Excretion in Insects
Excretion in Insects
- Excretion
CONTENTS
Introduction
The Malpighian tubules
Ultrastructure of Malpighian tubule cells
Formation of primary urine in Malpighian
tubules
A proton pump is the driving mechanism for
urine formation
Selective reabsorption in the hindgut
The role of the excretory system in
maintaining homeostasis
Cryptonephridial systems
Introduction
By analyzing the primary urine formed in the droplets, it was discovered that it
was isosmotic with the hemolymph, but with potassium concentrations up to 20
times higher.
Arrangement for Experimental
Perfusion of an Isolated Tubule
The Cumulative Formation of
Primary Urine by an Isolated Tubule
Anatomical Specialization of
Hindgut Epithelial Cells
The hindgut is the second system that completes
the excretion process by
– selectively reabsorbing some substances into the
hemolymph
– leaving others in the lumen
– actively secreting some substance into the hindgut lumen
The rectal cuticular lining has greater permeability
than the cuticular lining on foregut cells.
The epithelial cells of the hindgut are specialized for
both active secretion and active reabsorption.
Anatomical Specialization of
Hindgut Epithelial Cells
Rectal cells (or rectal pad cells, rectal papillae
cells) – a group cells in the rectum that have special
modifications for reabsorption. (next slide)
In Diptera, the cells of a rectal papilla are large,
usually cuboidal cells that surround a central
channel in the papilla that opens into the
hemolymph space through a valve. (next slide).
A Rectal Cell and Its Ion Transport
The rectum consists of the enlarged posterior-most
section of the hindgut, often containing specialized
structures called papillae or rectal pads that are enlarged
epithelial cells.
The rectum transports water and ions from the material
within the gut lumen into the hemolymph.
Secretion and Reabsorption in
the Ileum
The ileum is the most anterior part of the
hindgut, occurring just posterior to the origin of
the Malpighian tubules in most insects.
In locust S. gregaria, the ileum is a major site
for
– isosmotic fluid reabsorption
– active Na+ and Cl- reabsorption
– active secretion of proline as an energy source
Secretion and Reabsorption in
the Ileum
The driving mechanism for ion and water
reabsorption in the ileum is an electrogenic Cl-
pump.
A neuropeptide, the ion transport peptide (ITP)
stimulates Na+, Cl- and water reabsorption, and
promotes passive reabsorption of K+ by
electrical coupling.
The ileum plays a major role in acid-base
balance by secretion of H+ into the lumen,
formation of NH4+, and reabsorption of HCO3-.
Reabsorption in the Rectum
The rectum is the final and major site for
reabsorption of ions, water, and nutrients.
It is capable of reabsorbing fluid against strong
osmotic gradients, ultimately producing a very
concentrated hyperosmotic excreta in many
insects.
The driving mechanism for cation and water
reabsorption, as in the ileum, is an electrogenic
Cl- pump under the influence of a neuropeptide
hormone, chloride transport stimulating
hormone (CTSH), from the corpora cardiaca.
Fig. Ions are transported in and out of locust rectal cell by
numerous mechanisms.
Electrolyte Homeostasis
In mosquito A. aegypti, feeding on a blood meal
stimulates the release of mosquito natriuretic peptide
(MNP) from the CC, and cAMP is produced and acts
selectively to open Na+ channels in the basolateral
membrane of the Malpighian tubule cells.
Movement of water into tubule cells follows the
osmotic gradient.
The ion flex generated by MNP and cAMP is
specifically an increase in secretion of Na+. K+
movement is not influenced.
The Cl- load from the blood meal move from
hemolymph to tubule lumen in a passive transport
pathway between the cells (paracellular pathway).
Electrolyte Homeostasis
Larval A. aegypti live in fresh water, and in
response to an increase in salinity
– Secrete 5-hydroxytryptamine (serotonin) into the
hemolymph
– Increase cAMP in the Malpighian tubules
– Serotonin and cAMP stimulate fluid and ion (Na+
and K+) secretion rates in the tubules, but urine is
not concentrated with respect to the ions
Electrolyte Homeostasis
Beyenbach (1995) has reviewed three potential
physiological processes through which A. aegypti
may regulate rates of ion and fluid excretion
– The proton pump that supplies energy for Na+ and K+
secretion to the tubule lumen
– The resistance Rc across the tubule cells that control
ion channels in the basolateral membrane
– The resistance of the passive transport pathway for Cl-
movement
Water Homeostasis
Water excretion and retention are
regulated by hormones.
– Diuretic hormones promote fluid formation and
rapid excretion by the Malpighian tubules
Corticotropin-releasing factor (CRF)-related
peptides: range in size from 30-46 amino acids.
Insect kinins: small peptides of between 6 and 15
amino acids.
– Antidiuretic hormones act on the hindgut and
promote water reabsorption
Chloride transport-stimulating hormone (CTSH)
Ion transport peptide (ITP)
A Filter Chamber
In some Homopterans that feed exclusively on
plant juices containing low concentrations of
nutrients, the digestive tract forms an
arrangement known as a filter chamber.
Acid-Base Homeostasis
The excretory system is important in
maintaining the acid-base balance of
body fluids and tissues.
Acid-base regulation in S. gregaria
– Secretion of H+ and formation of NH4+ in the
ileum is a principal mechanism for excreting
excess acid equivalents.
– The ileum is a major site of ammoniagenesis
in locusts in which hindgut cells specifically
metabolize amino acids and glucose for
energy.
Acid-Base Homeostasis
Excretion of total ammonia nitrogen
serves several functions in locusts
– Ammonium urate (i.e. NH3 reacts with uric
acid) allows the insect to conserve Na+
– Conversion of NH3 to NH4+ in the ileal cells is
equivalent to removal of H+
– Excretion of NH3 by locusts conserves water
– Increases nitrogen excretion by 25% more
than excretion of only Na- or K-urate.
Nitrogen Homeostasis
The metabolism of proteins and nucleic acids
produce ammonia.
Some of this ammonia can be recycled into
amino acid synthesis by the formation of
glutamate from a-ketoglutarate and glutamine
from glutamate.
The excess ammonia that remains is highly
toxic unless it is diluted with water.
High levels of ammonia can interrupt nervous
transmission by substituting for necessary
potassium and can also alter carbohydrate and
lipid metabolism.
The Incorporation of Ammonia for
the Synthesis of Amino Acids
Nitrogen Homeostasis
Organisms must have excretory systems to
avoid the toxic accumulation of ammonia.
Because ammonia is very soluble in water, its
concentrations have to be maintained below
levels that are toxic.
Most terrestrial organisms have taken the
pathway of the incorporation of the nitrogen into
either urea or uric acid, which can be
concentrated in body fluid to a much greater
extent than can ammonia and require less
water for dilution.
Excretory Molecules that
Incorporate Nitrogen
Nitrogen Homeostasis
In insects, the need for water conservation may
have been the driving force for the incorporation
of their nitrogen wastes into uric acid.
The fat body is the primary site for uric acid
synthesis.
Uric acid does not dissolve well in water and
therefore fails to reach toxic levels in body fluids,
so it requires about 50 times less water to dilute
than does ammonia.
Insolubility of uric acid in water allows it to be
excreted in a dry form without having a significant
effect on water balance.
Nitrogen Homeostasis
Insects pay a high price for the benefits
they derive from employing uric acid as a
way to excrete nitrogen and still maintain
a positive water balance.
– The synthesis of uric acid results loss of
several carbon atoms.
– Eight ATP are required to first make the
intermediary metabolite, inosine
monophosphate (IMP)
Xanthine
dehygrogenase