Insect Physiology

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Insect Physiology

• Form and Function of Systems

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Outline

• Digestive
• Excretory
• Circulatory
• Respiratory
• Endocrine

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An insect uses its digestive system to extract
nutrients and other substances from the food it
consumes.   Most of this food is ingested in the
form of macromolecules and other complex
substances (such as proteins, polysaccharides,
fats, nucleic acids, etc.) which must be broken
down by catabolic reactions into smaller
molecules (i.e. amino acids, simple sugars, etc.)
before being used by cells of the body for
energy, growth, or reproduction.   This
break-down process is known as digestion.
                                                  
                              
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All insects have a complete digestive system.  
This means that food processing occurs within a
tube-like enclosure, the alimentary canal,
running lengthwise through the body from mouth to
anus. Ingested food usually travels in only one
direction.   This arrangement differs from an
incomplete digestive system (found in certain
lower invertebrates like hydra and starfish)
where a single opening to a pouch-like cavity
serves as both mouth and anus.   Most biologists
regard a complete digestive system as an
evolutionary improvement over an incomplete
digestive system because it permits functional
specialization -- different parts of the system
may be specially adapted for various functions of
food digestion, nutrient absorption, and waste
excretion.   In most insects, the alimentary
canal is subdivided into three functional
regions   foregut (stomodeum), midgut
(mesenteron), and hindgut (proctodeum).
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Stomodeum
In addition to the alimentary canal, insects also
have paired salivary glands and salivary
reservoirs. These structures usually reside in
the thorax (adjacent to the foregut).   Salivary
ducts lead from the glands to the reservoirs and
then forward, through the head, to an opening
(the salivarium) behind the hypopharynx.  
Movements of the mouthparts help mix saliva with
food in the buccal cavity.
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An insect's mouth, located centrally at the base
of the mouthparts, is a muscular valve
(sphincter) that marks the "front" of the
foregut.   Food in the buccal cavity is sucked
through the mouth opening and into the pharynx by
contractile action of cibarial muscles.   These
muscles, located between the head capsule and the
anterior wall of the pharynx, create suction by
enlarging the volume of the pharynx (like opening
a bellows).   This "suction pump" mechanism is
called the cibarial pump.   It is especially
well-developed in insects with piercing/sucking
mouthparts.                                     
                                            
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From the pharynx, food passes into the esophagus
by means of peristalsis (rhythmic muscular
contractions of the gut wall).   The esophagus is
just a simple tube that connects the pharynx to
the crop, a food-storage organ.   Food remains in
the crop until it can be processed through the
remaining sections of the alimentary canal.  
While in the crop, some digestion may occur as a
result of salivary enzymes that were added in the
buccal cavity and/or other enzymes regurgitated
from the midgut. In some insects, the crop
opens posteriorly into a muscular proventriculus.
  This organ contains tooth-like denticles that
grind and pulverize food particles.   The
proventriculus serves much the same function as a
gizzard in birds.   The stomodeal valve, a
sphincter muscle located just behind the
proventriculus, regulates the flow of food from
the stomodeum to the mesenteron.
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In a developing embryo, the foregut arises as a
simple invagination of the anterior body wall  
this means that all of its tissues and organs are
derived from embryonic ectoderm.   In effect, the
inside of the stomodeum is continuous with the
outside of the insect's body.   Since exoskeleton
is secreted to protect the insect externally, it
is not surprising to find that cells lining the
foregut produce a similar structure (known as the
intima) to protect themselves from abrasion by
food particles.   The hard denticles inside the
proventriculus are made from this same material.
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Mesenteron
The midgut begins just past the stomodeal valve.
  Near its anterior end, finger-like projections
(usually from 2 to 10) diverge from the walls of
the midgut.   These structures, the gastric
caecae, provide extra surface area for secretion
of enzymes or absorption of water (and other
substances) from the alimentary canal.   The rest
of the midgut is called the ventriculus -- it is
the primary site for enzymatic digestion of food
and absorption of nutrients.   Digestive cells
lining the walls of the ventriculus have
microscopic projections (microvilli) that
increase surface area for nutrient absorption.
                                                  
                              
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The midgut is derived from embryonic endoderm so
it is not protected by an intima.   Instead, the
midgut is lined with a semipermeable membrane
secreted by a cluster of cells (the cardial
epithelium) that lie just behind the stomodeal
valve.   This peritrophic membrane consists of
chitin fibrils embedded in a protein-carbohydrate
matrix.   It protects the delicate digestive
cells without inhibiting absorption of nutrient
molecules. The posterior end of the midgut is
marked by another sphincter muscle, the pyloric
valve.   It regulates the flow of material from
the mesenteron to the proctodeum.
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Proctodeum
The pyloric valve serves as a point of origin for
dozens to hundreds of Malpighian tubules.   These
long, spaghetti-like structures extend throughout
most of the abdominal cavity where they serve as
excretory organs, removing nitrogenous wastes
(principally ammonium ions, NH4) from the
hemolymph.   The toxic NH4 is quickly converted
to urea and then to uric acid by a series of
chemical reactions within the Malpighian tubules.
  The uric acid, a semi-solid, accumulates inside
each tubule and is eventually emptied into the
hindgut for elimination as part of the fecal
pellet.                                         
                                        
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The rest of the hindgut plays a major role in
homeostasis by regulating the absorption of water
and salts from waste products in the alimentary
canal.   In some insects, the hindgut is visibly
subdivided into an ileum, a colon, and a rectum.
  Efficient recovery of water is facilitated by
six rectal pads that are embedded in the walls of
the rectum.   These organs remove more than 90
of the water from a fecal pellet before it passes
out of the body through the anus. Embryonically,
the hindgut develops as an invagination of the
body wall (from ectodermal tissue). Just like the
foregut, it is lined with a thin, protective
layer of cuticle (intima) that is secreted by the
endothelial cells of the gut wall. When an insect
molts, it sheds and replaces the intima in both
the foregut and the hindgut.
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Circulatory System
Insects, like all other arthropods, have an
open circulatory system which differs in both
structure and function from the closed
circulatory system found in humans and other
vertebrates.   In a closed system, blood is
always contained within vessels (arteries, veins,
capillaries, or the heart itself).   In an open
system, blood (usually called hemolymph) spends
much of its time flowing freely within body
cavities where it makes direct contact with all
internal tissues and organs.
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The circulatory system is responsible for
movement of nutrients, salts, hormones, and
metabolic wastes throughout the insect's body.  
In addition, it plays several critical roles in
defense   it seals off wounds through a clotting
reaction, it encapsulates and destroys internal
parasites or other invaders, and in some species,
it produces (or sequesters) distasteful compounds
that provide a degree of protection against
predators.   The hydraulic (liquid) properties of
blood are important as well.   Hydrostatic
pressure generated internally by muscle
contraction is used to facilitate hatching,
molting, expansion of body and wings after
molting, physical movements (especially in
soft-bodied larvae), reproduction (e.g.
insemination and oviposition), and evagination of
certain types of exocrine glands.   In some
insects, the blood aids in thermoregulation   it
can help cool the body by conducting excess heat
away from active flight muscles or it can warm
the body by collecting and circulating heat
absorbed while basking in the sun.
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A dorsal vessel is the major structural component
of an insect's circulatory system.   This tube
runs longitudinally through the thorax and
abdomen, along the inside of the dorsal body
wall.   In most insects, it is a fragile,
membranous structure that collects hemolymph in
the abdomen and conducts it forward to the head.
                                                 
                           In the abdomen, the
dorsal vessel is called the heart.   It is
divided segmentally into chambers that are
separated by valves (ostia) to ensure one-way
flow of hemolymph.   A pair of alary muscles are
attached laterally to the walls of each chamber.
  Peristaltic contractions of the these muscles
force the hemolymph forward from chamber to
chamber.   During each diastolic phase
(relaxation), the ostia open to allow inflow of
hemolymph from the body cavity.   The heart's
contraction rate varies considerably from species
to species -- typically in the range of 30 to 200
beats per minute.   The rate tends to fall as
ambient temperature drops and rise as temperature
(or the insect's level of activity) increases.
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                                        In
front of the heart, the dorsal vessel lacks
valves or musculature.   It is a simple tube
(called the aorta) which continues forward to the
head and empties near the brain.   Hemolymph
bathes the organs and muscles of the head as it
emerges from the aorta, and then haphazardly
percolates back over the alimentary canal and
through the body until it reaches the abdomen and
re-enters the heart. To facilitate circulation
of hemolymph, the body cavity is divided into
three compartments (called blood sinuses) by two
thin sheets of muscle and/or membrane known as
the dorsal and ventral diaphragms.   The dorsal
diaphragm is formed by alary muscles of the heart
and related structures it separates the
pericardial sinus from the perivisceral sinus.  
The ventral diaphragm usually covers the nerve
cord it separates the perivisceral sinus from
the perineural sinus.
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In some insects, pulsatile organs are located
near the base of the wings or legs.   These
muscular "pumps" do not usually contract on a
regular basis, but they act in conjunction with
certain body movements to force hemolymph out
into the extremities. About 90 of insect
hemolymph is plasma   a watery fluid -- usually
clear, but sometimes greenish or yellowish in
color.   Compared to vertebrate blood, it
contains relatively high concentrations of amino
acids, proteins, sugars, and inorganic ions.  
Overwintering insects often sequester enough
ribulose, trehalose, or glycerol in the plasma to
prevent it from freezing during the coldest
winters.   The remaining 10 of hemolymph volume
is made up of various cell types (collectively
known as hemocytes) they are involved in the
clotting reaction, phagocytosis, and/or
encapsulation of foreign bodies.   The density of
insect hemocytes can fluctuate from less than
25,000 to more than 100,000 per cubic millimeter,
but this is significantly fewer than the 5
million red blood cells, 300,000 platelets, and
7000 white blood cells found in the same volume
of human blood.   With the exception of a few
aquatic midges, insect hemolymph does NOT contain
hemoglobin (or red blood cells).   Oxygen is
delivered by the tracheal system, not the
circulatory system.
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Respiratory System
The respiratory system is responsible for
delivering sufficient oxygen to all cells of the
body and for removing carbon dioxide (CO2) that
is produced as a waste product of cellular
respiration.   The respiratory system of insects
(and many other arthropods) is separate from the
circulatory system.   It is a complex network of
tubes (called a tracheal system) that delivers
oxygen-containing air to every cell of the body.
                                              
                              Air enters the
insect's body through valve-like openings in the
exoskeleton.   These openings (called spiracles)
are located laterally along the thorax and
abdomen of most insects -- usually one pair of
spiracles per body segment.   Air flow is
regulated by small muscles that operate one or
two flap-like valves within each spiracle --
contracting to close the spiracle, or relaxing to
open it.
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After passing through a spiracle, air enters a
longitudinal tracheal trunk, eventually diffusing
throughout a complex, branching network of
tracheal tubes that subdivides into smaller and
smaller diameters and reaches every part of the
body.   At the end of each tracheal branch, a
special cell (the tracheole) provides a thin,
moist interface for the exchange of gasses
between atmospheric air and a living cell.  
Oxygen in the tracheal tube first dissolves in
the liquid of the tracheole and then diffuses
into the cytoplasm of an adjacent cell.   At the
same time, carbon dioxide, produced as a waste
product of cellular respiration, diffuses out of
the cell and, eventually, out of the body through
the tracheal system. Each tracheal tube
develops as an invagination of the ectoderm
during embryonic development.   To prevent its
collapse under pressure, a thin, reinforcing
"wire" of cuticle (the taenidia) winds spirally
through the membranous wall.   This design
(similar in structure to a heater hose on an
automobile or an exhaust duct on a clothes dryer)
gives tracheal tubes the ability to flex and
stretch without developing kinks that might
restrict air flow.
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                                        The
absence of taenidia in certain parts of the
tracheal system allows the formation of
collapsible air sacs, balloon-like structures
that may store a reserve of air.   In dry
terrestrial environments, this temporary air
supply allows an insect to conserve water by
closing its spiracles during periods of high
evaporative stress.   Aquatic insects consume the
stored air while under water or use it to
regulate buoyancy.   During a molt, air sacs fill
and enlarge as the insect breaks free of the old
exoskeleton and expands a new one.   Between
molts, the air sacs provide room for new growth
-- shrinking in volume as they are compressed by
expansion of internal organs.
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Small insects rely almost exclusively on passive
diffusion and physical activity for the movement
of gasses within the tracheal system.   However,
larger insects may require active ventilation of
the tracheal system (especially when active or
under heat stress).   They accomplish this by
opening some spiracles and closing others while
using abdominal muscles to alternately expand and
contract body volume.   Although these pulsating
movements flush air from one end of the body to
the other through the longitudinal tracheal
trunks, diffusion is still important for
distributing oxygen to individual cells through
the network of smaller tracheal tubes.   In fact,
the rate of gas diffusion is regarded as one of
the main limiting factors (along with weight of
the exoskeleton) that prevents real insects from
growing as large as the ones we see in horror
movies!
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Endocrine System
A hormone is a chemical signal sent from cells in
one part of an organism to cells in another part
(or parts) of the same individual.   They are
often regarded as chemical messengers. Although
typically produced in very small quantities,
hormones may cause profound changes in their
target cells.   Their effect may be stimulatory
or inhibitory.   In some cases, a single hormone
may have multiple targets and cause different
effects in each target.   There are at least four
categories of hormone-producing cells in an
insect's body
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1) Endocrine glands -- secretory structures
adapted exclusively for producing hormones and
releasing them into the circulatory system. 2)
Neurohemal organs -- similar to glands, but they
store their secretory product in a special
chamber until stimulated to release it by a
signal from the nervous system (or another
hormone). 3) Neurosecretory cells --
specialized nerve cells (neurons) that respond to
stimulation by producing and secreting specific
chemical messengers.   Functionally, they serve
as a link between the nervous system and the
endocrine system 4) Internal organs --
hormone-producing cells are associated with
numerous organs of the body, including the
ovaries and testes, the fat body, and parts of
the digestive system. Together, these
hormone-secreting structures form an endocrine
system that helps maintain homeostasis,
coordinate behavior, and regulate growth,
development, and other physiological activities.
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In insects, the largest and most obvious
endocrine glands are found in the prothorax, just
behind the head.   These prothoracic glands
manufacture ecdysteroids, a group of
closely-related steroid hormones (including
ecdysone) that stimulate synthesis of chitin and
protein in epidermal cells and trigger a cascade
of physiological events that culminates in
molting.   For this reason, the ecdysteroids are
often called "molting hormones".   Once an insect
reaches the adult stage, its prothoracic glands
atrophy (wither away) and it will never molt
again. Prothoracic glands produce and release
ecdysteroids only after they have been stimulated
by another chemical messenger, prothoracicotropic
hormone (PTTH for short).   This compound is a
peptide hormone secreted by the corpora cardiaca,
a pair of neurohemal organs located on the walls
of the aorta just behind the brain.   The corpora
cardiaca release their store of PTTH only after
they receive a signal from neurosecretory cells
in the brain.   In a sense, they act as signal
amplifiers -- sending out a big pulse of hormone
to the body in response to a small message from
the brain.                                        
                        
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The corpora allata, another pair of neurohemal
organs, lie just behind the corpora cardiaca.  
They manufacture juvenile hormone (JH for short),
a compound that inhibits development of adult
characteristics during the immature stages and
promotes sexual maturity during the adult stage.
  Neurosecretory cells in the brain regulate
activity of the corpora allata -- stimulating
them to produce JH during larval or nymphal
instars, inhibiting them during the transition to
adulthood, and reactivating them once the adult
is ready for reproduction.   The chemical
structure of juvenile hormone is rather unusual
  it is a sesquiterpene compound -- more similar
to defensive chemicals found in pine trees than
to any other animal hormone.
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The neurosecretory cells are found in clusters,
both medially and laterally in the insect's
brain.   Axons from these cells can be traced
along tiny nerves that run to the corpora
cardiaca and corpora allata.   The cells produce
and secrete brain hormone, a low-molecular-weight
peptide that appears to be the same as (or very
similar to) prothoracicotropic hormone (PTTH)
manufactured by the corpora cardiaca.   Insect
physiologists suspect that brain hormone is bound
to a larger carrier protein while it is inside
the neurosecretory cell, and some believe that
each cluster of cells may produce as many as
three different brain hormones (or
hormone-carrier combinations).   Large numbers of
neurosecretory cells also occur in the ventral
ganglia of the nerve cord, but their function is
unknown. Many other tissues and organs of the
body also produce hormones.   Ovaries and testes,
for example, produce gonadal hormones that have
been shown to coordinate courtship and mating
behaviors.   Ventral ganglia in the nervous
system produce one compound (eclosion hormone)
that helps an insect shed its old exoskeleton and
another compound (bursicon) that causes hardening
and tanning of the new one.   There are still
other hormones that control the level of sugar
dissolved in the blood, adjust salt and water
balance, and regulate protein metabolism.
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When an immature insect has grown sufficiently to
require a larger exoskeleton, sensory input from
the body activates certain neurosecretory cells
in the brain.   These neurons respond by
secreting brain hormone which triggers the
corpora cardiaca to release their store of
prothoracicotropic hormone (PTTH) into the
circulatory system.   This sudden "pulse" of PTTH
stimulates the prothoracic glands to secrete
molting hormone (ecdysteroids).
(                                                 
           
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Molting hormone affects many cells throughout the
body, but its principle function is to stimulate
a series of physiological events (collectively
known as apolysis) that lead to synthesis of a
new exoskeleton.   During this process, the new
exoskeleton forms as a soft, wrinkled layer
underneath the hard parts (exocuticle plus
epicuticle) of the old exoskeleton.   The
duration of apolysis ranges from days to weeks,
depending on the species and its characteristic
growth rate.   Once new exoskeleton has formed,
the insect is ready to shed what's left of its
old exoskeleton.   At this stage, the insect is
said to be pharate, meaning that the body is
covered by two layers of exoskeleton.
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As long as ecdysteroid levels remain above a
critical threshold in the hemolymph, other
endocrine structures remain inactive (inhibited).
But toward the end of apolysis, ecdysteroid
concentration falls, and neurosecretory cells in
the ventral ganglia begin secreting eclosion
hormone. This hormone triggers ecdysis, the
physical process of shedding the old exoskeleton.
In addition, a rising concentration of eclosion
hormone stimulates other neurosecretory cells in
the ventral ganglia to secrete bursicon, a
hormone that causes hardening and darkening of
the integument (tanning) due to the formation of
quinone cross-linkages in the exocuticle
(sclerotization).
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In immature insects, juvenile hormone is
secreted by the corpora allata prior to each
molt.   This hormone inhibits the genes that
promote development of adult characteristics
(e.g. wings, reproductive organs, and external
genitalia), causing the insect to remain
"immature" (nymph or larva).   The corpora allata
become atrophied (shrink) during the last larval
or nymphal instar and stop producing juvenile
hormone.   This releases inhibition on
development of adult structures and causes the
insect to molt into an adult (hemimetabolous) or
a pupa (holometabolous). At the approach of
sexual maturity in the adult stage, brain
neurosecretory cells release a brain hormone that
"reactivates" the corpora allata, stimulating
renewed production of juvenile hormone.   In
adult females, juvenile hormone stimulates
production of yolk for the eggs.   In adult
males, it stimulates the accessory glands to
produce proteins needed for seminal fluid and the
case of the spermatophore.   In the absence of
normal juvenile hormone production, the adult
remains sexually sterile.