CROP PROTECTION 44

GENERAL PHYSIOLOGY AND TOXICOLOGY

INSECT
PHYSIOLOGY
THE DIGESTIVE SYSTEM
(ALIMENTARY SYSTEM)

RYAN CHRISTIAN B. GUIRITAN

Faculty In-charge, CPRT 44
Department of Entomology
DIGESTIVE SYSTEM (Alimentary System)
• Alimentary Canal – a tube where the insects possess, which
extends from the anterior oral opening, the mouth, to the posterior
anus.
– Functions:
• triturate
• lubricate
• store food material and expel the
undigested remains
• digest
• Absorb

• Length of the tube:

– short in carnivorous forms
– longer (often convoluted) in phytophagous forms
Digestion occurs by a series of progressive enzymatically
catalized steps
• polysaccharides are broken down into small chains,
disaccharides, and finally into simple absorbable
monosaccharides (e.g. glucose);
 • proteins are broken down into peptones,
small polypeptides, dipeptides, and finally
into amino acids which are absorbable.

Most absorption of organic molecules across the

midgut wall is passive, that is from a higher to a
lower concentration
Generalized insect alimentary system (adapted from Romoser and Stoffolano,1998)
MAJOR STRUCTURAL REGIONS OF GUT
FOREGUT
• During embryonic development, it develops from invaginating
ectodermal tissue at the anterior end of the body
• Epithelial Cells – usually flat squamous cells that do not secrete
enzymes into the lumen of the foregut; secrete a cuticular lining that is
attached to the surface of the cells on the lumen (apical) side
FOREGUT
• It can be divided into:
buccal cavity (mouth),
pharynx, esophagus,
crop, proventriculus, and
esophageal
invagination, and any
part may be highly
modified

At each molt, the old

cuticular lining from the
foregut is sloughed off into
the gut, and any
undigested residue is
eliminated with the feces.
EXTRA ORAL DIGESTION
• By injecting hydrolytic
enzymes into the food
source (animal or plant
material) and then
sucking back the
digested products,
insects utilize very high
percentages of the
nutrient value of the food
source
• Refluxing mixes
secretions and fluids and
extends the effective life
of the digestive enzymes
The left mandible (Md) of Dytiscus larva
(dorsal view), showing inner canal (c), a
groove on the inner face of the appendage
with confluent edges opening distally near
the tip (x) and proximally near the base (y)
of the jaw (after Snodgrass, 1993). Injection
of saliva with enzymes into the food source
and sucking back the digested product is
done through this kind of mandible.
THE PROVENTRICULUS (Gizzard)
• at the end of the foregut
• may be very muscular and
contains heavily
sclerotized teeth, ridges,
and spines for further
grinding and tearing of the
food, or it may be reduced
into a simple valve at the
entry of the midgut
• flap-like or valve-like
extensions of the
proventriculus sometimes
project into the midgut
• In worker honeybees: known as the
honey stopper
– consists of four converging fingers
projecting anteriorly into the crop
– the fingers, each bearing
intermeshing spines, open and
close rhythmically to capture pollen
grains and sweep them from the
crop into a bolus that later enters
the midgut for digestion
– nectar is strained through the
interlocking spines and retained in
the crop for later deposition in the
honeycomb
• In Orthopteroidea and usually in the Coleoptera: the
proventriculus functions as a grinding organ and as a valve, its
folds and teeth completely occluding the lumen when the circular
muscle sphincter is contained.
• In Carabidae: genera such as Calosoma and Carabus, which
feeds primarily on fluids following extra-intestinal digestion, have
long hairs lining the proventriculus.
• Hemiptera: are fluid feeders, have no proventriculus, but in the
carvivorous Hydrocorisae, the cibarial pump is modified for
grinding and filtering small fragments of prey which is ingested.
The cuticle becomes rigid, developed into a complex armature,
which grinds food as the two surfaces come together
MIDGUT
• the principal site for secretion of digestive enzymes and for
digestion and absorption in most insects.
• It has two main areas:
– tubular ventriculus
– gastric caeca (which arise at or near the origin of the
midgut, but they may be located at various points along the
midgut)
• to increase the surface area for either secretion of
digestive enzymes, or absorption of water, ions, glucose,
cholesterol, amino acids or probably other nutrients.
• digestion in certain insects also takes place in the gastric
caeca
• the evolutionary trend in the holometabolous insects is
toward the loss of the caeca
MIDGUT CELL TYPES

• Columnar cells
• Regenerative cells
• Goblet cells
COLUMNAR CELLS
• are the most numerous
cells in the midgut,
conduct most of
absorption of digested
products and secretion of
enzymes
• “Columnar” refers to the
tall shape of the cells, but
some insects have more
than one morphological
type
COLUMNAR CELLS
• Basic structure of columnar cell:
– Tall and columnar
– With regular microvilli, greatly increase the surface area
– The basal membranes of the cells, adjacent to the hemocoel,
are infolded with few openings
– Many mitochondria are enclosed by infoldings
– Endoplasmic reticulum with ribosomes assumed to be
concerned with the synthesis of digestive enzymes
– Laterally the cells are joined to each other by tight junctions and
septate desmosomes
REGENERATIVE CELLS
• replace midgut cells which wear out rapidly
• these cells grow into mature cells gradually and replace cells
lost through age, wear, and apocrine and holocrine secretion
GOBLET CELLS
• are, indeed goblet shaped, with a
large central cavity lined with
irregular microvilli and unlike the
microvilli of the columnar cells,
these often contain elongate
mitochondria.

• the sides of a goblet cell curve

around to enclose the cavity, and
at the apex the apical lips have
interdigiting microvilli that control
fluid exchange between cavity
and midgut lumen
GOBLET CELLS
• it has the apical border deeply
invaginated into the cell so that
a large cavity is formed
connecting with the gut lumen
by a narrow canal
• the cavity is bounded by
irregular microvilli

in caterpillars and in
Ephemeroptera and Plecoptera
goblet cells are distributed
between the columnar cells
PERITROPHIC MEMBRANE
• a thin sheath that separated the
midgut epithelium from the food
• consisting of a network of chitin
fibrils in a protein-glycoprotein
matrix (known as peritrophins)

• these proteins may have evolved

from gastrointestinal mucus proteins
by acquiring the ability to bind chitin
Suggested functions of peritrophic membrane include:
– Protection of the delicate microvilli on the surface of midgut
cells from contact with rough food particles.
– A barrier against entry of viruses, bacteria or other parasites
that would be too large to pass through the unbroken
peritrophic membrane
Suggested functions of peritrophic
membrane include:
– An aid in preventing the rapid
excretion of digestive
enzymes
– Compartmentalization of
digestion within the midgut
– Prevention of nonspecific
binding of undigested
materials or plant allelo-
chemicals to midgut
microvillar surfaces and/or
binding to transport proteins
at the midgut surface
HINDGUT
• develops in the embryo from ectodermal tissue, and,
consequently, hidgut cells have an attached cuticular lining on
their surface
• is most specialized in some insects that digest cellulose, such as
termites
• Malpighian tubules – usually mark the
beginning of the hindgut
• Ileum - The part of the hindgut
immediately past the Malpighian
tubules
• Sometimes the ileum simply
grades into the rectum, but in
some insects there is a distinct
middle region called the colon.
• Rectum - the terminal part of the
hindgut
• plays a critically important role in
the reabsorption of water, ions
and dissolved substances from
the primary urine flushed into the
hindgut by the Malpighian
tubules.
 DIGESTION

• Digestion in insects is both extra- and intra-cellular; the final

stages of chemical digestion of the food occur within the cells of
the midgut

• The initial stages, however occur in the gut lumen, predominantly

in the midgut, or in some species in the foregut

• Enzymes in the foregut may be products of the salivary glands or

be regurgitated from the midgut
SOURCES OF ENZYMES
• Salivary glands
– secrete digestive enzymes in most species
– salivary carbohydrases are common but proteases and
lipases also occur
– in some insects that practice external digestion by
salivating on the surface of food , or injecting saliva into a
victim and sucking up the soluble products, salivary glands
may be the sole source of enzymes
– in most insects, salivary secretion merely supplements the
activity of the midgut which is the main producer of
digestive enzymes
• Digestive enzymes of the midgut
• Microorganisms
Digestion of Carbohydrates
• Starch is broken down to maltose, and glycogen to glucose by
the action of amylase, which specifically catalyses the
hydrolysis of 1-4 alpha glycosidic linkages in polysaccharides,
but there are two types of amylase working in different ways
– exoamylase splits off maltose residues from the ends of
starch molecules leading to a rapid increase in the
concentration of maltose
– endoamylase attacks bonds within the starch molecule so
that there is only a slow build-up of maltose to start with
• The products are then further digested in the normal way by
alpha-glucosidases

• Most insects secrete enzymes capable of hydrolyzing simpler

carbohydrates like sucrose, maltose and amylase
Digestion of Proteins
• are broken down through peptones and peptides to amino acids.
• insects possess a series of proteases
• a trypsin-like proteinase which breaks down protein to peptones
and polypeptides is produced in the midgut
• are acted on by peptidases, some of which occur in the gut
lumen, but most of which are found in the epithelial cell indicating
that most of the polypeptides are absorbed before being digested
• There are different types of peptidase:
– Carboxypeptidase attacks the peptide chain from the –COOH
end, provided tyrosinase or other specific amino acids are
present in the chain

– Amino peptidase attacks the chain from the –NH2 end

– Dipeptidase hydrolyses all dipeptides
Digestion of Lipids

• Lipid is usually ingested mainly in the form of triglycerides

• Digestion involves the production of smaller molecules of
diglycerides, monoglycerides and free fatty acids
• Digestive lipases split fatty acids from triglycerides
 Digestion of Unusual Food:
CELLULOSE
• Cellulose, the structural component of plant cell walls, is an un-
branched glucose polymer
– Typically, it contain about 5000 glucose units
– Cotton: almost pure cellulose (95%); wood: about 50% cellulose
– The intestinal tracts of termites contain microorganisms that
produce cellulase
– Termites feed on and digest cellulose, as do some beetles, a few
cockroaches, and wood wasps in the family Siricidae
 Digestion of Unusual Food:
CHITIN
• Digestive chitinases have been identified in several predaceous
dragonflies, the carnivorous cockroach, and maybe of quite usual
occurrence in insectivorous species
Digestion of Unusual Food:
KERATIN
• Keratin: is the main protein in wool, hair and feathers
– It is made of polypeptide chains, including S-containing amino
acids, which are linked together by disulphide bonds (S-S)
rendering the whole protein stable
– Only some dermestid and tineid larvae and bird lice can digest
keratin
– Larvae of clothes moth (Tineola larva) have keratinase which
initiate the splitting of keratin.
– Digestion of wool by the clothes moth is aided by the
maintenance of extremely reducing conditions in the gut.
• Two enzymes involved are:
– cystine reductase which reduces cystine released from the
wool to cysteine

– sulphide desulfhydrase which splits hydrogen sulphide from

cysteine
• Cystine reductase & hydrogen sulphide: powerful reducing agents
capable of reducing the disulphide bridges of wool in the alkaline gut
contents, thereby rendering it more susceptible to attack by the gut
proteases

• Cysteine – the only standard amino acid that has a side that contains
a sulfhydryl group (-SH group)
 • Cysteine in the presence of mild oxidizing agents, readily dimerizes,
that is reacts with another cysteine molecule to form cystine
molecule. In cystine, the cysteine residues are linked via a covalent
disulfide bond

Cystine contains two cysteine residues linked by a disulfide bond. The covalent
disulfide bond of cystine is readily broken, using reducing agents, to regenerate
two cysteine molecules. This is the oxidation-reduction behavior involving
sulfhydryl groups and disulfide bonds (Wilbraham et al., 2000)
Digestion of Unusual Food:
COLLAGEN
• The larvae of Hypoderma
(Diptera) and some blowflies
are known to secrete a
collagenase which acts on the
collagen of animal tissues.
Hypoderma flies oviposit on
hairs of the host and the larvae
bore through the host’s skin
tissues.
Digestion of Unusual Food:
BEESWAX
– Beeswax which is not digested by
most animals, is the food of the wax
moth larva, Galleria melonella
– Paraffin constituents of the wax are
not digested by the moth’s own
enzymes, however, but by the
symbiotic bacteria that live in the gut.
• Microorganisms in other insects.
– Bacteria are the source of most carbohydrases found in the
digestive system of the cattle grub and in many calliphorids and
sarcophagids since these enzymes are absent in larvae reared
aseptically
Carbohydrate Absorption
– Glucose from digestion of carbohydrates is
rapidly absorbed passively by a process
known as facilitated diffusion
– Fat body cells, which can be found attached
in small groups on the hemolymph side of the
gut, rapidly synthesize absorbed glucose into
the disaccharide trehalose, keeping the
hemolymph concentration of glucose low in
most insects.
– Consequently, even with low concentration
of glucose in the gut, it can continue to be
absorbed passively

Trehalose – is the principal hemolymph sugar

in most insects
Amino Acid and Peptide Absorption
– Digestion of proteins in the intestinal lumen produces amino
acids and peptides of various sizes which are available for
absorption
– Peptide absorption occurs in insects suggesting that the final
stages of digestion are accomplished by membrane-bound
enzymes and amino acids are liberated intracellularly
Lipid Absorption
– Fatty acids undergo esterification as they traverse the gut cells
and are released into the hemolymph as diacylglycerols for
transport to fat body cells
– The diacylglycerols are picked up at the hemolymph side of the
gut by the lipoprotein complexes, lipophorins, that enable
transport of the absorbed lipids through the aqueous
hemolymph
– Lipids are mainly stored in fat body cells as triacylglycerols