidarians have nerve cells that contain a hormone stimulating
budding, regeneration, and growth. This hormone, known as the "head activator," accelerates
head regeneration and stimulates mitosis.

Platyhelminths (Flatworms)
• Neurosecretory Cells: These cells are found in the cerebral ganglion and along major nerve
cords.

• Functions: The neuropeptides produced by these cells regulate regeneration, asexual

reproduction, and gonad maturation. For instance, in tapeworms, neurosecretory cells in the
scolex control the shedding of proglottids or the initiation of strobilization.

Nemerteans (Ribbon Worms)

• Cephalization and Brain Structure: Nemerteans have a higher degree of cephalization and a
larger brain composed of dorsal and ventral ganglia connected by a nerve ring.

• Neuropeptides: The neuropeptides produced by these ganglia control gonadal development

and water balance.
 34

Nematodes (Roundworms)
• Neurosecretory Cells: These cells are associated with the central nervous system, even though
nematodes lack classical endocrine glands.

• Ecdysis Regulation: The neuropeptide produced by these cells controls ecdysis (shedding of
the cuticle). After a new cuticle is produced, the neuropeptide stimulates the excretory gland to
secrete an enzyme (leucine aminopeptidase) into the space between the old and new cuticles.
The fluid accumulation in this space causes the old cuticle to split and be shed.

• Hormones of Invertebrates

• Invertebrates rely on various hormones to regulate growth, maturation, and reproduction. Here's
an overview of the hormonal systems in some invertebrate groups:

Molluscs
• Neurosecretory Cells: Molluscs have a central nervous system with neurosecretory cells.

• Functions: The neuropeptides regulate heart rate, kidney function, and energy metabolism.

• Gastropods (e.g., Helix): Specific hormones stimulate spermatogenesis, egg development, and
the function of accessory sex organs. A growth hormone controls shell growth.

• Cephalopods (e.g., octopus, squid): The optic gland produces hormones that stimulate egg
development, proliferation of spermatogonia, and development of secondary sexual
characteristics.

Annelids
• Endocrine System: Annelids have a developed endocrine system due to their well-developed
nervous system, circulatory system, and coelom.

• Functions: Involved in morphogenesis, development, growth, regeneration, and gonadal

maturation.

• Polychaetes: Juvenile hormone inhibits gonads and stimulates growth and regeneration.
Gonadotropin stimulates egg development.

• Leeches: A neuropeptide stimulates gamete development and triggers color changes.

• Oligochaetes: Osmoregulatory hormones are reported, and a hyperglycemic hormone

maintains high blood glucose levels in Lumbricus.

Arthropods
• Endocrine System: Controls growth, maturation, and reproduction.

• Crustaceans (e.g., crayfish):

 35

• Ecdysis (Molting): Controlled by X-organs and Y-organs. X-organs produce molt-inhibiting

hormone (MIH), which keeps the Y-organ inactive. When MIH release stops, the Y-organ
releases ecdysone, leading to molting.

• Insects:

• Molting: The central nervous system activates

neurosecretory cells, which secrete ecdysiotropin. This
leads to the production of thoracotropic hormone by the
corpora cardiaca, stimulating the prothoracic glands to
release ecdysone for molting.

• Bursicon: Influences epidermal development, such as

tanning (hardening and darkening of the cuticle).

• Juvenile Hormone (JH): Produced by corpora allata.

High JH levels inhibit differentiation. Reduced JH
production leads to differentiation into a pupa and later
into an adult.

Echinoderms
• Hormonal System: Different from chordates, despite being deuterostomes.

• Neuropeptide: Radial nerves contain a gonad-stimulating

substance. This induces gamete shedding, spawning
behavior, and meiosis in oocytes when injected into a mature
sea star. It also releases a hormone called maturation-
inducing substance, affecting the reproductive system.

AN OVERVIEW OF THE
VERTEBRATE ENDOCRINE SYSTEM
The vertebrate endocrine system is the best-understood system of
hormonal control due to extensive study. As vertebrates evolved,
specialized hormone-producing cells and tissues emerged, controlled
in various ways:

• Nerve Cells and Brain Control: Certain endocrine tissues,

such as the medullary areas of the adrenal glands, are directed
by sets of nerve cells in the brain.

• Hypothalamus and Pituitary Gland Control: The

hypothalamus and the pituitary gland (hypophysis) exert
control over many other endocrine functions.

• Independent Endocrine Tissues: Some endocrine tissues operate independently of both nerve
signals and the pituitary gland.

• Types of Glands in Vertebrates

Vertebrates have two main types of glands:

Exocrine Glands:

• Definition: Secrete chemicals into ducts.

 36

• Function: These ducts then empty the chemicals into body cavities or onto body surfaces.

• Examples: Mammary glands (milk), salivary glands (saliva), and sweat glands (sweat).

Endocrine Glands:

• Definition: Do not have ducts.

• Function: Secrete hormones directly into the surrounding tissue space. From there, hormones
diffuse into the bloodstream, which transports them throughout the body to reach their target
cells.

Hormonal Control Mechanisms

• The endocrine system uses hormones as chemical messengers, which are crucial for regulating
various physiological processes. The hormones produced by endocrine glands reach their
specific target cells through the bloodstream, influencing functions such as growth, metabolism,
and reproduction.

• Understanding the vertebrate endocrine system highlights the complex interaction between
different control mechanisms—neural, hypothalamic-pituitary, and independent endocrine
functions—that collectively maintain homeostasis and coordinate bodily functions.

Endocrine Systems of Vertebrates Other Than Birds or Mammals

Since this text uses a phylogenetic approach to describe form and function in animals, endocrine
regulation in vertebrates other than birds or mammals is now discussed. Birds and mammals are the
subject of the last part of this chapter. Vertebrates other than birds or mammals have somewhat similar
endocrine systems, but differences do exist.

Species Differences in Endocrine Systems

Recent research has revealed the following three aspects of endocrinology that relate to species
differences among these vertebrates:

1. Hormone Variations: Hormones (or neuropeptides) with the same function in different species
may not be chemically identical.

2. Species-Specific Hormones: Certain hormones are species-specific with respect to their

function; conversely, some hormones produced in one species may be completely functional in
another species.

3. Different Responses to Hormones: A hormone from one species may elicit a different
response in the same target cell or tissue of a different species.

The examples that follow illustrate these three principles and also present a comparative survey of
endocrine function in selected vertebrates. When more ancient groups of vertebrates are compared with
more recent ones, one general tendency surfaces: older groups seem to have simpler endocrine systems.
For example, many of the hormones present in mammals are absent in fishes.
 37

Hormonal Control in Fish

In fishes, the brain and spinal cord are the most important producers of hormones, with other glands
being rudimentary.

Major Hormone-Secreting Regions in Fish

1. Pineal Gland (Epithalamus):

• Produces neuropeptides affecting pigmentation and inhibiting reproductive

development, both influenced by light.

• Produces melatonin, which affects metabolism by synchronizing activity patterns with

light intensity and day length.

2. Preoptic Nuclei (Hypothalamus):

• Produces various neuropeptides controlling functions such as growth, sleep, and

movement.

3. Urophysis:

• Located in the spinal cord of the tail.

• Produces neuropeptides that help control water and ion balance, blood pressure, and
smooth muscle contractions.

• Its absence in more complex vertebrates is not well understood.

 38

Pineal Gland and Skin Color

In many fishes, amphibians, and reptiles, hormones like melatonin from the pineal gland control
variations in skin color. Injecting melatonin from one species into another can cause dramatic color
changes, indicating chemical similarity among hormones despite distant evolutionary relationships.

Prolactin

Prolactin, produced by the pituitary gland, has multiple roles across species:

• Stimulates reproductive migrations in animals like salamanders.

• Causes brooding behavior in some fishes.

• Controls water and salt balances, essential for certain saltwater fishes to enter freshwater during
spawning.

Evolution of the Thyroid Gland

Evidence suggests that the thyroid gland in the earliest vertebrates evolved from a pouchlike structure
(endostyle) in the digestive tract. This explains its position in the neck, ventral to the pharynx, in all
vertebrates.

Thyroid Hormones

1. Thyroxine (T4) and Triiodothyronine (T3):

• Regulate metabolism, growth, and tissue differentiation in vertebrates.

• In amphibians, they also control metamorphosis.

Metamorphosis in Amphibians
 39

In frogs, the concentrations of three hormones—prolactin, thyroxine, and triiodothyronine—control

metamorphosis. Low thyroxine and triiodothyronine and high prolactin levels in young tadpoles prevent
metamorphosis. As tadpoles grow, the hypothalamus releases hormones that trigger the pituitary gland
to stop producing prolactin and start producing thyroid-stimulating hormone, leading to metamorphosis.

Ultimobranchial Glands and Calcitonin

In jawed fishes and primitive tetrapods, small ultimobranchial glands form near the esophagus and
produce calcitonin, which helps regulate blood calcium levels.

Adrenal Glands and Stress Response

Specialized endocrine cells (chromaffin tissue) or glands (adrenal glands) near the kidneys prepare
vertebrates for stress. They produce epinephrine (adrenaline) and norepinephrine (noradrenaline),
which cause vasoconstriction, increased blood pressure, changes in heart rate, and increased blood
glucose levels, contributing to "fight-or-flight" reactions.

Endocrine Systems of Birds and Mammals

With some minor exceptions, birds and mammals have a similar complement of endocrine glands
(figure 25.12). Table 25.1 summarizes the major hormones these vertebrates produce.

Birds
The endocrine glands in birds include the ovary, testes, adrenals, pituitary, thyroid, pancreas,
parathyroids, pineal, hypothalamus, thymus, ultimobranchial gland, and bursa of Fabricius
(figure 25.12a). Since the hormones that most of these glands produce and their effects on target tissues
are nearly the same as in mammals, they are discussed in the next section on mammals. A discussion
of some unique hormones and their functions in birds follows.

1. Prolactin:

• Secreted by the pituitary gland in some birds (e.g., pigeons and doves).

• Stimulates the production of "pigeon's milk" by desquamation (sloughing off cells) in

the pigeon’s crop.

• Stimulates and regulates broodiness and other parental behaviors.

• Along with estrogen, stimulates full development of the brood (incubation) patch,
which helps maintain the egg temperature between 33 and 37°C.

2. Thyroxine:

• Produced by the thyroid gland.

• Regulates the normal development of feathers and the molt cycle.

• Plays a role in the onset of migratory behavior.

3. Testosterone:

• Produced by the testes in male birds.

• Controls secondary sexual characteristics such as bright plumage color, comb (if
present), and spurs.

• Strongly influences sexual behavior.

 40

4. Calcitonin:

• Secreted by the ultimobranchial glands, small, paired structures in the neck just below
the parathyroid glands.

• Involved in regulating blood calcium concentrations.

5. Bursa of Fabricius:

• A sac that lies just dorsal to the cloaca and

empties into it.

• Well developed during the bird’s

embryological development, it begins to
shrink soon after hatching.

• Produces secretions responsible for the

maturation of white blood cells (B
lymphocytes), which play an important
role in immunological reactions.

Mammals
Zoologists know more about the endocrine organs, hormones, and target tissues of mammals than of
any other animal group. This is especially true for the human body. A brief overview of mammalian
endocrinology follows.

Pituitary Gland (Hypophysis)

The pituitary gland (also known as the hypophysis) is directly below the hypothalamus. The pituitary
has two distinct lobes: the anterior lobe (adenohypophysis) and the posterior lobe
(neurohypophysis). The two lobes differ in several ways:

1. The adenohypophysis is larger than the neurohypophysis.

2. Secretory cells called pituicytes are in the adenohypophysis but not in the neurohypophysis.

3. The neurohypophysis has a greater supply of nerve endings.

Pituicytes produce and secrete hormones directly from the adenohypophysis, whereas the
neurohypophysis obtains its hormones from the neurosecretory cells in the hypothalamus, storing and
releasing them when needed. These modified hypothalamic nerve cells project their axons down a stalk
of nerve cells and blood vessels, called the infundibulum, into the pituitary gland, directly linking the
nervous and endocrine systems.

The pituitary of many vertebrates (but not in humans, birds, and cetaceans) also has a functional
intermediate lobe (pars intermedia) of mostly glandular tissue. Its secretions (e.g., melanophore-
stimulating hormone) in response to external stimuli induce changes in the coloration of the body
surface of many animals.

Hormones of the Neurohypophysis

The neurohypophysis does not manufacture any hormones. Instead, the neurosecretory cells of the
hypothalamus synthesize and secrete two hormones, antidiuretic hormone and oxytocin, which move
down nerve axons into the neurohypophysis, where they are stored in the axon terminals until released.

Antidiuretic Hormone (ADH)

 41

• Function: ADH (vasopressin) decreases urine secretion.

• Mechanism: Released when a mammal becomes dehydrated, ADH increases water absorption
in the kidneys, reducing urine secretion and retaining water.

• Homeostasis: This negative feedback system restores water and solute balance.

Oxytocin

• Function: Plays a role in reproduction by affecting smooth muscle.

• Mechanism: Stimulates uterine contractions to aid in childbirth and promotes milk ejection
from mammary glands for newborn nourishment.

Evolutionary Perspective

• Common Ancestry: ADH and oxytocin are thought to have evolved from a similar ancestral
chemical messenger that controlled water loss and solute concentrations.

• Adaptation: The neurohypophysis is larger in animals living in arid regions, where water
conservation is crucial.

• Structural Similarity: ADH and oxytocin have similar structures, differing in only two amino
acids.

Hormones of the Adenohypophysis

The adenohypophysis is the true endocrine portion of
the pituitary gland, synthesizing six different hormones.
These hormones are all polypeptides, and all but two are
true tropic hormones that target other endocrine
glands. The two non-tropic hormones are growth
hormone and prolactin.

Non-Tropic Hormones

1. Growth Hormone (GH) / Somatotropin (STH)

• Function: Affects all parts of the body concerned

with growth.

• Mechanism: Directly induces cell division necessary

for growth and protein synthesis by stimulating amino
acid uptake, RNA synthesis, and ribosome activity.

2. Prolactin (PRL)

• Function: Plays a role in many aspects of reproduction.

• Mechanism: Stimulates reproductive migrations in mammals, enhances mammary

gland development, and promotes milk production. (Oxytocin stimulates milk ejection
from the mammary glands, but not its production.)

Tropic Hormones

1. Thyrotropin / Thyroid-Stimulating Hormone (TSH)

 42

• Function: Stimulates the thyroid gland.

• Mechanism: Promotes synthesis and secretion of thyroxine, the main thyroid hormone.

2. Adrenocorticotropic Hormone (ACTH)

• Function: Stimulates the adrenal gland.

• Mechanism: Promotes production and secretion of steroid hormones called

glucocorticoids (cortisol). ACTH secretion is regulated by corticotropin-releasing
factor from the hypothalamus, which is controlled by factors such as stress, insulin,
ADH, and other hormones.

3. Gonadotropins

• Luteinizing Hormone (LH)

• Function: Stimulates the gonads.

• Mechanism: In females, LH stimulates ovulation and the secretion of estrogen

and progesterone from the corpus luteum. In males, LH targets cells in the
testes to secrete testosterone.

• Follicle-Stimulating Hormone
(FSH)

• Function: Stimulates the

gonads.

• Mechanism: In females,
FSH stimulates follicular
cells in the ovaries to
develop into mature eggs and produce estrogen. In males, FSH stimulates the
cells of the testes to produce sperm.

Pineal Gland

The pineal gland (or pineal body) is named for its pine cone shape. Its distinctive cells evolved from
the photoreceptors of
lower vertebrates and
synthesize melatonin.
These cells are most
active in the dark, as light
inhibits the enzymes
needed for melatonin
synthesis.

• Function:
Melatonin affects
many
physiological
processes and
adjusts them to
diurnal and
seasonal cycles.
 43

• Evolutionary Adaptation: Melatonin ensures that periodic activities in mammals occur when
environmental conditions are optimal.

• Role in Humans: Decreased melatonin secretion may help trigger the onset of puberty, the age
at which reproductive structures start to mature.

Thyroid Gland and Parathyroid Glands

Thyroid Gland
The thyroid gland is located in the neck, in front of the trachea (see Figure 25.12). It produces two
important hormones: thyroxine and triiodothyronine. These hormones affect overall growth,
development, and metabolism.

Another hormone from the thyroid gland is calcitonin. Calcitonin helps control the level of calcium
ions (Ca2+) in the blood. When calcium levels are high, calcitonin promotes the storage of calcium in
the bones. When calcium levels return to normal, the thyroid gland reduces calcitonin production.

Parathyroid Glands
The parathyroid glands are small, pea-sized glands located in the
thyroid lobes, usually two in each lobe (see Figure 25.12). They
produce a hormone called parathormone (PTH). PTH regulates the
levels of calcium (Ca2+) and phosphate (HPO4 2-) ions in the
blood.

When blood calcium levels are low, the parathyroid glands increase
the production of PTH. PTH has several effects:

• It stimulates bone cells to break down bone tissue and

release calcium ions into the blood.

• It enhances calcium absorption from the small intestine

into the blood.

• It promotes calcium reabsorption by the kidneys, reducing

the amount of calcium lost in urine.

Figure 25.15 illustrates the negative feedback system for

parathormone.

Adrenal Glands
In mammals, the adrenal glands are located on top of the kidneys.
Each adrenal gland is made up of two different types of tissue: the
medulla (inner portion) and the cortex (outer portion) (see Figure
25.16).
 44

Adrenal Cortex

The adrenal cortex produces three types of steroid hormones:

glucocorticoids, mineralocorticoids, and sex hormones.

Glucocorticoids

• Cortisol is the main glucocorticoid.

• Functions: Regulates overall metabolism and blood

sugar levels. It also helps the body respond to infections
and injuries.

Mineralocorticoids

• Aldosterone is the primary mineralocorticoid.

• Functions: Maintains the balance of solutes (like sodium) in the extracellular fluid. It also
promotes sodium reabsorption in the kidneys, which leads to water reabsorption. This helps
maintain the homeostasis of extracellular fluid.

Sex Hormones

• aThe adrenal cortex produces sex hormones, including androgens (weak male hormones) and
estrogens (female hormones).

• Functions: These hormones usually have only a minor effect on the gonads (male and female
reproductive organs).

Adrenal Medulla
The adrenal medulla is controlled by the nervous system. It contains neurosecretory cells that release
two important hormones: epinephrine (adrenaline) and norepinephrine (noradrenaline). These
hormones help regulate heart rate and carbohydrate metabolism.

Neural Control

• Brain Centers and Hypothalamus: The brain centers and the hypothalamus control the
adrenal medulla through sympathetic nerves.

• Functions: During times of excitement, emergency, or stress, the adrenal medulla is activated
by the sympathetic nervous system.

Fight-or-Flight Response

In response to epinephrine and norepinephrine:

• The heart rate increases.

• Blood flow increases to many vital organs.

• The airways in the lungs dilate.

• More oxygen is delivered to all cells of the body.

These changes prepare the body to react quickly and effectively to emergencies, a response known as
the "fight-or-flight" response.

Pancreas
 45

The pancreas is an elongated, fleshy organ located behind the stomach. It has both exocrine and
endocrine functions.

Exocrine Function

As an exocrine gland, the pancreas secretes digestive enzymes

through ducts.

Endocrine Function

As an endocrine gland (ductless), the pancreas performs the

following functions:

• The endocrine portion constitutes about 1% of the gland.

• It consists of clusters of cells called pancreatic islets.

• There are 200,000 to 2,000,000 pancreatic islets scattered throughout the gland.

Pancreatic Islets

Each pancreatic islet contains four types of specialized cells:

1. Alpha (α) cells:

• Produce the hormone glucagon.

2. Beta (β) cells:

• Produce insulin.

3. Delta (δ) cells:

• Secrete somatostatin. Somatostatin

inhibits the secretion of glucagon and
insulin.

4. F cells:

• Secrete pancreatic polypeptide. This

polypeptide inhibits somatostatin
secretion, gallbladder contraction, and the
secretion of pancreatic digestive enzymes
after a meal.

Regulation of Blood Glucose

Insulin

• When blood glucose levels are high (e.g., after a meal), beta cells secrete insulin.

• Insulin promotes the uptake of glucose by body cells, including liver cells, where excess
glucose is converted to glycogen (a storage form of glucose).

Glucagon

• When blood glucose levels are low, alpha cells secrete glucagon.

• Glucagon stimulates the breakdown of glycogen into glucose, which is then released into the
bloodstream to raise blood glucose levels to the homeostatic range.
 46

Insulin and glucagon are crucial for regulating blood glucose concentrations through a negative
feedback system. This system maintains appropriate blood glucose levels, ensuring the body's energy
needs are met efficiently.

Gonads
The gonads (ovaries and testes) secrete hormones that regulate reproductive functions.

Male Gonads (Testes)

• Testosterone:

• Works with luteinizing hormone (LH) and follicle-stimulating hormone (FSH)

from the adenohypophysis to stimulate spermatogenesis.

• Necessary for the growth and maintenance of the male sex organs.

• Promotes the development and maintenance of sexual behavior.

• Stimulates the growth of facial and pubic hair and the enlargement of the larynx,
which deepens the voice.

• Inhibin:

• Inhibits the secretion of FSH.

Female Gonads (Ovaries)

• Estrogens (estrin, estrone, estradiol):

• Regulate the menstrual and estrus cycles.

• Aid in the development of the mammary glands and other female secondary sexual
characteristics.

• Progestins (primarily progesterone):

• Regulate the menstrual and estrus cycles.

 47

• Aid in the development of the mammary glands and placenta formation during
pregnancy.

• Relaxin:

• Softens the cervix at the time of delivery.

• Inhibin:

• Inhibits the secretion of FSH.

Thymus
The thymus gland is located near the heart. It is large and noticeable in young birds and mammals
but shrinks in size throughout adulthood.

• Major hormonal products:

• Thymopoietin (TP)

• Alpha1 thymosin

• Beta4 thymosin

• These hormones are essential for the normal development of the immune system.
 48

Circulation and Gas Exchange:

Internal Transport and Circulatory Systems

Introduction
All animals need to maintain a stable internal environment, or homeostasis. This involves the circulation
of nutrients, wastes, and gases throughout the body. Internal transport or circulatory systems reduce the
distance that these substances must travel by diffusion. The complexity and nature of these systems
vary depending on the size and lifestyle of the animal.

Transport Systems in Invertebrates

Protozoa

• Diffusion: Protozoa rely on simple diffusion due to their small size and high surface-area-to-
volume ratio.

• Medium: Materials diffuse through the plasma membrane and cytoplasm.

Sponges

• Water Circulation: Sponges circulate external water through their bodies instead of using an
internal fluid system.

Cnidarians

• Gastrovascular Cavity: Hydra and similar cnidarians use a fluid-filled gastrovascular cavity
to supply nutrients and oxygen to cells and remove wastes.

Flatworms

• Complex Gastrovascular Cavity: Planarians have a branched gastrovascular cavity that

shortens diffusion distances for nutrients, gases, and wastes. Body movements help distribute
materials.

Pseudocoelomate Invertebrates

• Coelomic Fluid: Rotifers, gastrotrichs,

and nematodes use the fluid in their
body cavity for transport. Body
movements ensure adequate
distribution.

Molluscs and Beyond

 49

• Separate
Circulatory
Systems: Beginning
with molluscs, many
invertebrates have
developed separate
circulatory systems.

• Types of Circulatory Systems:

• Open Circulatory System: Hemolymph is pumped by the heart into the body cavity,
where it bathes cells and tissues directly.

• Closed Circulatory System: Blood circulates within vessels, delivering nutrients and
removing wastes.

Examples

• Annelids (e.g.,
Earthworms): Have a
closed circulatory system
with blood confined to
vessels.

• Molluscs and
Arthropods: Most have
open circulatory systems
where hemolymph
bathes cells and tissues
directly. For example, insects have hearts that pump hemolymph through vessels that open into
a body cavity (hemocoel).

In summary, invertebrates use a variety of transport systems ranging from simple diffusion to complex
circulatory systems, depending on their size and complexity. These systems ensure that nutrients, gases,
and wastes are efficiently circulated to maintain homeostasis.

Characteristics of Invertebrate Coelomic Fluid, Hemolymph, and

Blood Cells
Coelomic Fluid
• Usage: Some invertebrates like echinoderms, annelids, and sipunculans use coelomic fluid for
transport within the body.

• Composition: Can be similar to interstitial fluids or may differ, especially in proteins and cells.

• Functions:

• Transports gases, nutrients, and wastes.

• Can act as a hydrostatic skeleton in certain invertebrates, such as annelids.

Hemolymph
 50

• Definition: Hemolymph is the circulating fluid in animals with open circulatory systems,
including most arthropods, ascidians, and many molluscs.

• Circulation:

• Pumped by the heart at low pressures through vessels into tissue spaces (hemocoel) and
sinuses.

• Circulation is generally slow, and hemolymph volume is high.

• Functions:

• Transports essential gases, nutrients, and wastes.

• Noncirculatory roles in some invertebrates:

• Insects: Hemolymph pressure assists in molting and wing inflation.

• Jumping Spiders: Provides a hydraulic mechanism for limb extension.

Blood Cells (Hemocytes)

• Presence: Found in coelomic fluid, hemolymph, or blood of most invertebrates.

• Types and Functions:

• Erythrocytes (Red Blood Cells): Contain respiratory pigments like hemoglobin to

facilitate oxygen transport.

• Non-pigmented Cells: Involved in various functions, such as blood clotting.

Examples by Invertebrate Group

1. Annelids:

• Blood contains phagocytic hemocytes.

• Coelomic fluid has coelomocytes (amoebocytes, eleocytes, lampocytes, linocytes)

involved in:

• Phagocytosis

• Glycogen storage

• Encapsulation

• Defense responses

• Excretion

2. Molluscs:

• Hemolymph contains two main types of hemocytes:

• Amoebocytes: Involved in immune responses and nacrezation (pearl

formation in some bivalves).

• Granulocytes: Perform various functions including phagocytosis.

3. Insects:
 51

• Hemolymph contains a variety of hemocyte types that function in:

• Phagocytosis

• Encapsulation

• Clotting

In summary, the internal transport systems of invertebrates feature a variety of fluids and cells adapted
to their specific needs, ranging from basic diffusion in simple organisms to more complex systems in
larger or more active animals. These systems ensure efficient transport of essential substances and
contribute to other physiological functions, aiding in the overall maintenance of homeostasis.

Transport Systems in Vertebrates

All vertebrates have a closed circulatory system in which the walls of the heart and blood vessels are
always contracted, and blood never leaves the blood vessels (figure 26.1g). Blood moves from the heart,
through arteries, smaller arteries (arterioles), tiny blood vessels (capillaries), small veins (venules),
veins, and back to the heart. Exchange between the blood and the fluid outside the cells only occurs at
the capillary level.

Characteristics of Vertebrate Blood and Blood Cells

Overall, vertebrate blood transports oxygen, carbon dioxide, and nutrients; defends against harmful
microorganisms, cells, and viruses; prevents blood loss through clotting; and helps regulate body
temperature and pH. Because it is a liquid, vertebrate blood is classified as a special type of connective
tissue. Like other connective tissues, blood contains a fluid matrix called plasma and cellular parts called
formed elements.

Plasma

Plasma (Gr., anything formed or molded) is the straw-colored, liquid part of blood. In mammals, plasma
is about 90% water and provides the liquid for dissolving and transporting nutrients. A group of proteins
(albumin, fibrinogen, and globulins) makes up another 7% of the plasma. The amount of these plasma
proteins affects the movement of water between the blood and the fluid outside the cells. Because
albumin makes up about 60% of the total plasma proteins, it plays important roles with respect to water
movement. Fibrinogen is needed for blood clotting, and the globulins include the immunoglobulins and
various metal-binding proteins. Serum is plasma from which the proteins involved in blood clotting
have been removed. The gamma globulin part functions in the immune response because it consists
mostly of antibodies. The remaining 3% of plasma is made up of electrolytes, amino acids, glucose and
other nutrients, various enzymes, hormones, metabolic wastes, and traces of many inorganic and organic
molecules.
 52

Formed Elements

The formed-element part (cellular

component) of vertebrate blood
consists of red blood cells
(erythrocytes), white blood cells
(leukocytes), and platelets
(thrombocytes) (figure 26.3). White
blood cells are present in lower
numbers than red blood cells,
generally being 1 to 2% of the blood
by volume. White blood cells are
divided into agranulocytes (without
granules in the cytoplasm) and
granulocytes (have granules in the
cytoplasm). The two types of
agranulocytes are lymphocytes and
monocytes. The three types of
granulocytes are eosinophils,
basophils, and neutrophils.
Fragmented cells are called platelets
(thrombocytes). Each of these cell
types is now discussed in more
detail.

Red Blood Cells

Red blood cells (erythrocytes; Gr.

erythros, red cells) vary greatly in
size, shape, and number in different
vertebrates (figures 26.4 and 26.5a).
For example, the RBCs of most
vertebrates are nucleated (have a
nucleus), but mammalian RBCs are enucleated (without a nucleus). Some fishes and amphibians also
have enucleated RBCs. Among all vertebrates, the salamander Amphiuma has the largest RBC (figure
26.4a). Bird RBCs (figure 26.4c) are oval-shaped, nucleated, and larger than mammalian RBCs.
Among birds, the ostrich has the largest RBC. Most mammalian RBCs are biconcave disks (figure
26.5a); however, the camel (figure 26.4e) and llama have elliptical RBCs. The shape of a biconcave
disk provides a larger surface area for gas diffusion than a flat disk or sphere. Generally, the lower
vertebrates tend to have fewer but larger RBCs than the higher vertebrates.

Almost the entire mass of an RBC consists of hemoglobin (Gr. haima, blood L. globulus, little globe),
an iron-containing protein. The major function of a red blood cell is to pick up oxygen from the
environment, bind it to hemoglobin to form oxyhemoglobin, and transport it to body tissues. Blood rich
in oxyhemoglobin is bright red. As oxygen diffuses into the tissues, blood becomes darker and appears
blue when observed through the blood vessel walls. However, when this less oxygenated blood is
exposed to oxygen (such as when a vein is cut and a mammal begins to bleed), it immediately turns
bright red. Hemoglobin also carries waste carbon dioxide (in the form of carbaminohemoglobin) from
the tissues to the lungs (or gills) for removal from the body.

White Blood Cells

White blood cells (leukocytes) (Gr. leukos, white cells) are scavengers that destroy microorganisms at
infection sites, remove foreign chemicals, and clear debris resulting from dead or injured cells. All
 53

WBCs are derived from immature cells (stem cells) in bone marrow through a process called
hematopoiesis (Gr. hemato, blood poiein, to make; see figure 26.3).

Granulocytes

Among the granulocytes, eosinophils are phagocytic, ingesting foreign proteins and immune complexes
rather than bacteria (figure 26.5b). In mammals, eosinophils also release chemicals that counteract the
effects of certain inflammatory chemicals during allergic reactions. Basophils are the least numerous
WBCs (figure 26.5c). When they encounter a foreign substance, their granules release histamine and
heparin. Histamine causes blood vessels to widen and leak fluid at sites of inflammation, while heparin
prevents blood clotting. Neutrophils are the most abundant of the white blood cells (figure 26.5d). They
are chemically drawn to sites of inflammation and are active phagocytes.

Agranulocytes

The two types of agranulocytes are monocytes and lymphocytes (figure 26.5e,f). Two distinct types of
lymphocytes are B cells and T cells, both crucial to the immune response. B cells originate in the bone
marrow and mature in lymphoid tissue, while T cells are associated with the thymus gland before
colonizing lymphoid tissue and contributing to the immune response. When B cells are activated, they
divide and differentiate to produce plasma cells.

Platelets (Thrombocytes)

Platelets (named for their flat, plate-like shape), or thrombocytes (Gr. thrombus, clot cells), are disk-
shaped cell fragments that initiate blood clotting. When a blood vessel is injured, platelets quickly move
to the site and clump, adhering to the damaged area and initiating the process of blood coagulation.

Vertebrate Blood Vessels

Arteries are elastic blood vessels that carry blood
away from the heart to the organs and tissues of the
body. The central canal of an artery (and all blood
vessels) is called a lumen. Surrounding the lumen of
an artery is a thick wall composed of three layers,
called tunics (L. tunica, covering) (figure 26.6a).

Most veins are relatively rigid, large vessels that

carry blood from the body tissues back to the heart.
The wall of a vein also contains the same three layers
(tunics) as arterial walls, but the middle layer is much
thinner, and one or more valves are present (figure 26.6b). Valves permit blood flow in only one
direction, which is crucial for returning the blood to the heart.

Arteries lead to terminal arterioles (those closest to a capillary). The arterioles then branch to form
capillaries (L. capillus, hair), which connect to venules and then to veins. Capillaries are generally
made up of a single layer of endothelial cells and are the most numerous blood vessels in an animal’s
body (figure 26.6c). An abundance of capillaries provides an enormous surface area for the exchange
of gases, fluids, nutrients, and wastes between the blood and nearby cells.

The Hearts and Circulatory Systems of Bony Fishes, Amphibians,

and Reptiles
The heart and blood vessels underwent significant changes as vertebrates transitioned from water to
land and as endothermy evolved. Let's delve into these transformations with examples from bony fishes,
amphibians, and reptiles.
 54

Bony Fishes:

The bony fish heart has two chambers—the atrium and ventricle (figure 26.7a). Blood exits the heart
through the ventral aorta, which leads to the gills. In the gills, blood becomes oxygenated, releases
carbon dioxide, and then enters the dorsal aorta. The dorsal aorta distributes blood to all body organs,
and blood eventually returns to the heart through the venous system. This system, called a single
circulation circuit, has the advantage of circulating oxygenated blood from the gills to the systemic
capillaries in all organs almost simultaneously. However, the circulation of blood through the gill
capillaries creates resistance to flow, leading to reduced blood pressure and flow rates to other organs.
This arrangement may not support the high metabolic rates present in some birds and mammals.

Amphibians:

In amphibians, a
double circulatory
circuit evolved,
where blood passes
through the heart
twice during its
circuit through the
body. Amphibians
typically have hearts
that are not fully
divided into two
chambers. A single
ventricle pumps
blood to both the
lungs and the rest of
the body (figure
26.7b). Since most
amphibians absorb
more oxygen through
their skin than
through their lungs or
gills, oxygenated blood returning from the skin also contributes to the ventricle, resulting in highly
oxygenated blood pumped out to the rest of the body.

Reptiles:

In most reptiles, such as turtles, the ventricle is partially divided into right and left sides (figure 26.7c).
Oxygenated blood from the lungs returns to the left side of the heart via the pulmonary vein and does
not mix much with deoxygenated blood in the right side of the heart. When the ventricles contract,
blood is pumped out to two aortae for distribution throughout the body and to the lungs. This incomplete
separation of the ventricles is an essential adaptation for reptiles, allowing blood to be diverted away
from the pulmonary circulation during diving and when the turtle withdraws into its shell. This
conserves energy and redirects blood to vital organs when the lungs cannot be ventilated.

The Hearts and Circulatory Systems of Birds and Mammals

While the physiological separation of blood in left and right ventricles is nearly complete in reptiles,
the complete anatomical separation of ventricles only occurs in crocodilians, birds, and mammals
(figure 26.7d). This enables the double circulation required to maintain high blood pressure. High
blood pressure is crucial for the rapid delivery of oxygenated, nutrient-rich blood to tissues with high
metabolic rates.
 55

Blood circulates throughout the avian and mammalian body

in two main circuits: the pulmonary circuit and the
systemic circuit (figure 26.8).

• The pulmonary circuit supplies blood only to the

lungs. It carries oxygen-poor (deoxygenated)
blood from the heart to the lungs, where carbon
dioxide is removed, and oxygen is added. Then, it
returns the oxygen-rich (oxygenated) blood to the
heart for distribution to the rest of the body.

• The systemic circuit supplies all cells, tissues, and

organs of the body with oxygen-rich blood and
returns oxygen-poor blood to the heart.

THE HUMAN HEART: A Powerful

Pump
Pumping Capacity

• Volume and Rate: Moves approximately 5 liters of blood per minute; handles around 8,000
liters daily through 96,000 km of blood vessels

• Frequency: Beats about 70 times per minute in an average adult, exceeding 100,000 times per
day

• Lifetime Activity: Over 2.6 billion beats in a typical 70-year lifespan without tiring

Heart Structure

• Myocardium: Dominant tissue type, consisting

of cardiac muscle

• Epicardium: Outer layer providing protective

fibrous connective tissue

• Endocardium: Inner lining, formed by

connective tissue and endothelium

Chamber Configuration and Valves

• Atria and Ventricles: Heart divided into left

and right halves, each with an atrium and
ventricle

• Valves: Ensure one-way blood flow; including tricuspid, bicuspid, pulmonary, and aortic
valves

Cardiac Cycle

• Initiation: Sinoatrial node (SA node) initiates each heartbeat

 56

• Propagation: Electrical signals travel through atrioventricular node (AV node) and bundle of
His

• Contraction: Ventricles
contract due to stimulation
by Purkinje fibers

Electrical Activity and Monitoring

• Pacemaker Function: SA
node generates electrical
impulse, initiating cardiac
cycle

• Recording:
Electrocardiogram (ECG or
EKG) monitors heart's
electrical activity

Systole and Diastole

• Contraction and Relaxation: Atria and ventricles undergo rhythmic cycles of contraction
(systole) and relaxation (diastole), facilitating blood movement

Blood Pressure Overview

Blood pressure is the fluid pressure
generated by ventricular contraction that
forces blood through the pulmonary and
systemic circuits. It is the force that blood
exerts against the inner walls of blood
vessels and is a crucial measurement for
assessing cardiovascular health.

Systemic Arterial Blood Pressure

Although blood pressure occurs throughout

the vascular system, the term typically
refers to systemic arterial blood pressure.
This pressure rises and falls in a pattern that
corresponds to the phases of the cardiac
cycle:

1. Ventricular Systole
(Contraction):

During this phase, the ventricles contract,

forcing blood into the pulmonary arteries
and the aorta. The pressure in these arteries
rises sharply. The highest pressure achieved
during this phase is known as the systolic
pressure.

2. Ventricular Diastole
(Relaxation):
 57

During this phase, the ventricles relax. The arterial pressure drops. The lowest pressure in the arteries
before the next ventricular contraction is known as the diastolic pressure.

Normal Blood Pressure Values

Systolic Pressure: The normal systolic pressure for a young adult is about 120 mm Hg. This is the
amount of pressure required to raise a column of mercury in a sphygmomanometer (sfig-mo-mah-no-
meter) to 120 mm.

Diastolic Pressure: The normal diastolic pressure is approximately 80 mm Hg.

These readings are conventionally expressed as 120/80 mm Hg.

Importance of Blood Pressure Monitoring

Monitoring blood pressure is essential for detecting and managing conditions such as hypertension (high
blood pressure) and hypotension (low blood pressure), both of which can lead to serious health
complications if left unaddressed. Regular check-ups can help maintain optimal cardiovascular health
and prevent long-term issues.

THE LYMPHATIC SYSTEM

The lymphatic system in vertebrates begins with lymphatic capillaries, which interact directly with the
extracellular fluid surrounding tissues. This system performs four primary functions:

1. Fluid Collection and Drainage: It collects and drains most of the fluid that seeps out of the
bloodstream and accumulates in the extracellular fluid.

2. Protein Return: It returns small amounts of proteins that have exited the cells.

3. Lipid Transport: It transports lipids absorbed from the small intestine.

4. Transport of Foreign Particles and Debris: It moves foreign particles and cellular debris to
lymph nodes, which are disposal centers.

Lymphatic capillaries merge to form larger vessels known as lymphatics, which are thin-walled and
equipped with valves to ensure the one-way flow of lymph. Lymph, the clear extracellular fluid in these
vessels, travels through lymph nodes on its way back to the heart. Lymph nodes are concentrated in
various body areas and play a crucial role in defending against disease.

In birds and mammals, the lymphatic system also includes lymphoid organs such as the spleen and,
depending on the species, either the bursa of Fabricius in birds or the thymus gland, tonsils, and adenoids
in mammals. This system is essential for an animal's defense against injury and infection.

Gas Exchange
To take advantage of the rich source of energy in earth’s organic matter, animals must solve two
practical problems. First, they must break down and digest the organic matter so that it can enter the
cells that metabolize it. Second, they must provide cells with both an adequate supply of oxygen
required for aerobic respiration and a way of eliminating the carbon dioxide that aerobic respiration
produces. This process of gas exchange with the environment, also called external respiration, is the
subject of the rest of this chapter.

Respiratory Surfaces
Protists and animals have five main types of respiratory systems (surfaces):
 58

1. Simple Diffusion across Plasma Membranes

2. Tracheae

3. Cutaneous (Skin) Exchange

4. Gills

5. Lungs

Invertebrate Respiratory Systems

• Single-celled Protists: In single-celled protists, such as protozoa, diffusion across the plasma
membrane moves gases into and out of the organism.

• Multicellular Invertebrates: Some multicellular invertebrates either have very flat bodies
(e.g., flatworms) in which all body cells are close to the body surface or are thin-walled and
hollow (e.g., Hydra). Gases diffuse into and out of these animals.

• Integumentary Exchange: Invertebrates such as earthworms that live in moist environments

use integumentary exchange. Earthworms have capillary networks just under their skin and
exchange gases with the air spaces among soil particles.

• Gills: Most aquatic invertebrates carry out gas exchange with gills. Sea stars have simple gills
that are small skin projections. Marine and annelid worms have prominent lateral projections
called parapodia that function as gills. Crustaceans and molluscs have compact gills protected
by hard coverings, divided into highly branched structures to maximize the area for gas
exchange.

• Tracheal Systems: Some terrestrial invertebrates (e.g., insects, centipedes, and some mites,
ticks, and spiders) have tracheal systems consisting of highly branched tubes called tracheae.
Tracheae open to the outside through spiracles, leading to branching tracheal trunks that give
rise to smaller branches called tracheoles. Most insects have ventilating mechanisms that move
air into and out of the trachea. Arachnids possess tracheae, book lungs, or both.

• Book Lungs: Book lungs are paired invaginations of the ventral body wall folded into a series
of leaflike lamellae. Air enters the book lung through a slitlike opening called a spiracle and
circulates between lamellae, with gases diffusing between the hemolymph and the air.

• Pulmonate Lungs: Land snails and slugs, part of the molluscan subclass Pulmonate, have a
lung that opens to the outside via a pore called a pneumostome. This lung is derived from the
mantle cavity common to molluscs. Primitive pulmonate snails are aquatic and close the
pneumostome during submergence, while most higher pulmonates are terrestrial and rely on
their lungs for gas exchange.

Vertebrate Respiratory Systems

Aquatic vertebrates (fish, amphibians, and some reptiles) rely on one or a combination of the following
surfaces for gas exchange: the cutaneous body surface, external filamentous gills, and internal
lamellar gills.

Bimodal Breathing is the ability of an organism to exchange respiratory gases simultaneously with
both air and water. A bimodal organism (e.g., some salamanders, crabs, barnacles, bivalve molluscs,
and fishes [lungfishes]) uses gills for water breathing and lungs for air breathing. However, some gas
exchange is always cutaneous, and some bimodal breathers are actually trimodal (skin, gills, and lungs).
 59

Bimodal breathing was an important respiratory adaptation that made possible the evolutionary
transition between aquatic and terrestrial habitats. Fundamental changes in the structure and function
of the respiratory organs accompanied the transition from water to air breathing.

In air-breathing terrestrial vertebrates (reptiles, birds, and mammals), lungs replaced gills. These
vertebrate surfaces and transitions are now discussed.

Cutaneous Exchange
Some vertebrates with lungs or gills, such as some aquatic turtles, salamanders with lungs, snakes,
fishes, and mammals, use cutaneous respiration or skin-based exchange to help with gas exchange.
However, cutaneous exchange is most developed in frogs, toads, lungless salamanders, and newts.

Amphibian skin has the simplest structure among the major vertebrate respiratory organs. In frogs, a
uniform network of tiny blood vessels lies just below the skin. This arrangement helps gas exchange
between the blood vessels and the environment through both diffusion and movement of air. A slimy
mucus layer keeps amphibian skin moist and protects it from injury, aiding in gas exchange. Some
amphibians get about 25% or more of their oxygen through this method, and lungless salamanders
perform all of their gas exchange through the skin and the mouth-throat area.

Gills
Gills are respiratory organs with a thin, moist, vascularized layer of skin to allow gas exchange across
thin gill membranes or a very thin skin layer over highly vascularized tissue. Larval forms of some fish
and amphibians have external gills that project
from their bodies. Adult fish have internal
gills, where gas exchange is very efficient.

Gas Exchange in Gills occurs as blood and

water move in opposite directions on either side
of the gill membranes. For example, water
passing over a gill first encounters blood
vessels carrying low-oxygen blood into the
body. Because the oxygen concentration is
lower in the blood than in the water, oxygen
diffuses into the blood. As water continues to
pass over vessels carrying higher-oxygen blood
from within the body, more oxygen diffuses
inward since the blood still has less oxygen
than the water. Carbon dioxide also diffuses
into the water because its concentration is
higher in the blood than in the water.

This countercurrent exchange mechanism

provides efficient gas exchange by maintaining
a concentration gradient between the blood and
water throughout the capillary bed.

Lungs
A lung is an internal, sac-shaped respiratory organ. In terrestrial vertebrates, the lung typically consists
of one or more internal blind pouches into which air is either drawn or forced. The respiratory lining of
the lungs is thin, well-supplied with blood vessels, and divided into many small units, significantly
increasing the surface area for gas exchange between the lung air and the blood. However, this design
 60

limits the efficiency of oxygen and carbon dioxide exchange with the atmosphere because only part of
the lung air is replaced with each breath.

Bird Lungs are an exception, as they have a highly efficient one-way airflow system. For example, a
mammal extracts about 25% of the oxygen from the air with each breath, while a bird extracts about
90%.

Evolution of Lungs and Swim Bladders: The evolution of vertebrate lungs is related to the evolution
of the swim bladder. The swim bladder is an air sac located above the digestive tract in many modern
fish. Evidence suggests that both lungs and swim bladders evolved from a lung-like structure present in
primitive fish, ancestors of both modern fish and tetrapods (amphibians, reptiles, birds, and mammals).
These ancestral fish likely had a ventral sac attached to their throat that served as a supplementary gas
exchange organ when gills couldn't provide enough oxygen, such as in stagnant or low-oxygen water.
By swimming to the surface and gulping air into this sac, the fish could exchange gases through its wall.

Evolutionary Paths: The further evolution of this sac followed two paths. In most modern bony fish,
the swim bladder lies above the digestive tract. In contrast, the lungs are located below the digestive
tract. A few present-day fish and all tetrapods have ventral lungs.

Adaptations in Lung Structure: The development of more complex lungs paralleled the evolution of
larger body sizes and higher metabolic rates in warm-blooded vertebrates (birds and mammals). These
adaptations required an increased lung surface area for gas exchange compared to the smaller body size
and lower metabolic rates of cold-blooded vertebrates.

Lung Ventilation
Ventilation is based on several basic principles for all air-breathing animals with lungs:

1. Air moves by bulk flow into and out of the lungs.

2. Oxygen and carbon dioxide diffuse across the respiratory surface of the lung tissue from
pulmonary capillaries.

3. At systemic capillaries, oxygen and carbon dioxide diffuse between the blood and interstitial
fluid in response to concentration gradients.

4. Oxygen and carbon dioxide diffuse between the interstitial fluid and body cells.

Vertebrates exhibit two different mechanisms for lung ventilation based on these principles:

• Positive Pressure Pumping: Amphibians and some reptiles push air into their lungs. The
muscles of the mouth and throat create a positive pressure to force air into the lungs.
 61

• Negative Pressure System: Most reptiles and all birds and mammals inhale by suction. They
expand the body cavity to decrease pressure in the lungs, drawing air in. Elastic recoil of the
lungs and movement of the ribs and body wall expel air.

In reptiles like snakes, lizards, and crocodiles, the body cavity expands with rib movement to ventilate
the lungs. Turtles, with their ribs as part of the shell,
exhale by contracting muscles that force the internal
organs upward, compressing the lungs. They inhale
by contracting muscles that increase the volume of
the visceral cavity, creating negative pressure to
draw air in.

Birds have the highest rate of oxygen consumption

due to their high metabolic rate from flying. They
use a negative pressure system for breathing, similar
to mammals, but have a unique lung ventilation
mechanism for one-way airflow over gas exchange
surfaces. This makes bird lungs more efficient than
mammal lungs. Birds have tunnel-like passages
called parabronchi leading to air capillaries for gas
exchange. A system of air sacs throughout the body
cavity makes one-way airflow possible. Inhaled air
bypasses the lungs and enters the posterior air sacs,
passes through the lungs to the anterior air sacs,
and is then exhaled. This process requires two
complete breathing cycles.

Human Respiratory System

The structure and function of external respiration in humans are typical of mammals. Thus, the human
respiratory system is used here to describe principles that apply to all air-breathing mammals.

Air-Conducting Portion
Nasal and Oral Cavities

• Air Entry and Exit: Air normally enters and leaves through either the nasal or oral cavities.

Pharynx

• Common Area: The pharynx is a common area for both the respiratory and digestive tracts.

• Connections: The pharynx connects to the larynx (voice box) and the esophagus (leading to
the stomach).

Epiglottis

• Function: A flap of cartilage that directs air into the trachea during breathing and covers the
trachea during swallowing to prevent food or water from entering.
 62

Trachea

• Windpipe: Air moves from the larynx

into the trachea.

Bronchi

• Branches: The trachea branches into a

right and left bronchus (plural: bronchi),
each entering a lung.

Bronchioles

• Smaller Tubes: Each bronchus branches

into smaller tubes called bronchioles.

• Terminal Bronchioles: These branch

further into even smaller tubes.

• Respiratory Bronchioles: Part of the

gas-exchange portion of the respiratory
system.

Gas-Exchange Portion
Alveolar Ducts

• Connection: Small tubes called alveolar ducts

connect the respiratory bronchioles to
grapelike outpouchings called alveoli.

Alveoli

• Structure: Alveoli cluster to form an alveolar sac surrounded by many capillaries.

• Function: Alveoli are the functional units of the lungs for gas exchange.

• Gas Exchange: Passive diffusion moves oxygen from the alveoli into the blood and carbon
dioxide from the blood into the alveoli.

• Surface Area: The alveoli provide a large surface area for gas exchange. If spread out, the
alveolar epithelium would cover the area of a tennis court.

Ventilation
Breathing (Pulmonary Ventilation)

• Phases: Breathing has two phases: inhalation (intake of air) and exhalation (outflow of air).

• Mechanism: Air movements result from changes in thoracic cavity volume, which lead to
pressure gradient reversals between the lungs and the atmosphere.

Inhalation
 63

1. Muscle Contraction: Several muscle sets,

mainly the diaphragm and intercostal muscles,
contract. The intercostal muscles pull the ribs
closer together, enlarging the thoracic cavity.

2. Diaphragm Action: The diaphragm contracts

and flattens, further enlarging the thoracic
cavity.

3. Pressure Drop: The increased size of the

thoracic cavity causes pressure to drop below
atmospheric pressure, allowing air to rush into
the lungs and inflate them.

Exhalation

1. Muscle Relaxation: The intercostal muscles

and the diaphragm relax, returning the thoracic
cavity to its original size and increasing the
pressure.

2. Abdominal Muscles: These muscles contract,

pushing the abdominal organs against the
diaphragm, increasing thoracic pressure.

3. Lung Compression: This action causes the

elastic lungs to contract and compress the air in
the alveoli. The increased alveolar pressure
expels air from the lungs.

Gas Transport
As noted in the previous discussion, oxygen must be transported from the sites of environmental gas
exchange to the cells of an animal’s body. Various systems (e.g., tracheae, cutaneous exchange, gills,
lungs) help accomplish this transport.

Limitations of Simple Diffusion

As animals became larger and acquired higher metabolic rates, simple diffusion became increasingly
inadequate as a means of delivering oxygen to the tissues.

Specialized Circulatory Systems

Consequently, in most animals with high metabolic rates and tissues more than a few millimeters from
respiratory surfaces, a specialized circulatory system circulates body fluids to aid in the internal
distribution of oxygen (see figure 26.1).

Increased Oxygen Demand in Active Animals

In general, more active animals have an increased demand for oxygen.

Challenges of Oxygen Solubility

However, simply creating a convection of a water-based body fluid does not in itself guarantee internal
transport of sufficient oxygen to meet this increased demand. The reason is the low solubility of oxygen
in water-based body fluids.
 64

Evolution of Respiratory Pigments

Thus, fluid-borne respiratory pigments specialized for reversibly binding large quantities of oxygen
evolved in most phyla.

Functions of Respiratory Pigments

Respiratory pigments help the various transport systems satisfy this increased oxygen demand. In
addition to oxygen transport, respiratory pigments may also function in short-term oxygen storage.

Respiratory Pigments
Respiratory pigments are organic compounds containing either metallic copper or iron that binds
oxygen. These pigments may be dissolved in the blood or body fluids, or they may be contained within
specific blood cells. Generally, pigments combine with oxygen in high oxygen concentrations and
release oxygen in low oxygen concentrations.

Types of Respiratory Pigments

1. Hemoglobin

• Metal: Iron

• Color: Reddish

• Distribution: Common in a variety of invertebrates (e.g., protozoa, platyhelminths,

nemerteans, nematodes, annelids, crustaceans, some insects, and molluscs) and almost
all vertebrates except a few fishes.

• Location: Carried within red blood cells (erythrocytes) or dissolved in blood or

coelomic fluid.

2. Hemocyanin

• Metal: Copper

• Color: Bluish when oxygenated

• Distribution: Most common in molluscs and certain crustaceans.

• Location: Always dissolved in hemolymph.

• Characteristic: Releases oxygen easily, providing a ready source of oxygen to tissues

as long as environmental oxygen concentrations are relatively high.

3. Hemerythrin

• Metal: Iron

• Color: Pink when oxygenated

• Distribution: Found in sipunculans, priapulids, a few brachiopods, and some

polychaetes.

• Location: In nucleated cells, rather than free in body fluids or hemolymph.

4. Chlorocruorin

• Metal: Iron
 65

• Color: Green in low oxygen concentrations, bright red in high oxygen concentrations

• Distribution: Found in several families of polychaete worms.

Function of Respiratory Pigments

Respiratory pigments significantly increase the oxygen-carrying capacity of body fluids beyond
what would be achieved by simple transport in a dissolved state.

Carbon Dioxide Transport

Carbon dioxide (CO2) concentrations in animal body fluids are higher than expected based on solubility
due to transport mechanisms:

• Bound to hemoglobin

• Dissolved state

• Carbonic acid (H2CO3) and bicarbonate ion (HCO3–) in reversible reactions:

• CO2 + H2O ⇌ H2CO3 ⇌ H+ + HCO3–

This "tying up" of carbon dioxide in other forms lowers its concentration in solution, thus increasing
the overall carrying capacity of a body fluid like blood.

Evolutionary Perspective

The occurrence of respiratory pigments among various taxa lacks a clear phylogenetic explanation.
Their sporadic distribution suggests that some pigments may have evolved multiple times through
parallel evolution. Notably, respiratory pigments are rare among successful insects, reflecting their use
of extensive tracheal systems to carry gases directly to the tissues. Insects without well-developed
tracheae carry oxygen in solution in the hemolymph.
 66

Nutrition and Digestion:

Animal Nutrition
Nutrition includes all processes by which an animal takes in, digests, absorbs, stores, and uses food
(nutrients) to meet its metabolic needs.

Digestion

Digestion (L. digestio, from dis, apart + gerere, to carry) is the chemical and/or mechanical
breakdown of food into particles that individual cells of an animal can absorb.

Overview of Animal Nutrition

This chapter discusses:

• Animal nutrition
• Different strategies animals use for consuming and using food
• Various animal digestive systems

Evolution of Nutrition
Nutrients in the food an animal eats provide the necessary chemicals for growth, maintenance, and
energy production. Overall, the nutritional requirements of an animal are inversely related to its ability
to make essential molecules for life. The fewer such biosynthetic abilities an animal has, the more
kinds of nutrients it must get from its environment.

Autotrophs and Heterotrophs

Green plants and photosynthetic protists have the fewest nutritional requirements because they can
make all their own complex molecules from simpler inorganic substances; they are autotrophs (Gr.
auto, self + trophe, nourishing).

Animals, fungi, and bacteria that cannot make many of their own organic molecules and must get them
by consuming other organisms or their products are heterotrophs (Gr. heteros, another or different +
trophe, nourishing).

Types of Diets in Animals

Animals such as rabbits that live entirely on plant material are herbivores (L. herba, plant + vorare, to
eat).

Carnivores (L. caro, flesh), such as hawks, are animals that eat only meat.

Omnivores (L. omnius, all), such as humans, bears, raccoons, and pigs, eat both plant and animal
matter.

Insectivores, such as bats, eat mainly arthropods.

Evolutionary Loss of Biosynthetic Abilities

Losses of biosynthetic abilities have marked much of animal evolution. Once an animal regularly gets
essential, complex organic molecules in its diet, it can afford to lose the ability to make those molecules.
 67

Moreover, losing this ability gives the animal an advantage because it stops using energy and resources
to make molecules that are already in its diet. Thus, as the diet of animals became more varied, they
tended to lose their abilities to make such widely available molecules as some of the amino acids.

The Metabolic Fates of Nutrients in Heterotrophs

When a heterotroph consumes nutrients, they can be categorized into macronutrients and
micronutrients.

Macronutrients are needed in large quantities and include carbohydrates, lipids, and proteins.

Micronutrients are required in small quantities and consist of organic vitamins and inorganic
minerals. Together, these nutrients fulfill the animal’s dietary requirements.

Calories and Energy

The energy value of food is measured in terms of calories or Calories.

A calorie (L. calor, heat) is the amount of energy needed to raise the temperature of 1 g of water by
1°C. A calorie, written with a small 'c', is also known as a gram calorie.

A kilocalorie, often referred to as a Calorie or kilogram calorie (kcal), is equal to 1,000 calories.

In everyday language, when we mention calories, we usually mean Calories because this larger unit is
more practical for measuring food energy.

For instance, if an advertisement states that a "light beer" contains 95 calories per 12 oz, it actually
means 95,000 calories, which translates to 95 Calories or 95 kcal.

Macronutrients
With a few exceptions, heterotrophs need organic molecules like carbohydrates, lipids, and proteins
in their diets. Enzymes break down these molecules into components that can be used for energy
production or as sources for life's building blocks.

Carbohydrates: Carbon and Energy from Sugars and Starches

The primary source of energy for heterotrophs is complex carbohydrates. Most carbohydrates
originate from plants. They can be in the form of various polysaccharides, disaccharides, or simple
sugars (monosaccharides). Carbohydrates also serve as a major carbon source for important organic
compounds.
 68

Lipids: Highly Compact Energy

Storage Nutrients
Neutral lipids (fats) or triacylglycerols are found in
fats and oils, meat and dairy products, nuts, and
certain fruits and vegetables rich in fats. Lipids are
the most concentrated source of food energy,
providing about 9 Calories (kcal) of usable energy
per gram, more than twice the energy available from
an equal mass of carbohydrate or protein.

Many heterotrophs have a definite dietary need for

lipids, sometimes for specific types. For instance,
many animals require unsaturated fatty acids,
which serve as precursor molecules for the synthesis
of sterols, such as cholesterol. Sterols are essential
for the synthesis of steroid hormones and are
incorporated into cell membranes. Other lipids function in insulating the bodies of some vertebrates and
help maintain a constant temperature.

Proteins: Basic to the Structure and Function of Cells

Animal sources of protein include other animals and milk. Plant sources include beans, peas, and nuts.
Proteins are needed for their amino acids, which heterotrophs use to build their own body proteins.

Micronutrients
Micronutrients typically consist of small ions, organic vitamins, inorganic minerals, and molecules
crucial for enzymatic reactions or as parts of certain proteins (e.g., copper in hemocyanin and iron in
hemoglobin). Despite being required in small amounts, animals cannot produce them rapidly, if at all;
therefore, they must obtain them from their diet.

Minerals
Some minerals are necessary in relatively large quantities and are termed essential minerals or
macrominerals. For instance, sodium and potassium are essential for the proper function of every
nerve and muscle in an animal’s body. Animals lose
significant amounts of these minerals, particularly
sodium, in their urine each day. Animals that sweat
to regulate body temperature also lose sodium in
their sweat. Adequate calcium intake is essential for
muscular activity and, along with phosphorus, for
bone formation. Table 27.2 outlines the functions of
the major essential minerals.
 69

Other minerals are referred to as trace minerals, trace elements, or microminerals. These are required
in very small amounts for various enzymatic functions. Table 27.3 provides a list of the functions of
some trace minerals.

Vitamins
Normal metabolic activity relies on small amounts of
over a dozen organic substances known as vitamins. The
term "vitamin" (L. vita, life) encompasses a variety of
chemically unrelated organic compounds found in many
foods in small quantities. They are necessary for normal
metabolic functioning. Vitamins can be water-soluble or
fat-soluble.

Water-Soluble Vitamins

Most water-soluble vitamins, such as the B vitamins

and vitamin C, are coenzymes required in metabolism.
These vitamins participate in various enzymatic reactions
essential for energy production and other metabolic
processes.

Fat-Soluble Vitamins

Fat-soluble vitamins have diverse functions.

These vitamins include vitamin A, vitamin D,
vitamin E, and vitamin K. They play roles in
various bodily functions, including vision, bone
health, antioxidant defense, and blood clotting.

Dietary Variations in Vitamin Requirements

The need for vitamin C and the fat-soluble

vitamins (A, D, E, and K) tends to be limited to
vertebrates. However, even within closely related
groups, vitamin requirements can differ. For
example, among vertebrates, humans and guinea
pigs require vitamin C, while rabbits do not.
Similarly, some birds require vitamin A, while
others do not.

Digestion
In some of the simplest life forms like protists
and sponges, certain cells directly take in whole
food particles from the environment through processes like diffusion, active transport, and/or
endocytosis. These cells then break down the food with enzymes to obtain nutrients. This approach is
known as intracellular digestion.

Intracellular digestion eliminates the need for mechanical breakdown of food or for a gut or other cavity
for chemical digestion. However, it limits an animal’s size and complexity because only very small
pieces of food can be utilized. Intracellular digestion provides nutrients in protozoa, sponges, cnidarians,
platyhelminths, rotifers, bivalve molluscs, and primitive chordates.

Extracellular Digestion
 70

Larger animals have evolved structures and mechanisms for extracellular digestion: the enzymatic
breakdown of larger food pieces into constituent molecules, typically in a specialized organ or cavity.
Nutrients from the food then pass into body cells lining the organ or cavity and can be used for energy
metabolism or biosynthesis.

Animal Strategies for Getting and Using Food

In most animals, nutrients cannot be absorbed directly from the external environment via intracellular
digestion. Therefore, animals must actively acquire their nutrients. The diversity of specializations for
food procurement (feeding) and extracellular digestion across animal species is vast. Here's a brief
overview of the major feeding strategies animals employ:

Continuous vs. Discontinuous Feeders

One aspect related to the structure of digestive systems is whether an animal is a continuous or
discontinuous feeder.

• Continuous Feeders: These animals are often slow-moving or sessile (remaining permanently
in one place). For instance, aquatic suspension feeders like tube worms and barnacles stay in
one place and continuously filter small food particles from the water.
• Discontinuous
Feeders:
Typically,
these animals
are active and
sometimes
highly mobile.
They often
possess more
digestive
specializations
than
continuous
feeders
because they consume large meals that require grinding up or storage, or both.
Carnivores
 71

Many carnivores pursue and capture relatively large prey. When successful, they consume large meals,
enabling them to avoid continuous hunting. Carnivores have digestive systems that allow for the storage
and gradual digestion
of large, infrequent
meals.

Herbivores

Herbivores spend more

time eating than
carnivores but are also
discontinuous feeders.
They must move from
one area to another
when food is depleted
and, in natural
environments, must
limit their grazing time
to avoid excessive
exposure to predators.
Herbivores have
digestive systems that
facilitate relatively
rapid food gathering
and gradual digestion.

Suspension
Feeders
Suspension feeding
involves the removal of
suspended food particles from the surrounding water using a capture, trapping, or filtration structure.
This feeding strategy includes three steps:

1. Transport of water past the feeding structure

 72

2. Removal of nutrients
from the water
3. Transport of the
nutrients to the mouth
of the digestive system
Suspension feeders include:

• Sponges
• Ascidians
• Branchiopods
• Ectoprocts
• Entoprocts
• Phoronids
• Most bivalves
• Many crustaceans
• Polychaetes
• Gastropods
• Some nonvertebrate
chordates

Deposit Feeders
Deposit feeding primarily involves omnivorous animals obtaining nutrients from sediments of soft-
bottom habitats or terrestrial soils. There are two main types of deposit feeding:

• Direct deposit feeding: Animals swallow large quantities of sediment, digest usable nutrients,
and excrete the remains.
• Indirect deposit feeding: Animals use tentacle-like structures to consume sediment.
Examples of deposit feeders include:

• Polychaete annelids
• Some snails
• Some sea urchins
• Most earthworms
• Sea cucumbers
• Most sipunculans
• Certain clams
• Several types of polychaetes

Herbivory
 73

Herbivory involves the consumption of macroscopic plants. This feeding strategy requires the ability
to bite and chew large pieces of plant matter (macroherbivory). Invertebrates that evolved
macroherbivory include:

• Molluscs
• Polychaete worms
• Arthropods
• Sea urchins
Examples of herbivores include insects and crustaceans, which have large, powerful mandibles capable
of biting off plant material and grinding and chewing it before ingestion.

Predation
Predation (L. praedator, a plunderer, pillager) is one of the most sophisticated feeding strategies as it
involves the capture of live prey. Predators can be classified based on how they capture their prey:

• Motile Stalkers: These predators actively pursue their prey. Examples include ciliate protozoa,
nemerteans, polychaete worms, gastropods, octopuses, squids, crabs, sea stars, and many
vertebrates.
• Lurking Predators: These predators sit and wait for their prey to come within seizing distance.
Examples include certain species of praying mantises, shrimp, crabs, spiders, polychaetes, and
many vertebrates.
• Sessile Opportunists: These predators are usually not very mobile and can only capture prey
when the prey organism comes into contact with them. Examples include certain protozoa,
barnacles, and cnidarians.
• Grazing Carnivores: These predators move about the substrate picking up small organisms,
primarily sessile and slow-moving animals like sponges, ectoprocts, tunicates, snails, worms,
and small crustaceans.

Surface Nutrient Absorption

Some highly specialized animals have evolved to absorb nutrients directly from the external medium
across their body surfaces, dispensing entirely with mechanisms for prey capture, ingestion, and
digestive processes. Examples include:

• Free-living Protozoa: Some protozoa, such as Chilomonas, absorb all their nutrients across
their body surface.
• Endoparasitic Organisms: Endoparasitic protozoa, cestode worms, endoparasitic gastropods,
and crustaceans lack mouths and digestive systems, absorbing all nutrients across their body
surface.
• Nonparasitic Multicellular Animals: Certain animals like gutless bivalves and pogonophoran
worms also lack a mouth and digestive system, absorbing nutrients across their body surface.
Pogonophoran worms, for instance, also supplement their nutrition with organic carbon fixed
by symbiotic bacteria within their tissues.

Fluid Feeders
Fluid feeding involves consuming the biological fluids of animals and plants, which are rich sources
of nutrients. This feeding strategy is particularly characteristic of some parasites and external parasites.
 74

• Parasites: Intestinal nematodes and other parasites feed by biting, rasping off host tissue, or
sucking blood.
• External Parasites (Ectoparasites): Leeches, ticks, mites, lampreys, and certain crustaceans
are examples of external parasites that use various mouthparts to feed on body fluids.
Examples of Fluid Feeders:

• Sea Lamprey: This parasite has a funnel-like structure surrounding its mouth, lined with over
200 rasping teeth and a rasplike tongue. It grips its fish host with the funnel, rasps a hole in the
fish’s body wall with its tongue, and sucks blood and body fluids from the wound.
• Insects: Insects have highly developed sucking structures for fluid feeding. Butterflies, moths,
aphids, and blood-sucking mosquitoes are examples. Mosquitoes have complex mouthparts
with piercing stylets, while butterflies and moths have tubelike mouthparts to suck up plant
fluids.
• Birds: Pollen- and nectar-feeding birds, like some butterflies, have long bills and tongues
specialized for particular types of flowers. Some birds have brushlike or hollow-tipped tongues
to collect nectar, while others make holes in flowers to access nectar with their tongues.
• Mammals: The only mammals that feed exclusively on blood are vampire bats, such as
Desmodus, found in tropical South and Central America. They use sharp front teeth to pierce
surface blood vessels and lap at the wound. Nectar-feeding bats and mammals like the honey
possum also have specialized adaptations for fluid feeding.

Diversity in Digestive Structures: Invertebrates

In primitive multicellular animals like cnidarians, the gut is a blind (closed) sac known as a
gastrovascular cavity, possessing only one opening serving as both entrance and exit. This structure
constitutes an incomplete digestive tract. Some specialized cells within the cavity secrete digestive
enzymes to initiate extracellular digestion, while other phagocytic cells lining the cavity engulf food
material and continue intracellular digestion within food vacuoles. A similar digestive pattern is
observed in some flatworms.

An evolutionary breakthrough in digestive structures occurred with the development of the anus and
complete digestive tract in the aschelminths. A complete digestive tract facilitates the one-way flow of
ingested food without mixing it with previously ingested food or waste. Moreover, complete digestive
tracts allow for progressive digestive processing in specialized regions along the system, enabling
efficient digestion in a series of distinctly different steps. Variations in the basic plan of a complete
digestive tract correlate with different food-gathering mechanisms and diets.

Examples of Digestive Systems in Protozoa and Invertebrates:

1. Incomplete Digestive System in Ciliated Protozoan: This represents an example of an

intracellular digestive system.
2. Bivalve Mollusc: Bivalve molluscs showcase both intracellular and extracellular digestion.
3. Insect: Insects exhibit extracellular digestion and possess a complete digestive tract.
These examples highlight the diversity of digestive structures in protozoa and invertebrates, each
adapted to specific feeding mechanisms and dietary needs.
 75

Protozoa
protozoa exhibit various nutritional modes, including
autotrophic, saprozoic, or heterotrophic (ingestion of
food particles). Among them, ciliated protozoa serve
as prime examples of organisms employing
heterotrophic nutrition.

Heterotrophic Nutrition in Ciliated Protozoa:

Ciliary action plays a crucial role in directing food

from the environment into the buccal cavity and
cytostome. The cytostome, in turn, leads to the
cytopharynx, which expands as food enters and then
pinches off a food-containing vacuole. As the
detached food vacuole moves through the cytoplasm,
excess water is eliminated, and the contents are
subjected to acidification followed by alkalization.
Additionally, digestive enzymes are added by
lysosomes. Within the vacuole, food particles
undergo digestion, and the resulting nutrients are absorbed into the cytoplasm. Waste products are
excreted via the cytopyge from the residual vacuole.

This process illustrates the intricacies of heterotrophic nutrition in ciliated protozoa, highlighting the
coordinated action of various cellular components for food ingestion, digestion, and waste excretion.

Bivalve Molluscs
Many bivalve molluscs engage in suspension feeding, ingesting small food particles from their
environment. Their digestive tract consists of several parts, including a short esophagus, stomach,
midgut, hindgut, and rectum.

Digestive Tract Structure:

• Esophagus: Connects the mouth to the stomach.

• Stomach: Contains a crystalline style, gastric shield, and diverticulated region.
• Diverticulae: Blind-ending sacs that increase surface area for absorption and intracellular
digestion.
 76

• Midgut, Hindgut, and Rectum: These parts function in extracellular digestion and
absorption.
Digestive Processes:

• Feeding Cycle: Involves the mechanical and enzymatic breakdown of food.

• Extracellular
Digestion
Cycle: Prepares
for digestion
and absorption.
• Intracellular
Digestion
Cycle: Releases
nutrients into
the blood and
produces
fragmentation
spherules for
waste excretion.
Integration with Tidal
Patterns:

• These cycles are

synchronized
with the tidal
immersion and
emersion of the
mollusc, ensuring efficient feeding and digestion based on environmental conditions.
This coordination of feeding and digestion processes reflects the adaptation of bivalve molluscs to their
suspension-feeding lifestyle, optimizing nutrient acquisition in their aquatic habitat.

Insects: Grasshopper Digestive System

Overview: The grasshopper serves as a typical example of an insect with a complete digestive tract
and extracellular digestion.

Feeding Process:

1. Mastication: Food is broken down by the mandibles and maxillae.

2. Salivary Secretion: Salivary glands secrete saliva containing the enzyme amylase to begin
carbohydrate digestion.
3. Transportation to Crop: Food is passed into the crop via the esophagus for temporary storage.
4. Enzymatic Digestion: Additional enzymes (carbohydrases, lipases, proteases) from the
midgut aid in digestion.
5. Stomach Processing: Food particles are mechanically reduced and sorted in the stomach.
6. Extracellular Digestion: Small particles move to the gastric cecae for completion of
extracellular digestion.
7. Nutrient Absorption: Most nutrient absorption occurs in the intestine.
 77

8. Water and Ion Absorption: In the rectum, water and ions are absorbed from undigested food.
9. Fecal Pellet Formation: Solid fecal pellets are formed and expelled through the anus.
Regulation and Control:

• Throughout the feeding process, the nervous system and endocrine system play crucial roles
in regulating enzyme production at different stages of digestion.
• Presence of food also influences enzyme secretion and digestive activity in various parts of the
digestive tract.
The coordinated action of the nervous and endocrine systems, along with the digestive processes,
ensures efficient digestion and nutrient absorption in grasshoppers and other insects.

Diversity in Digestive Structures: Vertebrates

The vertebrate digestive tract, also known as the gut tube, is specialized in both structure and function
to digest a wide variety of foods. The basic structures of the gut tube include the buccal cavity, pharynx,
esophagus, stomach, small intestine, large intestine, rectum, and anus or cloaca. Additionally, three
important glandular systems are associated with the digestive tract: the salivary glands, the liver
(including the gallbladder and bile duct), and the pancreas (including the pancreatic duct).

Feeding as a Universal Activity

Most vertebrates spend the majority of their time acquiring food, making feeding a universal pastime.
The structure of the oral cavity, teeth, intestines, and other major digestive structures usually reflects
how an animal gathers food, the type of food it eats, and the way it digests that food. These major
digestive structures illustrate the diversity of form and function among different vertebrates.

Tongues
A tongue or tongue-like structure develops in the floor of the oral cavity in many vertebrates:

• Lampreys: Have a protrusible tongue with horny teeth that rasp the flesh of their prey.
• Fishes: May have a primary tongue that bears teeth to help hold prey, though it is not muscular.
• Tetrapods: Have evolved mobile tongues for gathering food. For example, frogs, salamanders,
and some lizards can rapidly project part of their tongue to capture insects. Woodpeckers have
long, spiny tongues for gathering insects and grubs. Ant- and termite-eating mammals use long,
sticky tongues. Carnivores like cats have spiny papillae on their tongues to help rasp flesh from
bones.

Teeth
Most vertebrates have teeth, except for birds, turtles, and baleen whales:

• Birds: Lack teeth, likely to reduce body weight for flight.

 78

• Snakes: Have backward-sloping teeth to aid in

retaining prey while swallowing.
• Wolves: Have specialized canine teeth for ripping food.
• Herbivores: Such as deer, have predominantly
grinding teeth. Beavers use their front teeth for
chiseling trees and branches. Elephants have two upper
front teeth specialized as tusks for weapons and moving
objects.
• Omnivores: Like humans, pigs, bears, and raccoons
have versatile teeth capable of tearing, ripping,
chiseling, and grinding.

Salivary Glands
The presence and function of salivary glands vary among
vertebrates:

• Fishes: Generally lack salivary glands, with the

exception of lampreys, which have glands that secrete
an anticoagulant to keep prey's blood flowing.
• Snakes: Some have modified salivary glands that produce venom to immobilize prey.
• Amphibians and Reptiles: Typically lack significant salivary glands since oral digestive
enzymes are not crucial.
• Birds: Mostly lack salivary glands.
• Mammals: Universally possess salivary glands for moistening food and initiating digestion.

Esophagi
The length of the esophagus varies:

• Fishes and Amphibians: Have short

esophagi.
• Amniotes: Possess longer esophagi
due to their longer necks. Grain- and
seed-eating birds have a crop that
develops from the caudal portion of
the esophagus, ensuring a
continuous supply of food to the
stomach and intestine, thus reducing
the frequency of feeding while
maintaining a high metabolic rate.

Stomachs
The stomach is an ancestral vertebrate
structure:

• Function: Evolved to store food and

possibly to kill bacteria with
hydrochloric acid (HCl). The enzyme pepsinogen, essential for digestion, may have evolved
later.
 79

Gizzards
Some vertebrates possess a gizzard:

• Fishes, Reptiles (like crocodilians), and Birds: Have a gizzard for grinding food. In birds, the
gizzard develops from the posterior part of the stomach (ventriculus). Swallowed pebbles (grit)
retained in the gizzard facilitate the grinding process, especially in grain-eating birds.
These adaptations illustrate how vertebrates have evolved diverse digestive structures to efficiently
process the wide variety of foods available in their environments.

Ruminant Mammals
Ruminant mammals—animals that “chew their cud”, such as cows, sheep, and deer—exhibit some
of the most unusual modifications of the stomach. This digestion method allows animals to eat large
amounts of food quickly and chew it later at a more comfortable or safer location. The ruminant
stomach also supports large populations of microorganisms that digest the cellulose in grass and other
vegetation.

Cellulose Digestion

Cellulose contains a significant amount of energy, but animals generally cannot produce cellulase, the
enzyme needed to digest cellulose. Gut microorganisms can produce cellulase, making the
herbivorous lifestyle more effective.

Stomach Structure

In ruminants, the stomach has four compartments:

1. Rumen: The upper portion that expands to form a large pouch where food encounters
microorganisms.
2. Reticulum: A smaller compartment where partially digested food is regurgitated as “cud” for
thorough chewing.
3. Omasum: A small antechamber where the more liquid digested food moves after being
swallowed again.
4. Abomasum: The “true” stomach, where digestive enzymes are first encountered, and
digestion continues.
Digestive Process

1. Food enters the rumen: Encounters microorganisms that begin partial digestion, aided by fluid
secretions, body heat, and churning.
2. Moves to the reticulum: Mouthfuls are regurgitated as cud and thoroughly chewed.
3. Reswallowed food reenters the rumen: Becomes more liquid in consistency.
4. Flows to the omasum: Digested material moves to the omasum as it becomes very liquid.
5. Enters the abomasum: Encounters digestive enzymes, continuing digestion.

CECA
 80

In nonruminant herbivores,
microbial action on cellulose occurs
after digestion in the cecum:

• Rabbits, horses, and rats:

Maintain a population of
microorganisms in their large
cecum, the blind pouch
extending from the colon.
• Coprophagy: Some, like
mice and rabbits, eat their
own feces to reprocess
remaining materials such as
vitamins.

LIVERS AND GALLBLADDERS

In vertebrates with a gallbladder, it is closely associated with the liver:

• Liver: Manufactures bile, containing bile salts and bile pigments.

• Bile salts: Emulsify dietary fat, aiding digestion by lipase.
• Bile pigments: Result from phagocytosis of red blood cells, producing bilirubin (“red bile”)
and biliverdin (“green bile”).
• Gallbladder size: Large in carnivores and vertebrates with fat-rich diets; reduced or absent in
bloodsuckers (e.g., lamprey) and primarily plant-eating animals (e.g., some teleosts, many
birds, and rats).

PANCREATA
All vertebrates have a pancreas:

• In lampreys and lungfishes: Embedded in the wall of the intestine, not visible.
• Pancreatic fluid: Contains many enzymes, empties into the small intestine via the pancreatic
duct.

INTESTINES
Configuration and divisions of the small and large intestines vary greatly among vertebrates,
influenced by:

• Type of food
• Body size
• Levels of activity
Examples of Intestinal Variations

• Cyclostomes, chondrichthian fishes, primitive bony fishes: Have short, nearly straight
intestines extending from the stomach to the anus.
• More advanced bony fishes: Intestines increase in length and begin to coil.
• Amphibians and reptiles: Possess moderately long intestines.
• Birds and mammals: Have longer intestines with more surface area.
 81

Specific Features

• Birds: Typically have two ceca.

• Mammals: Have a single cecum at the beginning of the large intestine.
• Large intestine: Much longer in mammals than in birds, emptying into the cloaca in most
vertebrates.

The Mammalian Digestive System

Omnivores such as humans, pigs, bears, and raccoons possess a digestive system capable of processing
a variety of foods. The following sections discuss gastrointestinal motility control, major parts of the
alimentary canal, and accessory organs of digestion.

Digestive Process Steps

1. Ingestion: Eating
2. Peristalsis: Involuntary, sequential muscular contractions moving nutrients along the digestive
tract
3. Segmentation: Mixing the contents in the digestive tract
4. Secretion: Release of hormones, enzymes, and specific ions and chemicals involved in
digestion
5. Digestion: Conversion of large nutrient particles or molecules into smaller particles or
molecules
 82

6. Absorption: Passage of usable

nutrient molecules from the small
intestine into the bloodstream and
lymphatic system to body cells
7. Defecation: Elimination of
undigested and unabsorbed material
as waste
Key Components

• Control of gastrointestinal
motility: Regulation of movements
within the digestive system
• Major parts of the alimentary
canal: Includes the mouth, pharynx,
esophagus, stomach, small
intestine, and large intestine
• Accessory organs: Salivary glands,
liver, gallbladder, and pancreas
This comprehensive process ensures the
efficient breakdown, absorption, and
excretion of nutrients and waste in
mammals.

Gastrointestinal Motility and Its

Control
The function of the gastrointestinal (GI)
tract is determined by its tissue types,
which maintain a consistent anatomical structure throughout its length.

Anatomical Structure

1. Serosa: Outer layer of connective tissue, forming the moist epithelial sheet called the
peritoneum.
• The peritoneum lines the abdominal cavity and covers internal organs, encompassing
the coelom.
2. Muscle Layers:
• Longitudinal smooth muscle layer
• Circular smooth muscle layer
3. Submucosa: Connective tissue containing blood and lymphatic vessels.
4. Mucosa: Faces the central opening called the lumen.
Types of Movements

1. Peristalsis:
• Function: Advances food through the GI tract.
• Mechanism: Rings of circular smooth muscle contract behind and relax in front of the
food, similar to squeezing icing from a pastry tube.
 83

2. Segmentation:
• Function: Mixes food with digestive secretions, enhancing absorption efficiency.
• Mechanism: Rings of smooth muscles contract and relax, creating an oscillating back-
and-forth movement in the same place.
Sphincters

• Role: Control the flow of

material and prevent
backflow.
• Examples:
• Cardiac
sphincter:
Between
esophagus and
stomach.
• Pyloric sphincter:
Between stomach
and small intestine.
Control Mechanisms

Local Control:

• Mechanical receptors:
Stimulated by gut
distension due to ingested
food.
• Chemical receptors:
Stimulated by digestion of
carbohydrates, lipids, and
proteins.
• Nerve Plexuses: Signals
travel through these in the
gut wall to control muscular
contractions (peristalsis and
segmentation) and
secretion of substances
(e.g., mucus, enzymes).
Central Nervous System (CNS) Control:

• Long-distance nerve pathways: Connect receptors and effectors with the CNS, functioning to
maintain homeostasis.
Endocrine Control:

• Hormones: Produced by endocrine cells in the GI tract to regulate secretion, digestion, and
absorption.

Oral Cavity
 84

The oral cavity (mouth) is protected by a pair of lips, highly vascularized skeletal muscle tissue rich in
sensory nerve endings. Lips aid in food retention during chewing and contribute to sound modification
during phonation.

Components

1. Lips: Highly vascularized, skeletal muscle tissue with sensory nerve endings.
2. Tongue and Teeth: Mammals can mechanically process a wide range of foods due to enamel-
covered teeth, the hardest material in the body, and strong jaw and tooth forces.
3. Saliva: Continuously bathes the oral cavity, secreted by at least three pairs of salivary glands.
• Functions: Moistens food, binds it
with mucins (glycoproteins),
forming a moist mass (bolus).
Contains bicarbonate ions (HCO3)
for buffering mouth chemicals,
thiocyanate ions (SCN) and
lysozyme for microbial control,
and amylase for carbohydrate
digestion initiation.

Pharynx and Esophagus

Swallowing Process

• Initiation: Voluntary or involuntary

swallowing reflex.
• Mechanism: Sequential, involuntary
contractions of smooth muscles in the
esophageal walls propel the bolus or
liquid to the stomach.
• Epiglottis: Temporarily seals off the
tracheal opening (glottis) to prevent
food entry into the trachea during
swallowing.
Digestive Contribution

• Neither the pharynx nor the

esophagus contribute to digestion.
They serve as the common passageway
for both the digestive and respiratory
tracts.

Stomach
The mammalian stomach is a muscular, distensible sac with three main functions:

1. Storage and Mixing: Receives food bolus from the esophagus and mixes it.
2. Secretion: Releases substances (enzymes, mucus, hydrochloric acid [HCl]) initiating protein
digestion.
 85

3. Control of Food Movement: Regulates the rate of food passage into the small intestine via the
pyloric sphincter.
Structure and Function

• Mucous Membrane: Contains thousands of gastric glands.

• Cell Types:
• Parietal Cells: Secrete a solution containing HCl.
• Chief Cells: Secrete pepsinogen, the precursor of pepsin.
• Mucous Cells: Secrete mucus to protect the stomach from HCl and digestive
enzymes.
• Hormonal Regulation: Endocrine cells release gastrin hormone, stimulating gastric gland
activity.
• Digestive Process:
• Bolus Entry: Food bolus distends stomach walls, stimulating gastric gland secretion
of HCl and pepsinogen.
• Activation: HCl converts pepsinogen into active enzyme pepsin.
• Mixing and Churning: Smooth mucosal muscles contract, churning and mixing the
food bolus, initiating protein breakdown.
• Chyme Formation: After about three to four hours, stomach contents become semiliquid mass
called chyme (Gr. chymos, juice).
• Pyloric Sphincter: Regulates release of chyme into the small intestine.
• Peristaltic Waves: Cease when stomach is empty, but may occur after about 10 hours of
fasting, causing "hunger pangs" due to sensory nerve impulses to the brain.

Small Intestine: Main Site of Digestion

Most of the ingested food in mammals is digested and absorbed in the small intestine. Here's a
breakdown of its structure and function:

• Dimensions: Approximately 4 cm in diameter and 7 to 8 m in length in humans, providing a

substantial surface area for nutrient absorption.
• Surface Area: Enhanced by circular folds and villi, with each square centimeter of mucosa
bearing thousands of fingerlike villi.
• Epithelial Cells: Covered with simple columnar epithelial cells, each with numerous
microvilli, further increasing surface area.
• Functional Segments:
• Duodenum: Primarily functions in digestion, containing many digestive enzymes
secreted by intestinal glands and the pancreas.
 86

• Jejunum and Ileum: Absorb end products of digestion, including amino acids, simple
sugars, fatty
acids, and
glycerol.
• Absorption
Mechanisms:
• Active
Transport:
Sugars and
amino acids
absorbed
into
capillaries
of villi.

• Chylomicron Formation: Free fatty acids enter epithelial cells of villi, recombine with
glycerol to form triglycerides, coated with proteins to form chylomicrons, which enter
lacteals of villi and eventually the bloodstream.
• Water and Mineral Absorption: Small intestine absorbs about 9 liters of water per day, with
the large intestine absorbing the rest.

Large Intestine
• Structure: Lacks circular folds, villi, or microvilli, resulting in a smaller surface area compared
to the small intestine.
• Cecum: Blind-ended sac where the small intestine joins the large intestine; serves as a storage
site and possibly an evolutionary remnant.
• Appendix: Contains abundant lymphoid tissue, possibly functioning as part of the immune
system.
• Functions:
• Water and Mineral Reabsorption: Essential for maintaining hydration and
electrolyte balance.
 87

• Feces Formation: Stores and compacts waste material.

• Peristalsis: Moves food residue along, facilitating water and mineral absorption.
• Microbiota: Contains symbiotic bacteria and fungi that further break down organic molecules
and secrete amino acids and vitamin K.
• Feces: Mixture of bacteria, fungi, undigested plant fiber, sloughed-off intestinal cells, and other
waste products.

Role of the Pancreas in Digestion

The pancreas is a vital organ located just ventral to the stomach, serving both endocrine and exocrine
functions. Here's an overview of its role in digestion:

• Exocrine Function:
• Secretion of Digestive Enzymes: Exocrine cells in the pancreas release digestive
enzymes into the pancreatic duct.
• Composition of Enzymes:
• Protein Digestion: Trypsin, carboxypeptidase, and chymotrypsin break down
proteins into small peptides and individual amino acids.
• Lipid Digestion: Pancreatic lipases split triglycerides into absorbable glycerol
and free fatty acids.
• Carbohydrate Digestion: Pancreatic amylase converts polysaccharides into
disaccharides and monosaccharides.
• Bicarbonate Secretion:
• The pancreas also secretes bicarbonate (HCO3 ) ions into the pancreatic duct.
• Neutralization of Acidity: Bicarbonate helps neutralize the acidic food residue from
the stomach, raising the pH from 2 to 7 for optimal digestion.
• Facilitates Enzyme Function: Without neutralization, pancreatic enzymes would not
function effectively.
 88

Role of the Liver and Gallbladder in Digestion

Liver:
• Location: The liver, the largest organ in the mammalian body, is situated just under the
diaphragm.
• Function of Hepatocytes:
• Nutrient Processing: Millions of specialized cells called hepatocytes in the liver
absorb nutrients from the intestines and release them into the bloodstream.
• Blood Protein Synthesis: Hepatocytes manufacture blood proteins like prothrombin
and albumin.
• Major Metabolic Functions:
• Removal of amino acids from organic compounds.
• Formation of urea from proteins and conversion of excess amino acids into urea to
reduce ammonia levels in the body.
• Synthesis of plasma proteins, formation of fetal erythrocytes, and destruction of worn-
out erythrocytes.
• Synthesis of nonessential amino acids.
• Conversion of galactose and fructose to glucose.
• Oxidation of fatty acids.
 89

• Formation of lipoproteins, cholesterol, and phospholipids.

• Conversion of carbohydrates and proteins into fat.
• Modification and detoxification of waste products, drugs, and toxins.
• Synthesis of vitamin A from carotene and participation in vitamin D activation.
• Maintenance of stable body temperature by raising the blood temperature.
• Manufacture of bile salts used for fat emulsification in the small intestine.

Gallbladder:
• Location: The gallbladder is a small organ near the liver.
• Function:
• Bile Storage: The gallbladder stores bile produced continuously by liver cells.
• Composition of Bile: Bile is alkaline and contains pigments, cholesterol, lecithin,
mucin, bilirubin, and bile salts.
• Emulsification of Fats: Bile salts act as detergents to emulsify fats, aiding in fat
digestion and absorption.
• Formation of Micelles: Bile salts combine with end products of fat digestion to form
micelles, facilitating fat absorption by crossing the intestinal epithelium.
The liver and gallbladder work together to produce and store bile, which is essential for the digestion
and absorption of fats in the small intestine. Additionally, the liver performs numerous metabolic
functions crucial for overall physiological homeostasis.
 90

Cell Biology:

Introduction to Cell Biology

What is Cell Biology?

Cell biology, also known as cytology, is a branch of biology that focuses on the structure, function, and
behavior of cells. Cells are the fundamental units of life, the building blocks from which all living
organisms are constructed. Understanding cells, their components, and how they function is essential
for all biological sciences.

Historical Background

The study of cells began with the invention of the microscope in the 17th century. Robert Hooke, in
1665, was the first to coin the term "cell" when he observed the cell walls in cork tissue. Later, Antonie
van Leeuwenhoek's improvements in microscopy allowed for the observation of living cells, including
bacteria and protozoa.

The Cell Theory

The development of cell theory in the 19th century was a pivotal moment in biology. The cell theory
states:

1. All living organisms are composed of one or more cells.

2. The cell is the basic unit of life.

3. All cells arise from pre-existing cells.

Types of Cells

Cells come in various forms and functions, but they are primarily categorized into two types:

1. Prokaryotic Cells: These cells lack a true nucleus and membrane-bound organelles. Bacteria
and archaea are examples of prokaryotic organisms.

2. Eukaryotic Cells: These cells have a true nucleus enclosed by a nuclear membrane and contain
various membrane

Eukaryotic Cell Definition

“Eukaryotic cells are the cells that contain a membrane bound nucleus
and organelles.”

What is a Eukaryotic Cell?

Eukaryotic cells have a nucleus enclosed within the nuclear membrane and form large and complex
organisms. Protozoa, fungi, plants, and animals all have eukaryotic cells. They are classified under the
kingdom Eukaryota.

They can maintain different environments in a single cell that allows them to carry out various metabolic
reactions. This helps them grow many times larger than the prokaryotic cells.

Characteristics of Eukaryotic Cells

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The features of eukaryotic cells are as follows:

1. Eukaryotic cells have the nucleus enclosed within the nuclear membrane.

2. The cell has mitochondria.

3. Flagella and cilia are the locomotory organs in a eukaryotic cell.

4. A cell wall is the outermost layer of the eukaryotic cells.

5. The cells divide by a process called mitosis.

6. The eukaryotic cells contain a cytoskeletal structure.

7. The nucleus contains a single, linear DNA, which carries all the genetic information.

Structure Of Eukaryotic Cell

The eukaryotic cell structure comprises the following:

Plasma Membrane

• The plasma membrane separates the cell from the outside environment.

• It comprises specific embedded proteins, which help in the exchange of substances in and out
of the cell.

Cell Wall

• A cell wall is a rigid structure present outside the plant cell. It is, however, absent in animal
cells.

• It provides shape to the cell and helps in cell-to-cell interaction.

• It is a protective layer that protects the cell from any injury or pathogen attacks.

• It is composed of cellulose, hemicellulose, pectins, proteins, etc.

Cytoskeleton

The cytoskeleton is present inside the cytoplasm, which consists of microfilaments, microtubules, and
fibres to provide perfect shape to the cell, anchor the organelles, and stimulate the cell movement.

Endoplasmic Reticulum

It is a network of small, tubular structures that divides the cell surface into two parts: luminal and
extraluminal.

• Rough Endoplasmic Reticulum contains ribosomes.

• Smooth Endoplasmic Reticulum that lacks ribosomes and is therefore smooth.

Nucleus

• The nucleoplasm enclosed within the nucleus contains DNA and proteins.

• The nuclear envelop consists of two layers- the outer membrane and the inner membrane. Both
the membranes are permeable to ions, molecules, and RNA material.

• Ribosome production also takes place inside the nucleus.

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Golgi Apparatus

• It is made up of flat disc-shaped structures called cisternae.

• It is absent in red blood cells of humans and sieve cells of plants.

• They are arranged parallel and concentrically near the nucleus.

• It is an important site for the formation of glycoproteins and glycolipids.

Ribosomes

These are the main site for protein synthesis and are composed of proteins and ribonucleic acids.

Mitochondria

• These are also known as “powerhouse of cells” because they produce energy.

• It consists of an outer membrane and an inner membrane. The inner membrane is divided into
folds called cristae.

• They help in the regulation of

cell metabolism.

Lysosomes

They are known as “suicidal bags”

because they possess hydrolytic
enzymes to digest protein, lipids,
carbohydrates, and nucleic acids.

Plastids

These are double-membraned structures

and are found only in plant cells. These
are of three types:

• Chloroplast that contains

chlorophyll and is involved in
photosynthesis.

• Chromoplast that contains a pigment called carotene that provides the plants yellow, red, or
orange colours.

• Leucoplasts that are colourless and store oil, fats, carbohydrates, or proteins.

Eukaryotic Cell Diagram

Eukaryotic cell diagram mentioned below depicts different cell organelles present in eukaryotic cells.
The nucleus, endoplasmic reticulum, cytoplasm, mitochondria, ribosomes, lysosomes are clearly
mentioned in the diagram.

Prokaryotic Cell Definition

“Prokaryotic cells are the cells that do not have a true nucleus and membrane-bound organelles.”

What is a Prokaryotic Cell?

 93

Prokaryotic cells are single-celled microorganisms known to be the earliest on earth. Prokaryotes
include Bacteria and Archaea. The photosynthetic prokaryotes include cyanobacteria that perform
photosynthesis.

A prokaryotic cell consists of a single membrane and therefore, all the reactions occur within the
cytoplasm. They can be free-living or parasites.

Characteristics of Prokaryotic Cell

Prokaryotic cells have different characteristic features. The characteristics of the prokaryotic cells are
mentioned below.

1. They lack a nuclear membrane.

2. Mitochondria, Golgi bodies, chloroplast, and lysosomes are absent.

3. The genetic material is present on a single chromosome.

4. The histone proteins, the important constituents of eukaryotic chromosomes, are lacking in
them.

5. The cell wall is made up of carbohydrates and amino acids.

6. The plasma membrane acts as the mitochondrial membrane carrying respiratory enzymes.

7. They divide asexually by binary fission. The sexual mode of reproduction involves conjugation.

Prokaryotic Cell Structure

A prokaryotic cell does not have a nuclear membrane. However, the genetic material is present in a
region in the cytoplasm known as the nucleoid. They may be spherical, rod-shaped, or spiral. A
prokaryotic cell structure is as follows:

1. Capsule– It is an outer protective covering found in the bacterial cells, in addition to the cell
wall. It helps in moisture retention, protects the cell when engulfed, and helps in the attachment
of cells to nutrients and surfaces.

2. Cell Wall– It is the outermost layer of the cell which gives shape to the cell.

3. Cytoplasm– The cytoplasm is mainly composed of enzymes, salts, cell organelles and is a gel-
like component.

4. Cell Membrane– This layer surrounds the cytoplasm and regulates the entry and exit of
substances in the cells.

5. Pili– These are hair-like outgrowths that attach to the surface of other bacterial cells.

6. Flagella– These are long structures in the form of a whip, that help in the locomotion of a cell.

7. Ribosomes– These are involved in protein synthesis.

8. Plasmids– Plasmids are non-chromosomal DNA structures. These are not involved in
reproduction.

9. Nucleoid Region– It is the region in the cytoplasm where the genetic material is present.

A prokaryotic cell lacks certain organelles like mitochondria, endoplasmic reticulum, and Golgi bodies.

Prokaryotic Cell Diagram

 94

The prokaryotic cell diagram

given below represents a
bacterial cell. It depicts the
absence of a true nucleus and
the presence of a flagellum
that differentiates it from a
eukaryotic cell.

Difference between
Prokaryotic and
Eukaryotic Cells
Though these two classes of
cells are quite different, they
do possess some common
characteristics. For
instance, both possess cell
membranes and ribosomes,
but the similarities end there.
The complete list of
differences between prokaryotic and eukaryotic cells is summarized as follows:

Prokaryotes Eukaryotes

Type of Cell Always unicellular Unicellular and multi-cellular

Cell size Ranges in size from 0.2 μm – 2.0 μm in Size ranges from 10 μm – 100 μm in
diameter diameter

Cell wall Usually present; chemically complex in When present, chemically simple in
nature nature

Nucleus Absent. Instead, they have a nucleoid region Present

in the cell

Ribosomes Present. Smaller in size and spherical in Present. Comparatively larger in size
shape and linear in shape

DNA arrangement Circular Linear

Mitochondria Absent Present

Cytoplasm Present, but cell organelles absent Present, cell organelles present
 95

Endoplasmic Absent Present

reticulum

Plasmids Present Very rarely found in eukaryotes

Ribosome Small ribosomes Large ribosomes

Lysosome Lysosomes and centrosomes are absent Lysosomes and centrosomes are
present

Cell division Through binary fission Through mitosis

Flagella The flagella are smaller in size The flagella are larger in size

Reproduction Asexual Both asexual and sexual

Example Bacteria and Archaea Plant and Animal cell

What are Cell Organelles?

The cellular components are called cell organelles. These cell organelles include both membrane and
non-membrane bound organelles, present within the cells and are distinct in their structures and
functions. They coordinate and function efficiently for the normal functioning of the cell. A few of them
function by providing shape and support, whereas some are involved in the locomotion and
reproduction of a cell. There are various organelles present within the cell and are classified into three
categories based on the presence or absence of membrane.

Organelles without membrane: The Cell wall, Ribosomes, and Cytoskeleton are non-membrane-
bound cell organelles. They are present both in the prokaryotic cell and the eukaryotic cell.

Single membrane-bound organelles: Vacuole, Lysosome, Golgi Apparatus, Endoplasmic Reticulum

are single membrane-bound organelles present only in a eukaryotic cell.

Double membrane-bound organelles: Nucleus, mitochondria and chloroplast are double membrane-
bound organelles present only in a eukaryotic cell.

Let us learn more in detail about the different cell organelles in brief.
 96

Ultrastructure of the Cell

The ultrastructure of a cell refers to its fine structural details, typically observed using electron
microscopy. This level of detail reveals the intricate organization and function of various cellular
components. Here's an overview of the ultrastructure of key cell organelles:

1. Plasma Membrane

• Structure:

• Phospholipid Bilayer: Composed of two layers of phospholipids with hydrophilic

(water-attracting) heads facing outward and hydrophobic (water-repelling) tails facing
inward.

• Proteins: Integral (span the bilayer) and peripheral (attached to the surface) proteins
involved in transport, signaling, and structural support.

• Cholesterol: Interspersed within the bilayer, providing fluidity and stability.

• Carbohydrates: Attached to proteins and lipids on the extracellular surface, forming

glycoproteins and glycolipids that play roles in cell recognition and adhesion.

2. Nucleus

• Structure:

• Nuclear Envelope: Double membrane with nuclear pores that control the movement
of molecules in and out of the nucleus.

• Nucleoplasm: Semi-fluid matrix inside the nucleus containing chromatin and the
nucleolus.

• Chromatin: Complex of DNA and histone proteins, which condenses to form

chromosomes during cell division.
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• Nucleolus: Dense region within the nucleus where ribosomal RNA (rRNA) synthesis
and ribosome assembly occur.

3. Endoplasmic Reticulum (ER)

• Rough Endoplasmic Reticulum (RER):

• Structure: Network of membranous sacs and tubules studded with ribosomes.

• Function: Synthesis of proteins destined for secretion, incorporation into the plasma
membrane, or use in lysosomes.

• Smooth Endoplasmic Reticulum (SER):

• Structure: Network of membranous tubules without ribosomes.

• Function: Synthesis of lipids, metabolism of carbohydrates, detoxification of drugs

and poisons, and storage of calcium ions.

4. Golgi Apparatus

• Structure:

• Cisternae: Stacked, flattened membranous sacs.

• Vesicles: Small, membrane-bound compartments that transport materials to and from

the Golgi.

• Function: Modifies, sorts, and packages proteins and lipids received from the ER for secretion
or delivery to other organelles.

5. Mitochondria

• Structure:

• Double Membrane: Outer membrane is smooth, while the inner membrane is highly
folded into cristae to increase surface area.

• Matrix: Inner space containing mitochondrial DNA, ribosomes, and enzymes involved
in metabolic processes.

• Function: Site of cellular respiration and ATP production through oxidative phosphorylation.

6. Lysosomes

• Structure:

• Membrane-bound Vesicles: Contain hydrolytic enzymes.

• Function: Degrade macromolecules, damaged organelles, and foreign particles through

enzymatic digestion.

7. Peroxisomes

• Structure:

• Membrane-bound Organelles: Contain enzymes such as catalase and oxidases.

• Function: Break down fatty acids and detoxify harmful substances, such as hydrogen peroxide.
 98

8. Cytoskeleton

• Structure:

• Microfilaments: Thin filaments composed of actin, involved in cell movement and

shape.

• Intermediate Filaments: Rope-like fibers providing mechanical support.

• Microtubules: Hollow tubes composed of tubulin, involved in cell division,

intracellular transport, and maintaining cell shape.

• Function: Provides structural support, facilitates cell movement, and aids in intracellular
transport and cell division.

9. Ribosomes

• Structure:

• Composed of rRNA and Proteins: Consist of two subunits (large and small).

• Free or Bound: Found free in the cytoplasm or bound to the RER.

• Function: Site of protein synthesis, translating mRNA into polypeptide chains.

10. Chloroplasts (in Plant Cells)

• Structure:

• Double Membrane: Outer and inner membranes enclose the stroma.

• Thylakoids: Membranous sacs stacked into grana where photosynthesis occurs.

• Stroma: Fluid containing enzymes, ribosomes, and chloroplast DNA.

• Function: Conduct photosynthesis to convert light energy into chemical energy stored in
glucose.

11. Vacuoles

• Structure:

• Membrane-bound Sacs: Larger in plant cells (central vacuole) than in animal cells.

• Function: Store nutrients, waste products, and maintain turgor pressure in plant cells.

Endoplasmic Reticulum
“Endoplasmic Reticulum is a complex network of tubular membranes exclusively present in the
cytoplasm of the eukaryotic cell.”
 99

The endoplasmic reticulum transpires in two forms: a type with a ribosome-studded surface and another
with a smooth surface. The latter is called the smooth endoplasmic reticulum, and the former is called
the rough endoplasmic reticulum. These membranes form continuous folds, eventually joining the
outer layer of the nuclear
membrane. Except for sperm cells
and red blood cells, the
endoplasmic reticulum is observed
in every other type of eukaryotic
cell.

Endoplasmic Reticulum
Diagram

The below diagram shows the

variants of the endoplasmic
reticulum:

• Rough ER

• Smooth ER

Rough endoplasmic reticulum has

ribosomes embedded within its
structure, giving a “rough”
appearance. The smooth
endoplasmic reticulum does not have these ribosomes, hence appearing “smooth.”

Structure of Endoplasmic Reticulum

The structure of the endoplasmic reticulum is shaped like a sac. Since ER is of two types, each has its
own distinguishing features:

Rough Endoplasmic Reticulum Structure

• The rough endoplasmic reticulum is named so because of its appearance.

• It is a series of connected flattened sacs having several ribosomes on its outer surface, hence
the name.

• It synthesizes and secretes proteins in the liver, hormones and other substances in the glands.

• Rough ER is prominent in cells where protein synthesis happens (such as hepatocytes)

Role in Protein Synthesis

• Ribosome Attachment: Ribosomes on the RER synthesize proteins destined for secretion,
incorporation into the plasma membrane, or use in lysosomes.
• Co-translational Translocation: As a protein is synthesized by a ribosome, it is threaded into
the lumen of the RER through a translocon channel.
• Protein Folding: Inside the RER lumen, proteins fold into their correct three-dimensional
shapes with the assistance of chaperone proteins.
• Post-translational Modifications: Proteins undergo initial modifications, such as
glycosylation (addition of carbohydrate groups), forming glycoproteins.
• Quality Control: The RER ensures that only properly folded and assembled proteins proceed
to the Golgi apparatus. Misfolded proteins are retained and eventually degraded.
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Smooth Endoplasmic Reticulum Structure

• The smooth endoplasmic reticulum, on the other hand, does not have ribosomes.

• The smooth endoplasmic reticulum has a tubular form.

• It participates in the production of phospholipids, the chief lipids in cell membranes and are
essential in the process of metabolism.

• Smooth ER transports the products of the rough ER to other cellular organelles, especially the
Golgi apparatus.

Role in Lipid Metabolism and Drug Detoxification

• Lipid Synthesis: The SER is involved in the synthesis of lipids, including phospholipids and
steroids, which are essential components of cellular membranes and hormones.
• Carbohydrate Metabolism: The SER helps in the metabolism of carbohydrates, including the
conversion of glycogen to glucose in liver cells.
• Calcium Storage: The SER stores calcium ions, which are crucial for muscle contraction and
other cellular signaling processes.
• Detoxification: The SER contains enzymes, such as cytochrome P450, that are involved in the
detoxification of drugs, alcohol, and other harmful substances. These enzymes catalyze
reactions that make these substances more water-soluble and easier to excrete from the body.
Functions of Endoplasmic Reticulum

As stated above, the endoplasmic reticulum is categorised into two types, and both these types of ER
perform specific functions:

Smooth Endoplasmic Reticulum Function:

• Smooth ER is responsible for the synthesis of essential lipids such as phospholipids and
cholesterol.

• Smooth ER is also responsible for the production and secretion of steroid hormones.

• It is also responsible for the metabolism of carbohydrates.

• The smooth ER store and releases calcium ions. These are quite important for the nervous
system and muscular systems.

Rough Endoplasmic Reticulum Function:

• The majority of the functions of rough ER is associated with protein synthesis.

• The rough endoplasmic reticulum also plays a vital role in protein folding.

• Also ensures quality control (regarding correct protein folding).

• The second most important function after protein synthesis and protein folding is protein
sorting.

Golgi Apparatus
The Golgi apparatus, also known as the Golgi complex or Golgi body, is a vital organelle in eukaryotic
cells responsible for modifying, sorting, and packaging proteins and lipids for secretion or delivery to
 101

other organelles. It plays a crucial role in the post-translational processing of proteins and the formation
of lysosomes.
Structure
• Cisternae: The Golgi apparatus consists of a series of flattened, membrane-bound sacs called
cisternae. These stacks are typically organized into three main regions:
• Cis-Golgi Network (CGN): The entry face, located nearest the endoplasmic reticulum
(ER). Vesicles from the ER fuse with the CGN.
• Medial Golgi: The central layers where most of the processing occurs.
• Trans-Golgi Network (TGN): The exit face, where modified molecules are sorted and
packaged into vesicles for transport to their final destinations.
• Vesicles: Small, membrane-bound sacs that transport materials between the ER, Golgi
apparatus, and other cellular compartments. Vesicles bud off from and fuse with the cisternae.

Role in synthesis of glycoprotein.

The Golgi apparatus is integral to the synthesis and maturation of glycoproteins. Glycoproteins are
proteins that have carbohydrate groups covalently attached to them, which are crucial for proper protein
folding, stability, signaling, and cell-cell recognition.

Steps in Glycoprotein Synthesis

1. Initial Glycosylation in the Endoplasmic Reticulum (ER):

• N-linked Glycosylation: The process begins in the ER, where an oligosaccharide is

attached to the nitrogen atom (N) of an asparagine residue in the nascent protein. This
oligosaccharide is a pre-assembled 14-sugar structure (often referred to as a "core
glycan") that is transferred from a lipid carrier (dolichol phosphate) to the protein.

• Quality Control: The ER ensures that only properly folded proteins proceed to the
Golgi apparatus. Misfolded proteins are retained and eventually degraded.

2. Transport to the Golgi Apparatus:

• Vesicular Transport: Once initial glycosylation and folding are complete, the
glycoproteins are packaged into COPII-coated vesicles and transported from the ER to
the cis-Golgi network (CGN).

3. Processing in the Golgi Apparatus:

• Cis-Golgi Network (CGN): The CGN receives the vesicles containing glycoproteins
from the ER. Initial modifications may begin here.

• Medial Golgi: As the glycoproteins move through the medial Golgi, extensive
remodeling of the oligosaccharide chains occurs. This can include the removal of
mannose residues and the addition of various sugars such as N-acetylglucosamine,
galactose, and sialic acid.

• Trans-Golgi Network (TGN): In the TGN, further modifications and sorting occur.
Terminal sugars are often added here, and the glycoproteins are sorted into different
vesicles based on their final destinations.

Specific Modifications in the Golgi Apparatus

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• Trimming and Addition of Sugars:

• Enzymes in the Golgi apparatus trim the initial oligosaccharide added in the ER and
sequentially add new sugars in a highly regulated manner.

• Enzymes Involved:

• Glycosidases: Remove specific sugar residues.

• Glycosyltransferases: Add sugars to specific positions on the growing

oligosaccharide chain.

• Complex and Hybrid N-glycans:

• In the Golgi, N-linked glycans can be processed into complex or hybrid forms, which
have different functional properties compared to the high-mannose forms typically
found in the ER.

• O-linked Glycosylation:

• Unlike N-linked glycosylation, O-linked glycosylation occurs entirely in the Golgi

apparatus. In this process, sugars are attached to the hydroxyl groups (O) of serine or
threonine residues in the protein. This type of glycosylation is common in mucins and
certain hormones.

Functional Importance of Glycoprotein Synthesis

• Protein Stability and Folding:

• Glycosylation helps in proper folding of the protein and increases its stability by
protecting it from proteolytic degradation.

• Cell-Cell and Cell-Matrix Interactions:

• Glycoproteins play critical roles in cell-cell recognition, signaling, and adhesion

processes. For example, they are important in immune responses where glycoproteins
on the surface of cells can act as antigens.

• Targeting and Trafficking:

• Proper glycosylation is crucial for the targeting of glycoproteins to their correct

destinations, such as lysosomes, plasma membrane, or for secretion outside the cell.

Mitochondria: Cell Respiration and Significance as a

Semi-Autonomous Organelle
Structure of Mitochondria
Double Membrane:

Outer Membrane:

• Smooth and permeable to small molecules and ions.

Inner Membrane:

• Highly folded into structures called cristae, increasing the surface area for cellular
respiration.
 103

Intermembrane Space:

The region between the outer and inner membranes.

Matrix:

• The innermost compartment containing mitochondrial DNA, ribosomes, and enzymes

necessary for the Krebs cycle and other metabolic pathways.

Cell Respiration

• Mitochondria are the powerhouses of the cell, responsible for producing ATP through
the process of cellular respiration. This process consists of three main stages:

Glycolysis (Cytoplasm):

• Glucose is broken down into pyruvate, yielding a small amount of ATP and NADH.

• Pyruvate is transported into the mitochondria for further processing.

Krebs Cycle (Citric Acid Cycle) (Matrix):

• Acetyl-CoA, derived from pyruvate, enters the Krebs cycle.

• This cycle produces ATP, NADH, FADH2, and releases CO2 as a byproduct.

• NADH and FADH2 are electron carriers that transfer electrons to the electron transport
chain (ETC).

Electron Transport Chain (ETC) and Oxidative Phosphorylation (Inner Membrane):

• NADH and FADH2 donate electrons to the ETC, which is composed of a series of
protein complexes and electron carriers embedded in the inner mitochondrial
membrane.

• As electrons move through the ETC, protons are pumped from the matrix into the
intermembrane space, creating a proton gradient (proton motive force).

• Protons flow back into the matrix through ATP synthase, driving the synthesis of ATP
from ADP and inorganic phosphate (Pi).

• Oxygen serves as the final electron acceptor, combining with electrons and protons to
form water.

Significance as a Semi-Autonomous Organelle

• Mitochondria exhibit several unique features that classify them as semi-autonomous
organelles:

Own DNA:

• Mitochondria contain their own circular DNA (mtDNA), which encodes some of the
proteins and RNAs required for mitochondrial function.

• mtDNA is inherited maternally and is similar to bacterial DNA, supporting the

endosymbiotic theory.

Ribosomes:
 104

• Mitochondria possess their own ribosomes (similar to bacterial ribosomes), allowing

them to synthesize some of their own proteins.

Replication and Division:

• Mitochondria can replicate independently of the cell cycle through a process similar to
binary fission in bacteria.

• The number of mitochondria within a cell can change in response to the cell’s energy
needs.

Protein Synthesis:

• While mitochondria can produce some proteins, the majority of mitochondrial proteins
are encoded by nuclear DNA, synthesized in the cytoplasm, and imported into the
mitochondria.

• Mitochondrial biogenesis requires the coordinated expression of both nuclear and

mitochondrial genomes.

Dual Genetic Control:

• Mitochondrial function is governed by both mitochondrial and nuclear genomes,

necessitating intricate coordination between these genetic systems.

Functional Significance
Energy Production:

• Mitochondria generate most of the cell’s ATP through oxidative phosphorylation,

providing the energy necessary for various cellular processes.

Metabolic Integration:

• Mitochondria play key roles in metabolic pathways, including the Krebs cycle, fatty
acid oxidation, and amino acid metabolism.

Apoptosis Regulation:

• Mitochondria are involved in the intrinsic pathway of apoptosis (programmed cell

death) by releasing cytochrome c, which activates caspases that dismantle the cell.

Calcium Homeostasis:

• Mitochondria help regulate intracellular calcium levels, crucial for signal transduction,
muscle contraction, and other cellular activities.

Reactive Oxygen Species (ROS) Production:

• Mitochondria are a source of ROS, which can act as signaling molecules but also cause
oxidative damage if not properly regulated.
 105

Lysosomes: Diverse Roles Due to Hydrolytic Activity of Enzymes

Lysosomes are membrane-bound organelles in eukaryotic cells that contain a wide variety of hydrolytic
enzymes capable of breaking down different types of biomolecules. These enzymes include proteases,
lipases, nucleases, and glycosidases, all of which function optimally at an acidic pH. Lysosomes are
often referred to as the cell's "digestive system" because of their crucial role in degrading and recycling
cellular waste, macromolecules, and foreign substances.
Structure of Lysosomes
• Membrane: A single phospholipid bilayer that separates the acidic interior of the lysosome
from the cytoplasm.
• Enzymes: Contains over 50 different hydrolytic enzymes.
• Acidic Environment: Maintained by proton pumps (H+-ATPases) that actively transport
hydrogen ions (protons) into the lysosome.

Diverse Roles of Lysosomes

1. Intracellular Digestion:
• Autophagy: Lysosomes degrade and recycle damaged or obsolete cellular
components. During autophagy, portions of the cytoplasm and organelles are enclosed
in autophagosomes, which then fuse with lysosomes, where their contents are digested.
• Heterophagy: Involves the digestion of extracellular material taken up by the cell via
endocytosis or phagocytosis. Lysosomes fuse with endosomes or phagosomes, forming
endolysosomes or phagolysosomes, where the ingested material is degraded.
2. Pathogen Destruction:
• Phagocytosis: Lysosomes play a crucial role in the immune response by destroying
pathogens. Phagocytic cells, such as macrophages and neutrophils, engulf pathogens
into phagosomes, which then fuse with lysosomes to form phagolysosomes where
pathogens are digested by hydrolytic enzymes.
3. Apoptosis:
 106

• Lysosomes can contribute to programmed cell death (apoptosis) by releasing their

enzymes into the cytoplasm, leading to cellular breakdown and death.
4. Membrane Repair:
• Lysosomes are involved in repairing the plasma membrane. When the membrane is
damaged, lysosomes
can fuse with the
membrane to patch
up the lesion.
5. Metabolic
Regulation:
• Lysosomes
participate in
nutrient sensing and
metabolic regulation
through the mTOR
signaling pathway.
They help regulate
cellular metabolism
by responding to the
availability of
nutrients and growth
factors.
6. Secretion:
• Some
specialized cells use
lysosome-related organelles for the regulated secretion of their contents. For example,
in immune cells like cytotoxic T cells and natural killer (NK) cells, lysosomes contain
cytotoxic molecules that are secreted to kill target cells.
Hydrolytic Enzymes in Lysosomes
• Proteases: Break down proteins into amino acids.
• Lipases: Degrade lipids into fatty acids and glycerol.
• Nucleases: Digest nucleic acids into nucleotides.
• Glycosidases: Hydrolyze carbohydrates into simple sugars.
Diseases Related to Lysosomal Dysfunction
Defects in lysosomal enzymes or in the transport of enzymes to lysosomes can lead to lysosomal storage
diseases (LSDs), which are characterized by the accumulation of undigested substrates. Examples
include:
• Tay-Sachs Disease: Caused by a deficiency in the enzyme hexosaminidase A, leading to the
accumulation of GM2 ganglioside in nerve cells.
• Gaucher Disease: Caused by a deficiency in the enzyme glucocerebrosidase, leading to the
accumulation of glucocerebroside.
• Pompe Disease: Caused by a deficiency in the enzyme acid alpha-glucosidase, leading to the
accumulation of glycogen in cells.
 107

Peroxisomes: Metabolism of Hydrogen Peroxide

Peroxisomes are membrane-bound organelles found in eukaryotic cells that are involved in various
metabolic processes, including the metabolism of hydrogen peroxide (H2O2). They contain several
enzymes that participate in the breakdown of H2O2, a toxic byproduct of various cellular reactions, into
water and oxygen. Peroxisomes play a crucial role in maintaining cellular redox balance and protecting
cells from oxidative damage.

Enzymes Involved in Hydrogen Peroxide Metabolism

1. Catalase:
• Catalase is the key enzyme responsible for the breakdown of hydrogen peroxide into
water and oxygen.
• It catalyzes the reaction:
2 H2O2 → 2 H2O + O2
• Catalase achieves this by facilitating the decomposition of hydrogen peroxide into
water and molecular oxygen, effectively neutralizing its toxicity.
• Peroxisomes contain a high concentration of catalase, making them the primary site for
hydrogen peroxide metabolism in the cell.
2. Peroxidases:
• Peroxisomes also contain peroxidases, such as peroxiredoxins, which can reduce
hydrogen peroxide and other organic peroxides using reducing equivalents from
molecules like NADPH.
• These peroxidases play a complementary role to catalase in detoxifying hydrogen
peroxide and other reactive oxygen species (ROS) in the cell.

Importance of Hydrogen Peroxide Metabolism

1. Protection Against Oxidative Stress:
• Hydrogen peroxide is a reactive oxygen species (ROS) generated as a byproduct of
various metabolic reactions, including cellular respiration.
• If not neutralized, hydrogen peroxide can damage cellular components such as DNA,
proteins, and lipids through oxidative stress, leading to cell dysfunction and death.
• Peroxisomes help protect cells from oxidative damage by efficiently breaking down
hydrogen peroxide into harmless water and oxygen.
2. Regulation of Cellular Redox Balance:
• Peroxisomes contribute to the regulation of cellular redox balance by controlling the
levels of hydrogen peroxide and other ROS.
• They participate in maintaining an optimal cellular environment for metabolic reactions
and signaling pathways by preventing ROS accumulation.
3. Cellular Signaling:
• Reactive oxygen species, including hydrogen peroxide, can also act as signaling
molecules involved in various cellular processes, such as apoptosis, inflammation, and
cell proliferation.
 108

• Peroxisomes help regulate the levels of hydrogen peroxide, thereby influencing cellular
signaling pathways and responses to stress and environmental stimuli.

Disorders Related to Peroxisomal Dysfunction

• Peroxisomal Biogenesis Disorders (PBDs):
• PBDs are a group of genetic disorders characterized by defects in peroxisome
formation and function.
• Examples include Zellweger syndrome, neonatal adrenoleukodystrophy, and infantile
Refsum disease.
• These disorders result in the accumulation of toxic substances and impaired
metabolism of various molecules, leading to severe neurological, developmental, and
metabolic abnormalities.

Glyoxysomes
Glyoxysomes are specialized organelles found in plants and some microorganisms, particularly in
germinating seeds, where they play a crucial role in lipid metabolism. They are essentially specialized
peroxisomes with additional enzymes required for the glyoxylate cycle, a variant of the Krebs cycle.
The glyoxylate cycle enables plants to convert fatty acids into carbohydrates, which are essential for
seedling growth before photosynthesis begins. Glyoxysomes contain enzymes such as isocitrate lyase
and malate synthase, which are absent in typical peroxisomes but are necessary for converting fatty
acids into sugars.

Plasma Membrane and Its Functions

The plasma membrane, also known as the cell membrane, is a phospholipid bilayer that surrounds the
cytoplasm of a cell, separating it from the extracellular environment. The plasma membrane is a
dynamic structure that performs several essential functions for the cell:
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1. Cellular Barrier:
• The plasma membrane acts as a semi-permeable barrier, regulating the movement of
substances into and out of the cell. It controls the passage of ions, nutrients, waste
products, and signaling molecules.
2. Cellular Communication:
• The plasma membrane contains proteins, such as receptors and channels, that facilitate
communication between the cell and its environment. These proteins allow the cell to
detect and respond to external signals, such as hormones, neurotransmitters, and
environmental cues.
3. Cell Adhesion:
• Specialized proteins in the plasma membrane, such as integrins and cadherins, mediate
cell adhesion and interactions with neighboring cells and the extracellular matrix. Cell
adhesion is crucial for maintaining tissue structure and integrity.
4. Cellular Recognition:
• Carbohydrate molecules, often attached to proteins and lipids on the extracellular
surface of the plasma membrane, serve as markers for cell recognition and
identification. These markers are involved in processes such as immune responses,
tissue development, and cell-cell signaling.
5. Transport:
• The plasma membrane contains various protein channels, carriers, and pumps that
regulate the movement of ions, nutrients, and other molecules across the membrane.
This transport is essential for maintaining cellular homeostasis and supporting cellular
processes such as nutrient uptake, ion exchange, and waste removal.
6. Cellular Signaling:
• The plasma membrane is involved in signal transduction, whereby extracellular signals
are detected by receptors on the cell surface and transmitted to the cell interior, leading
to cellular responses. This signaling regulates processes such as cell growth,
differentiation, and survival.
7. Electrical Properties:
• The plasma membrane maintains a resting membrane potential, which is essential for
the generation and propagation of electrical signals in excitable cells such as neurons
and muscle cells.

The plasma membrane

The plasma membrane, also known as the cell membrane, is a dynamic and selectively permeable
barrier that surrounds the cytoplasm of a cell, separating it from the extracellular environment. It is
composed of a complex arrangement of lipids, proteins, and carbohydrates that form a fluid mosaic
structure.

Chemical Composition of the Plasma Membrane

1. Phospholipids:
• Phospholipids are the primary structural components of the plasma membrane. They
consist of a hydrophilic ("water-loving") phosphate head and two hydrophobic ("water-
fearing") fatty acid tails.
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• The arrangement of phospholipids in a bilayer forms the basic structure of the

membrane, with the hydrophobic tails facing inward and the hydrophilic heads facing
outward, towards the aqueous environments inside and outside the cell.
2. Cholesterol:
• Cholesterol molecules are interspersed within the phospholipid bilayer of the plasma
membrane.
• Cholesterol helps maintain the fluidity and stability of the membrane by preventing
phospholipids from packing too closely together or becoming too fluid.
3. Proteins:
• Integral Proteins: Embedded within the lipid bilayer, integral proteins traverse the
membrane and may span from one side to the other. They can serve as channels,
carriers, receptors, or enzymes.
• Peripheral Proteins: Found on the inner or outer surface of the membrane, peripheral
proteins are not embedded within the lipid bilayer. They often interact with integral
proteins or the cytoskeleton to help maintain the membrane's structure and function.
4. Carbohydrates:
• Carbohydrates are attached to proteins (glycoproteins) or lipids (glycolipids) on the
extracellular surface of the plasma membrane.
• These carbohydrate chains, also known as glycocalyx, play roles in cell-cell
recognition, adhesion, and communication.

Structure of the Plasma Membrane

• Fluid Mosaic Model:
• The plasma membrane is described by the fluid mosaic model, which depicts the
membrane as a fluid, dynamic structure composed of a mosaic of lipids, proteins, and
carbohydrates.
• The fluidity of the membrane allows for the lateral movement of phospholipids and
proteins within the bilayer, enabling various cellular processes such as membrane
trafficking, signal transduction, and cell-cell interactions.
• Lipid Bilayer:
• The plasma membrane consists of two layers of phospholipids arranged with their
hydrophobic tails facing inward and their hydrophilic heads facing outward.
• This lipid bilayer provides the basic structure of the membrane and serves as a barrier
to the passage of hydrophilic molecules and ions.
• Protein Distribution:
• Integral proteins are embedded within the lipid bilayer, with some spanning the entire
membrane (transmembrane proteins) and others extending partially into the
hydrophobic core.
• Peripheral proteins are found on the inner or outer surface of the membrane and interact
with integral proteins or the cytoskeleton.
• Asymmetry:
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• The distribution of lipids and proteins across the plasma membrane is asymmetric, with
different compositions on the inner and outer leaflets.
• This asymmetry is essential for various cellular processes, including membrane
trafficking, cell signaling, and cell recognition.

Cell Permeability
Cell permeability refers to the ability of molecules to pass through the cell membrane. The plasma
membrane is selectively permeable, meaning it allows certain substances to pass through while
restricting others. The permeability of the membrane depends on factors such as the size, charge, and
lipid solubility of the molecules.
Passive Transport: In passive transport, molecules move across the cell membrane without the
expenditure of energy by the cell. This includes processes such as:
Diffusion: The movement of molecules from an area of higher concentration to an area of lower
concentration until equilibrium is reached.
Facilitated Diffusion: The passive movement of molecules across the membrane facilitated by
transport proteins, such as channels and carriers.
Osmosis: The diffusion of water across a selectively permeable membrane from an area of lower solute
concentration to an area of higher solute concentration.
Active Transport: Active transport requires the expenditure of energy by the cell to move molecules
against their concentration gradient, from an area of lower concentration to an area of higher
concentration. This process is typically mediated by transport proteins called pumps, such as the
sodium-potassium pump.

Endocytosis
Endocytosis is a cellular process by which cells engulf extracellular materials by forming vesicles
derived from the plasma membrane. There are several types of endocytosis:
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Phagocytosis: In phagocytosis, large particles, such as bacteria, cell debris, or other solid materials, are
engulfed by the cell. Phagocytosis is primarily performed by specialized cells such as macrophages and
neutrophils as part of the immune response.
Pinocytosis: Also known as "cell drinking," pinocytosis involves the non-selective uptake of small
dissolved molecules and extracellular fluid into the cell. It is a continuous process that occurs in most
cells to sample the extracellular environment and regulate the composition of the cell's internal
environment.
Receptor-Mediated Endocytosis: This process involves the selective uptake of specific molecules
bound to receptor proteins on the cell surface. Ligands bind to cell surface receptors, triggering the
formation of clathrin-coated vesicles that are internalized by the cell.

Phagocytosis
Phagocytosis is a specialized form of endocytosis in which cells engulf large particles or
microorganisms into vesicles called phagosomes. Phagocytosis is primarily performed by specialized
cells called phagocytes, including macrophages, neutrophils, and dendritic cells. The process of
phagocytosis involves several steps:
Recognition and Binding: The phagocyte recognizes and binds to the particle or microorganism
through receptors on its surface.
Engulfment: The plasma membrane extends outward to surround the particle, forming a phagosome.
Phagosome Formation: The phagosome, containing the engulfed particle, is internalized within the
cell.
Phagosome Maturation: The phagosome fuses with lysosomes, forming a phagolysosome. The
lysosomal enzymes within the phagolysosome digest the particle, breaking it down into smaller
components.
Digestion and Degradation: The particle is broken down into its constituent molecules, which can
then be utilized by the cell for energy or other cellular processes.
Exocytosis: After digestion is complete, residual materials are expelled from the cell through
exocytosis.

cytoskeleton
The cytoskeleton is a dynamic network of protein filaments found in the cytoplasm of eukaryotic cells.
It provides structural support, maintains cell shape, facilitates cell movement, and is involved in various
cellular processes such as cell division and intracellular transport. The cytoskeleton is composed of
three main types of protein filaments: microfilaments (actin filaments), microtubules, and intermediate
filaments.

Microfilaments (Actin Filaments)

• Composition: Microfilaments are thin, flexible filaments composed of actin protein subunits
arranged in a double helix structure.
• Function:
• Microfilaments play a crucial role in cell motility, including cell crawling, contraction,
and cytokinesis (cell division).
• They are involved in maintaining cell shape and providing mechanical support to the
cell's plasma membrane.
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• Microfilaments are also important in intracellular transport processes, such as the

movement of organelles and vesicles along actin filaments.
• Examples of Cellular Structures: Microvilli in epithelial cells, contractile ring during
cytokinesis, muscle contraction in muscle cells.

Microtubules
• Composition: Microtubules are hollow, cylindrical structures composed of tubulin protein
subunits arranged in a protofilament structure.
• Function:
• Microtubules serve as tracks for intracellular transport, facilitating the movement of
organelles, vesicles, and molecular motors within the cell.
• They are essential for maintaining cell shape and organization, as well as for
positioning organelles within the cell.
• Microtubules are involved in cell division, forming the mitotic spindle that separates
chromosomes during mitosis and meiosis.
• Examples of Cellular Structures: Mitotic spindle during cell division, cilia and flagella for
cell motility, centrioles and basal bodies.

Intermediate Filaments
• Composition: Intermediate filaments are fibrous proteins that form ropelike structures
composed of various types of proteins, such as keratins, vimentins, and neurofilaments.
• Function:
• Intermediate filaments provide mechanical strength and structural support to cells,
particularly in tissues subjected to mechanical stress.
• They contribute to the maintenance of cell shape and integrity, as well as the anchorage
of organelles within the cytoplasm.
• Intermediate filaments are
important for cell-cell
adhesion and tissue
organization.
• Examples of Cellular Structures:
Desmosomes, nuclear lamina,
keratin filaments in epithelial cells.

Comparison

Microfilaments (Actin
Feature Filaments) Microtubules Intermediate Filaments

Composition Actin protein subunits Tubulin protein subunits Various fibrous proteins

Structure Thin, flexible filaments Hollow, cylindrical tubes Ropelike structures

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Cell motility, shape

maintenance, intracellular Intracellular transport, cell Mechanical strength, structural
Function transport division, shape maintenance support, cell-cell adhesion

Microvilli, contractile ring, Desmosomes, nuclear lamina,

Examples muscle contraction Mitotic spindle, cilia, flagella keratin filaments

Replication
Definition: Replication is the process by which DNA molecules make exact copies of themselves.

Key Steps:
1. Initiation: DNA replication begins at specific sites called origins of replication, where the
DNA double helix unwinds and separates into two strands.

2. Elongation: Enzymes called DNA polymerases add complementary nucleotides to each of the
original DNA strands, synthesizing two new daughter strands.

3. Termination: Replication proceeds bidirectionally along the DNA strands until the entire
molecule is copied. The process ends when replication forks from neighboring origins meet or
reach the end of the DNA molecule.

Significance: DNA replication ensures that genetic information is faithfully transmitted from parent to
daughter cells during cell division. It is essential for growth, development, and inheritance in all living
organisms.

Transcription
Definition: Transcription is the process by which the information encoded in DNA is copied into RNA
molecules.

Key Steps:
1. Initiation: RNA polymerase binds to a specific DNA sequence called the promoter region,
marking the beginning of transcription.

2. Elongation: RNA polymerase moves along the DNA template strand, synthesizing a
complementary RNA strand by adding ribonucleotides according to the base-pairing rules (A-
U and G-C).

3. Termination: Transcription ends when RNA polymerase reaches a termination signal in the
DNA sequence, causing RNA polymerase to dissociate from the DNA and release the newly
synthesized RNA molecule.

Significance: Transcription is the first step in gene expression, where the genetic information stored in
DNA is converted into functional RNA molecules such as messenger RNA (mRNA), transfer RNA
(tRNA), and ribosomal RNA (rRNA). These RNA molecules play crucial roles in protein synthesis and
various cellular processes.

Translation
Definition: Translation is the process by which the information encoded in RNA molecules is used to
synthesize proteins.
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Key Steps:
1. Initiation: The small ribosomal subunit binds to the mRNA molecule at a specific sequence
called the start codon (usually AUG), and the initiator tRNA carrying methionine binds to the
start codon.

2. Elongation: The ribosome moves along the mRNA molecule in a 5' to 3' direction, catalyzing
the formation of peptide bonds between amino acids carried by tRNA molecules in the A site
and P site. The ribosome then translocates to the next codon, shifting the tRNA molecules to
the P and E sites.

3. Termination: Translation ends when a stop codon (UAA, UAG, or UGA) is encountered on
the mRNA molecule. Release factors bind to the stop codon, causing the ribosome to release
the newly synthesized polypeptide chain and dissociate from the mRNA.

Significance: Translation is essential for protein synthesis, where the sequence of nucleotides in mRNA
is translated into the sequence of amino acids in a protein. Proteins are the functional molecules that
carry out most cellular processes, including enzymatic reactions, structural support, and cell signaling.