The Malpighian tubule system is a type of

excretory Excretion is a process in which metabolic waste is eliminated from an organism. 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 tasks aft ...

and

osmoregulatory Osmoregulation is the active regulation of the osmotic pressure of an organism's body fluids, detected by osmoreceptors, to maintain the homeostasis of the organism's water content; that is, it maintains the fluid balance and the concentration o ...

system found in some

insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...

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. The fossil record of myriapods reaches back into the late Silurian, ...

arachnid Arachnida () is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegar ...

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 Kleiner Wasserb� ...

s. The system consists of branching

tubule In biology, a tubule is a general term referring to small tube or similar type of structure. Specifically, tubule can refer to: * a small tube or fistular structure * a minute tube lined with glandular epithelium * any hollow cylindrical body stru ...

s 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 GI tract contains all the major organs of the digestive system, in humans and ...

that absorbs

solutes In chemistry, a solution is a special type of homogeneous mixture composed of two or more substances. In such a mixture, a solute is a substance dissolved in another substance, known as a solvent. If the attractive forces between the solven ...

, water, and wastes from the surrounding

hemolymph Hemolymph, or haemolymph, is a fluid, analogous to the blood in vertebrates, that circulates in the interior of the arthropod (invertebrate) body, remaining in direct contact with the animal's tissues. It is composed of a fluid plasma in which ...

. 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 . It forms hydrates , where ''n'' varies from 1 to 3. Anhydrous and all hydrated forms are colorless or white. The monohydrate ...

. The system is named after

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

, 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 300px, Stained cells of an insect fat body Fat body is a highly dynamic insect tissue composed primarily of storage cells. It is distributed throughout the insect's internal body cavity; the haemocoel, in close proximity to the epidermis, dig ...

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 ov ...

for structural support and

microvilli Microvilli (singular: 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, ...

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 forcep-like pincers on their abdomen, and membranous wings folded ...

and

Thysanoptera Thrips (order Thysanoptera) are minute (mostly long or less), slender insects with fringed wings and unique asymmetrical mouthparts. Different thrips species feed mostly on plants by puncturing and sucking up the contents, although a few are ...

do not possess these muscles and

Collembola Springtails (Collembola) form the largest of the three lineages of modern hexapods that are no longer considered insects (the other two are the Protura and Diplura). Although the three orders are sometimes grouped together in a class called ...

and

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

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/crop ...

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 medium containing ions that is electrically conducting through the movement of those ions, but not conducting electrons. This includes most soluble salts, acids, and bases dissolved in a polar solvent, such as water. Upon di ...

are transported through the tubule walls. Wastes such as

urea Urea, also known as carbamide, is an organic compound with chemical formula . This amide has two amino groups (–) joined by a carbonyl functional group (–C(=O)–). It is thus the simplest amide of carbamic acid. Urea serves an important ...

and

amino acids Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha am ...

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

sodium Sodium is a chemical element with the symbol Na (from Latin ''natrium'') and atomic number 11. It is a soft, silvery-white, highly reactive metal. Sodium is an alkali metal, being in group 1 of the periodic table. Its only stable ...

and

potassium Potassium is the chemical element with the symbol K (from Neo-Latin '' kalium'') and atomic number19. Potassium is a silvery-white metal that is soft enough to be cut with a knife with little force. Potassium metal reacts rapidly with atmos ...

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 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 metabolic breakdown ...

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 from a region of high water potential (region of lower solute concentration) to a region of low water potential (region ...

. Uric acid is left to mix with feces, 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 order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describe ...

and

Lepidoptera Lepidoptera ( ) is an order of insects that includes butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 families and 46 superfamilies, 10 percent of the total described speci ...

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'' 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 Allelopathy is a biological phenomenon by which an organism produces one or more biochemicals that influence the germination, growth, survival, and reproduction of other organisms. These biochemicals are known as allelochemicals and can have ben ...

, 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