The Malpighian tubule system is a type of

excretory Excretion is elimination of metabolic waste, which is an essential process in all organisms. In vertebrates, this is primarily carried out by the lungs, kidneys, and skin. This is in contrast with secretion, where the substance may have specific ...

and osmoregulatory system found in some

insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...

myriapods Myriapods () are the members of subphylum Myriapoda, containing arthropods such as millipedes and centipedes. The group contains about 13,000 species, all of them terrestrial. Although molecular evidence and similar fossils suggests a diversifi ...

arachnid Arachnids are arthropods in the Class (biology), class Arachnida () of the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, opiliones, harvestmen, Solifugae, camel spiders, Amblypygi, wh ...

s and

tardigrade Tardigrades (), known colloquially as water bears or moss piglets, are a phylum of eight-legged segmented micro-animals. They were first described by the German zoologist Johann August Ephraim Goeze in 1773, who called them . In 1776, th ...

s. It has also been described in some

crustacean Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquatic arthrop ...

species, and is likely the same organ as the posterior caeca which has been described in crustaceans. The system consists of branching tubules extending from the

alimentary canal The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The tract is the largest of the body's systems, after the cardiovascular system. ...

that absorbs

solutes In chemistry, a solution is defined by IUPAC as "A liquid or solid phase containing more than one substance, when for convenience one (or more) substance, which is called the solvent, is treated differently from the other substances, which are ...

, water, and wastes from the surrounding

hemolymph Hemolymph, or haemolymph, is a fluid, similar to the blood in invertebrates, that circulates in the inside of the arthropod's body, remaining in direct contact with the animal's tissues. It is composed of a fluid plasma in which hemolymph c ...

. The wastes then are released from the organism in the form of solid nitrogenous compounds and

calcium oxalate Calcium oxalate (in archaic terminology, oxalate of lime) is a calcium salt of oxalic acid with the chemical formula or . It forms hydrates , where ''n'' varies from 1 to 3. Anhydrous and all hydrated forms are colorless or white. The monohydr ...

. The system is named after

Marcello Malpighi Marcello Malpighi (10 March 1628 – 30 November 1694) was an Italians, Italian biologist and physician, who is referred to as the "founder of microscopical anatomy, histology and father of physiology and embryology". Malpighi's name is borne by ...

, a seventeenth-century anatomist.

Structure

Malpighian tubules are slender tubes normally found in the posterior regions of arthropod alimentary canals. Each tubule consists of a single layer of cells that is closed off at the distal end with the proximal end joining the alimentary canal at the junction between the midgut and hindgut. Most tubules are normally highly convoluted. The number of tubules varies between species although most occur in multiples of two. Tubules are usually bathed in

and are in proximity to fat body tissue. They contain

actin Actin is a family of globular multi-functional proteins that form microfilaments in the cytoskeleton, and the thin filaments in muscle fibrils. It is found in essentially all eukaryotic cells, where it may be present at a concentration of ...

for structural support and

microvilli Microvilli (: microvillus) are microscopic cellular membrane protrusions that increase the surface area for diffusion and minimize any increase in volume, and are involved in a wide variety of functions, including absorption, secretion, cellula ...

for propulsion of substances along the tubules. Malpighian tubules in most insects also contain accessory musculature associated with the tubules which may function to mix the contents of the tubules or expose the tubules to more hemolymph. The insect orders,

Dermaptera Earwigs make up the insect order Dermaptera. With about 2,000 species in 12 families, they are one of the smaller insect orders. Earwigs have characteristic cerci, a pair of forceps-like pincers on their abdomen, and membranous wings folde ...

and

Thysanoptera Thrips (order Thysanoptera) are minute (mostly long or less), slender insects with fringed wings and unique asymmetrical mouthparts. Entomologists have described approximately 7,700 species. They fly only weakly and their feathery wings are ...

do not possess these muscles and

Collembola Springtails (class Collembola) form the largest of the three lineages of modern hexapods that are no longer considered insects. Although the three lineages are sometimes grouped together in a class called Entognatha because they have internal m ...

and

Hemiptera Hemiptera (; ) is an order of insects, commonly called true bugs, comprising more than 80,000 species within groups such as the cicadas, aphids, planthoppers, leafhoppers, assassin bugs, bed bugs, and shield bugs. They range in size from ...

Aphididae The Aphididae are a very large insect family in the aphid superfamily ( Aphidoidea), of the order Hemiptera. These insects suck the sap from plant leaves. Several thousand species are placed in this family, many of which are considered plant/cr ...

completely lack a Malpighian tubule system.

General mode of action

Pre-urine is formed in the tubules, when

nitrogenous waste Metabolic wastes or excrements are substances left over from metabolic processes (such as cellular respiration) which cannot be used by the organism (they are surplus or toxic), and must therefore be excreted. This includes nitrogen compounds ...

and

electrolytes An electrolyte is a substance that conducts electricity through the movement of ions, but not through the movement of electrons. This includes most soluble salts, acids, and bases, dissolved in a polar solvent like water. Upon dissolving, t ...

are transported through the tubule walls. Wastes such as

urea Urea, also called carbamide (because it is a diamide of carbonic acid), is an organic compound with chemical formula . This amide has two Amine, amino groups (–) joined by a carbonyl functional group (–C(=O)–). It is thus the simplest am ...

and

amino acids Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although over 500 amino acids exist in nature, by far the most important are the Proteinogenic amino acid, 22 α-amino acids incorporated into p ...

are thought to diffuse through the walls, while ions such as

sodium Sodium is a chemical element; it has Symbol (chemistry), symbol Na (from Neo-Latin ) and atomic number 11. It is a soft, silvery-white, highly reactive metal. Sodium is an alkali metal, being in group 1 element, group 1 of the peri ...

and

potassium Potassium is a chemical element; it has Symbol (chemistry), symbol K (from Neo-Latin ) and atomic number19. It is a silvery white metal that is soft enough to easily cut with a knife. Potassium metal reacts rapidly with atmospheric oxygen to ...

are transported by active pump mechanisms. Water follows thereafter. The pre-urine, along with digested food, merge in the hindgut. At this time,

uric acid Uric acid is a heterocyclic compound of carbon, nitrogen, oxygen, and hydrogen with the Chemical formula, formula C5H4N4O3. It forms ions and salts known as urates and acid urates, such as ammonium acid urate. Uric acid is a product of the meta ...

precipitates out, and sodium and potassium ions are actively absorbed by the rectum, along with water via

osmosis Osmosis (, ) is the spontaneous net movement or diffusion of solvent molecules through a selectively permeable membrane, selectively-permeable membrane from a region of high water potential (region of lower solute concentration) to a region of ...

. Uric acid is left to mix with faeces, which are then excreted.

Alternative modes of action

Complex cycling systems of Malpighian tubules have been described in other insect orders.

n insects use tubules that permit movement of solutes into the distal portion of the tubules while reabsorption of water and essential ions directly to the hemolymph occurs in the proximal portion and the rectum. Both

Coleoptera Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...

and

Lepidoptera Lepidoptera ( ) or lepidopterans is an order (biology), order of winged insects which includes butterflies and moths. About 180,000 species of the Lepidoptera have been described, representing 10% of the total described species of living organ ...

use a cryptonephridial arrangement where the distal end of the tubules are embedded in fat tissue surrounding the rectum. Such an arrangement may serve to increase the efficiency of solute processing in the Malpighian tubules.

Other uses

Although primarily involved in excretion and osmoregulation, Malpighian tubules have been modified in some insects to serve accessory functions. Larvae of all species in genus ''

Arachnocampa ''Arachnocampa'' is a genus of nine fungus gnat species which have a bioluminescent larval stage, akin to the larval stage of glowworm beetles. The species of ''Arachnocampa'' are endemic to Australia and New Zealand, dwelling in caves and grotto ...

'' use modified and swollen Malpighian tubules to produce a blue-green lightGreen, L.B.S. (1979) The fine structure of the light organ of the New Zealand glow-worm ''Arachnocampa luminosa'' (Diptera: Mycetophilidae). ''Tissue and Cell'' 11: 457–465. attracting prey towards mucus-coated trap lines. In insects which feed on plant material containing noxious allelochemicals, Malpighian tubules also serve to rapidly excrete such compounds from the hemolymph.

References

*Gullan, P.J. and Cranston, P.S. (2000) ''The Insects: An Outline of Entomology.'' Blackwell Publishing UK *Romoser, W.S. and Stoffolano Jr., J.G. (1998) ''The Science of Entomology.'' McGraw-Hill Singapore *Bradley, T.J. The excretory system: structure and physiology. In: Kerkut, G.A. and Gilbert, L.I. eds. ''Comprehensive insect physiology, biochemistry and pharmacology.'' Vol.4 Pergamon Press New York {{ISBN, 0-08-030807-4 pp. 421–465 Arthropod anatomy Arachnid anatomy Insect anatomy

Structure

General mode of action

Alternative modes of action

Other uses

See also

References