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Earthworms
""Earthworm.JPG
A species of earthworm with a well-developed clitellum
"Scientific classification
Kingdom: "Animalia
Phylum: "Annelida
Class: "Oligochaeta
Subclass: "Oligochaeta (but disputed, see "taxonomy) or "Haplotaxida
Order: Megadrilacea
Suborder: Lumbricina + Moniligastrida
"NODC v. 8.0, 1996[1]
"Families

An earthworm is a tube-shaped, segmented "worm found in the phylum "Annelida. Earthworms are commonly found living in soil, feeding on live and dead organic matter. An earthworm's digestive system runs through the length of its body. It conducts respiration through its skin. It has a double transport system composed of "coelomic fluid that moves within the fluid-filled "coelom and a simple, closed blood circulatory system. It has a central and a peripheral nervous system. The central nervous system consists of two ganglia above the mouth, one on either side, connected to a nerve cord running back along its length to motor neurons and sensory cells in each segment. Large numbers of chemoreceptors are concentrated near its mouth. Circumferential and longitudinal muscles on the periphery of each segment enable the worm to move. Similar sets of muscles line the gut, and their actions move the digesting food toward the worm's anus.[2]

Earthworms are "hermaphrodites–each individual carries both male and female sex organs. They lack either an internal "skeleton or "exoskeleton, but maintain their structure with fluid-filled coelom chambers that function as a "hydrostatic skeleton.

"Earthworm" is the common name for the largest members of "Oligochaeta (which is either a class or a subclass depending on the author). In classical systems, they were placed in the order Opisthopora, on the basis of the male pores opening posterior to the female pores, though the internal male segments are anterior to the female. Theoretical "cladistic studies have placed them, instead, in the suborder Lumbricina of the order Haplotaxida, but this may again soon change. Folk names for the earthworm include "dew-worm", "rainworm", "night crawler", and "angleworm" (due to its use as "fishing bait).

Larger terrestrial earthworms are also called megadriles (which translates to "big worms"), as opposed to the microdriles ("small worms") in the semiaquatic families "Tubificidae, "Lumbricidae, and "Enchytraeidae, among others. The megadriles are characterized by having a distinct "clitellum (which is more extensive than that of microdriles) and a vascular system with true capillaries.

Earthworms are far less abundant in disturbed environments and are typically active only if water is present.[3]

Contents

Anatomy[edit]

Form and function[edit]

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Earthworm head

Depending on the species, an adult earthworm can be from 10 mm (0.39 in) long and 1 mm (0.039 in) wide to 3 m (9.8 ft) long and over 25 mm (0.98 in) wide, but the typical "Lumbricus terrestris grows to about 360 mm (14 in) long.[4] Probably the longest worm on confirmed records is "Amynthas mekongianus that extends up to 3 m (10 ft) [5] in the mud along the banks of the 4,350 km (2,703 mi) Mekong River in Southeast Asia.

From front to back, the basic shape of the earthworm is a cylindrical tube, divided into a series of segments (called "metamerisms) that compartmentalize the body. Furrows are generally[6] externally visible on the body demarking the segments; dorsal pores and nephridiopores exude a fluid that moistens and protects the worm's surface, allowing it to breathe. Except for the mouth and anal segments, each segment carries bristle-like hairs called lateral "setae[7] used to anchor parts of the body during movement;[8] species may have four pairs of setae on each segment or more than eight sometimes forming a complete circle of setae per segment.[7] Special ventral setae are used to anchor mating earthworms by their penetration into the bodies of their mates.["citation needed]

Generally, within a species, the number of segments found is consistent across specimens, and individuals are born with the number of segments they will have throughout their lives. The first body segment (segment number 1) features both the earthworm's mouth and, overhanging the mouth, a fleshy lobe called the "prostomium, which seals the entrance when the worm is at rest, but is also used to feel and chemically sense the worm's surroundings. Some species of earthworm can even use the prehensile prostomium to grab and drag items such as grasses and leaves into their burrow.

An adult earthworm develops a belt-like glandular swelling, called the "clitellum, which covers several segments toward the front part of the animal. This is part of the reproductive system and produces egg capsules. The "posterior is most commonly cylindrical like the rest of the body, but depending on the species, may also be quadrangular, octagonal, trapezoidal, or flattened. The last segment is called the "periproct; the earthworm's anus, a short vertical slit, is found on this segment.[7]

""A segment of an earthworm posterior to the clitellum including all of the segmental structures

The exterior of an individual segment is a thin "cuticle over skin, commonly pigmented red to brown, which has specialized cells that secrete mucus over the cuticle to keep the body moist and ease movement through soil. Under the skin is a layer of nerve tissue, and two layers of muscles—a thin outer layer of circular muscle, and a much thicker inner layer of longitudinal muscle.[9] Interior to the muscle layer is a fluid-filled chamber called a "coelom[10] that by its pressurization provides structure to the worm's boneless body. The segments are separated from each other by septa (the plural of "septum")[11] which are perforated transverse walls, allowing the coelomic fluid to pass between segments.[12] A pair of structures called "nephrostomes are located at the back of each septum; a nephric tubule leads from each nephrostome through the septum and into the following segment. This tubule then leads to the main body fluid filtering organ, the "nephridium or metanephridium, which removes metabolic waste from the "coelomic fluid and expels it through pores called nephridiopores on the worm's sides; usually two nephridia (sometimes more) are found in most segments.[13] At the center of a worm is the "digestive tract, which runs straight through from mouth to anus without coiling, and is flanked above and below by blood vessels (the dorsal blood vessel and the ventral blood vessel as well as a subneural blood vessel) and the "ventral nerve cord, and is surrounded in each segment by a pair of pallial blood vessels that connect the dorsal to the subneural blood vessels.

Many earthworms can eject coelomic fluid through pores in the back in response to stress; Australian Didymogaster sylvaticus (known as the "blue squirter earthworm") can squirt fluid as high as 30 cm (12 in).[12]

Nervous system[edit]

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Nervous system of the anterior end of an earthworm

The earthworm's nervous system has three parts: the "central nervous system (CNS), "peripheral nervous system and the "sympathetic nervous system.[14][15]

Central nervous system[edit]

The CNS consists of a bilobed "brain (cerebral "ganglia, or supra-pharyngeal ganglion), sub-pharyngeal ganglia, circum-pharyngeal connectives and a "ventral nerve cord.

Earthworms' brains consist of a pair of pear-shaped cerebral ganglia. These are located in the dorsal side of the "alimentary canal in the third segment, in a groove between the "buccal cavity and "pharynx.

A pair of circum-pharyngeal connectives from the brain encircle the pharynx and then connect with a pair of sub-pharyngeal ganglia located below the pharynx in the fourth segment. This arrangement means the brain, sub-pharyngeal ganglia and the circum-pharyngeal connectives form a nerve ring around the pharynx.

The ventral nerve cord (formed by nerve cells and nerve fibres) begins at the sub-pharyngeal ganglia and extends below the alimentary canal to the most posterior body segment. The ventral nerve cord has a swelling, or ganglion, in each segment, i.e. a segmental ganglion, which occurs from the fifth to the last segment of the body. There are also three "giant axons, one medial giant axon (MGA) and two lateral giant axons (LGAs) on the mid-dorsal side of the ventral nerve cord. The MGA is 0.07 mm in diameter and transmits in an anterior-posterior direction at a rate of 32.2 m/s. The LGAs are slightly wider at 0.05 mm in diameter and transmit in a posterior-anterior direction at 12.6 m/s. The two LGAs are connected at regular intervals along the body and are therefore considered one giant axon.[16][17]

Peripheral nervous system[edit]

Sympathetic nervous system[edit]

The sympathetic nervous system consists of nerve plexuses in the epidermis and alimentary canal. (A plexus is a web of nerve cells connected together in a two dimensional grid.) The nerves that run along the body wall pass between the outer circular and inner longitudinal muscle layers of the wall. They give off branches that form the intermuscular plexus and the subepidermal plexus. These nerves connect with the circumpharyngeal connective.

Movement[edit]

On the surface, crawling speed varies both within and among individuals. Earthworms crawl faster primarily by taking longer "strides" and a greater frequency of strides. Larger Lumbricus terrestris worms crawl at a greater absolute speed than smaller worms. They achieve this by taking slightly longer strides but with slightly lower stride frequencies.[18]

Touching an earthworm, which causes a "pressure" response as well as (often) a response to the dehydrating quality of the salt on human skin (toxic to earthworms), stimulates the subepidermal nerve plexus which connects to the intermuscular plexus and causes the longitudinal muscles to contact, thereby the writhing movements when we pick up an earthworm. This behaviour is a "reflex and does not require the CNS; it occurs even if the nerve cord is removed. Each segment of the earthworm has its own nerve plexus. The plexus of one segment is not connected directly to that of adjacent segments. The nerve cord is required to connect the nervous systems of the segments.[19]

The giant axons carry the fastest signals along the nerve cord. These are emergency signals that initiate reflex escape behaviours. The larger dorsal giant axon conducts signals the fastest, from the rear to the front of the animal. If the rear of the worm is touched, a signal is rapidly sent forwards causing the longitudinal muscles in each segment to contract. This causes the worm shorten very quickly as an attempt to escape from a predator or other potential threat. The two medial giant axons connect with each other and send signals from the front to the rear. Stimulation of these causes the earthworm to very quickly retreat (perhaps contracting into its burrow to escape a bird).

The presence of a nervous system is essential for an animal to be able to experience "nociception or "pain. However, other physiological capacities are also required such as opioid sensitivity and central modulation of responses by analgesics.[20] "Enkephalin and "α-endorphin-like substances have been found in earthworms. Injections of "naloxone (an opioid antagonist) inhibit the escape responses of earthworms. This indicates that opioid substances play a role in sensory modulation, similar to that found in many vertebrates.[21]

Senses[edit]

Photosensitivity[edit]

Earthworms do not have eyes (although some worms do), however, they do have specialised photosensitive cells called "light cells of Hess". These photoreceptor cells have a central intracellular cavity (phaosome) filled with "microvilli. As well as the microvilli, there are several sensory "cilia in the phaosome which are structurally independent of the microvilli.[22] The photoreceptors are distributed in most parts of the epidermis but are more concentrated on the back and sides of the worm. A relatively small number occur on the ventral surface of the 1st segment. They are most numerous in the prostomium and reduce in density in the first three segments; they are very few in number past the third segment.[19]

Digestive system[edit]

The "gut of the earthworm is a straight tube which extends from the worm's mouth to its anus. It is differentiated into a buccal cavity (generally running through the first one or two segments of the earthworm), pharynx (running generally about four segments in length), esophagus, crop, gizzard (usually) and intestine.[23]

Food enters in the mouth. The pharynx acts as a suction pump; its muscular walls draw in food. In the pharynx, the pharyngeal glands secrete mucus. Food moves into the esophagus, where calcium (from the blood and ingested from previous meals) is pumped in to maintain proper blood calcium levels in the blood and food pH. From there the food passes into the crop and gizzard. In the gizzard, strong muscular contractions grind the food with the help of mineral particles ingested along with the food. Once through the gizzard, food continues through the intestine for digestion. The intestine secretes "Pepsin to digest proteins, "Amylase to digest polysaccharides, "Cellulase to digest cellulose, and lipase to digest fats.[2] Earthworms use, in addition to the digestive proteins, a class of surface active compounds called "drilodefensins, which help digest plant material.[24] Instead of being coiled like a mammalian intestine, an earthworm's intestine increases surface area to increase nutrient absorption by having many folds running along its length. The intestine has its own pair of muscle layers like the body, but in reverse order—an inner circular layer within an outer longitudinal layer.[25]

Circulatory system[edit]

The earthworm has a dual circulatory system in which both the coelomic fluid and a closed circulatory system carry the food, waste, and respiratory gases. The closed circulatory system has five main blood vessels: the dorsal (top) vessel, which runs above the digestive tract; the ventral (bottom) vessel, which runs below the digestive tract; the subneural vessel, which runs below the ventral nerve cord; and two lateroneural vessels on either side of the nerve cord.[26] The dorsal vessel moves the blood forward, while the other four longitudinal vessels carry the blood rearward. In segments seven through eleven, a pair of aortic arches rings the coelom and acts as hearts, pumping the blood to the ventral vessel that acts as the aorta. The blood consists of ameboid cells and hemoglobin dissolved in the plasma. The second circulatory system derives from the cells of the digestive system that line the coelom. As the digestive cells become full, they release non-living cells of fat into the fluid-filled coelom, where they float freely but can pass through the walls separating each segment, moving food to other parts and assist in wound healing.[27]

Excretory system[edit]

The excretory system contains a pair of "nephridia in every segment, except for the first three and the last ones.[28] The three types of nephridia are: integumentary, septal, and pharyngeal. The integumentary nephridia lie attached to the inner side of the body wall in all segments except the first two. The septal nephridia are attached to both sides of the septa behind the 15th segment. The pharyngeal nephridia are attached to fourth, fifth and sixth segments.[28] The waste in the coelom fluid from a forward segment is drawn in by the beating of "cilia of the "nephrostome. From there it is carried through the septum (wall) via a tube which forms a series of loops entwined by blood capillaries that also transfer waste into the tubule of the nephrostome. The excretory wastes are then finally discharged through a pore on the worm's side.[29]

Respiration[edit]

Earthworms have no special respiratory organs. Gases are exchanged through the moist skin and capillaries, where the oxygen is picked up by the hemoglobin dissolved in the blood plasma and carbon dioxide is released. Water, as well as salts, can also be moved through the skin by active transport.

Reproduction[edit]

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Earthworm copulation
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Earthworm cocoons from L. terrestris
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An earthworm cocoon from L. rubellus

Mating occurs on the surface, most often at night. Earthworms are "hermaphrodites; that is, they have both male and female sexual organs. The sexual organs are located in segments 9 to 15. Earthworms have one or two pairs of "testes contained within sacs. The two or four pairs of "seminal vesicles produce, store and release the sperm via the male pores. Ovaries and oviducts in segment 13 release eggs via female pores on segment 14, while sperm is expelled from segment 15. One or more pairs of "spermathecae are present in segments 9 and 10 (depending on the species) which are internal sacs that receive and store sperm from the other worm during copulation. As a result, segment 15 of one worm exudes sperm into segments 9 and 10 with its storage vesicles of its mate. Some species use external "spermatophores for sperm transfer.

In Hormogaster samnitica and Hormogaster elisae "transcriptome DNA libraries were sequenced and two sex "pheromones, Attractin and Temptin, were detected in all tissue samples of both "species.[30] Sex pheromones are probably important in earthworms because they live in an environment where chemical signaling may play a crucial role in attracting a partner and in facilitating outcrossing. Outcrossing would provide the benefit of masking the expression of deleterious recessive mutations in progeny.[31] (Also see "complemenation.)

"Copulation and "reproduction are separate processes in earthworms. The mating pair overlap front ends "ventrally and each exchanges sperm with the other. The "clitellum becomes very reddish to pinkish in color. Some time after copulation, long after the worms have separated, the clitellum (behind the spermathecae) secretes material which forms a ring around the worm. The worm then backs out of the ring, and as it does so, it injects its own eggs and the other worm's sperm into it. As the worm slips out of the ring, the ends of the cocoon seal to form a vaguely lemon-shaped incubator ("cocoon) in which the embryonic worms develop. They emerge as small, but fully formed earthworms, but lack their sex structures, which develop in about 60 to 90 days. They attain full size in about one year. Scientists predict that the average lifespan under field conditions is four to eight years, while most garden varieties live only one to two years. Several common earthworm species are mostly "parthenogenetic.

Among lumbricid earthworms, parthenogenesis arose from sexual relatives many times.[32] Parthenogenesis in some Aporrectodea trapezoides lineages arose 6.4 to 1.1 million years ago from sexual ancestors.[33]

Regeneration[edit]

Earthworms have the ability to regenerate lost segments, but this ability varies between species and depends on the extent of the damage. Stephenson (1930) devoted a chapter of his monograph to this topic, while G.E. Gates spent 20 years studying regeneration in a variety of species, but “because little interest was shown”, Gates (1972) only published a few of his findings that, nevertheless, show it is theoretically possible to grow two whole worms from a bisected specimen in certain species. Gates’s reports included:

An unidentified Tasmanian earthworm shown growing a replacement head has been reported.[36]

Locomotion and importance to soil[edit]

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Close up of an earthworm in garden soil

Earthworms travel underground by the means of waves of muscular contractions which alternately shorten and lengthen the body ("peristalsis). The shortened part is anchored to the surrounding soil by tiny claw-like bristles ("setae) set along its segmented length. In all the body segments except the first, last and clitellum, there is a ring of S-shaped setae embedded in the epidermal pit of each segment (perichaetine). The whole burrowing process is aided by the secretion of lubricating mucus. Worms can make gurgling noises underground when disturbed as a result of their movement through their lubricated tunnels. Earthworms move through soil by expanding crevices with force; when forces are measured according to body weight, hatchlings can push 500 times their own body weight whereas large adults can push only 10 times their own body weight.[37]

Earthworms work as biological ""pistons" forcing air through the tunnels as they move. Thus earthworm activity aerates and mixes the soil, and is conducive to mineralization of nutrients and their uptake by vegetation. Certain species of earthworm come to the surface and graze on the higher concentrations of organic matter present there, mixing it with the mineral soil. Because a high level of organic matter mixing is associated with soil fertility, an abundance of earthworms is generally considered beneficial by farmers and gardeners.[38][39] In fact, as long ago as 1881 "Charles Darwin wrote: "It may be doubted whether there are many other animals which have played so important a part in the history of the world, as have these lowly organized creatures."[40]

Benefits[edit]

The major benefits of earthworm activities to soil fertility can be summarized as:

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"Faeces in form of casts

Earthworms accelerate nutrient cycling in the soil-plant system through fragmentation & mixing of plant debris – physical grinding & chemical digestion.[3] The earthworm's existence cannot be taken for granted. Dr. "W. E. Shewell-Cooper observed "tremendous numerical differences between adjacent gardens", and worm populations are affected by a host of environmental factors, many of which can be influenced by good management practices on the part of the gardener or farmer.[42]

Darwin estimated that "arable land contains up to 53,000 worms per acre (13/m2), but more recent research from "Rothamsted Experimental Station has produced figures suggesting that even poor soil may support 250,000/acre (62/m2), whilst rich fertile farmland may have up to 1,750,000/acre (432/m2), meaning that the weight of earthworms beneath a farmer's soil could be greater than that of the livestock upon its surface.

The ability to break down organic materials and excrete concentrated nutrients makes the earthworm a functional contributor in restoration projects. In response to ecosystem disturbances, some opencast mining sites have utilized earthworms to prepare soil for the return of native flora. Sites using this method have observed advances in the return of ecosystem services that previously took much longer to re-establish.[43] Research from the Station d'écologie Tropicale de Lamto asserts that the earthworms positively influence the rate of macroaggregate formation, an important feature for soil structure.[44] The stability of aggregates in response to water was also found to be improved when constructed by earthworms.[44]

As an invasive species[edit]

From a total of around 7,000 species, only about 150 species are widely distributed around the world. These are the peregrine or cosmopolitan earthworms.[45]

Special habitats[edit]

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Permanent vertical burrow
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"Devil's coach horse beetle preying on Lumbricus sp.

While, as the name earthworm suggests, the main habitat of earthworms is in soil, the situation is more complicated than that. The brandling worm "Eisenia fetida lives in decaying plant matter and manure. Arctiostrotus vancouverensis from "Vancouver Island and the "Olympic Peninsula is generally found in decaying conifer logs. Aporrectodea limicola, "Sparganophilus spp., and several others are found in mud in streams. Some species are arboreal, some aquatic and some euryhaline (salt-water tolerant) and littoral (living on the sea-shore, e.g. Pontodrilus litoralis). Even in the soil species, special habitats, such as soils derived from "serpentine, have an earthworm fauna of their own.

Ecology[edit]

Earthworms are classified into three main ecophysiological categories: (1) leaf litter- or compost-dwelling worms that are nonburrowing, live at soil-litter interface and eat decomposing OM (called Epigeic) e.g. Eisenia fetida; (2) topsoil- or subsoil-dwelling worms that feed (on soil), burrow and cast within soil, creating horizontal burrows in upper 10–30 cm of soil (called Endogeics); and (3) worms that construct permanent deep vertical burrows which they use to visit the surface to obtain plant material for food, such as leaves (called Anecic (meaning "reaching up")), e.g. Lumbricus terrestris.[46]

Earthworm populations depend on both physical and chemical properties of the soil, such as temperature, moisture, pH, salts, aeration, and texture, as well as available food, and the ability of the species to reproduce and disperse. One of the most important environmental factors is "pH, but earthworms vary in their preferences. Most favor neutral to slightly acidic soils. Lumbricus terrestris is still present in a pH of 5.4 and Dendrobaena octaedra at a pH of 4.3 and some "Megascolecidae are present in extremely acidic humic soils. Soil pH may also influence the numbers of worms that go into "diapause. The more acidic the soil, the sooner worms go into diapause and the longer they remain in diapause at a pH of 6.4.

Earthworms form the base of many "food chains. They are preyed upon by many species of "birds (e.g. "starlings, "thrushes, "gulls, "crows, "European robins and "American robins), snakes, mammals (e.g. "bears, "foxes, "hedgehogs, "pigs, "moles) and invertebrates (e.g. "ground beetles and other "beetles, "snails, "slugs). Earthworms have many internal "parasites, including "protozoa, "platyhelminthes, and "nematodes; they can be found in the worms' "blood, "seminal vesicles, "coelom, or "intestine, or in their "cocoons.

Nitrogenous fertilizers tend to create "acidic conditions, which are fatal to the worms, and dead specimens are often found on the surface following the application of substances such as "DDT, "lime sulphur, and "lead arsenate. In "Australia, changes in farming practices such as the application of "superphosphates on "pastures and a switch from "pastoral farming to "arable farming had a devastating effect on populations of the giant "Gippsland earthworm, leading to their classification as a "protected species.

The addition of organic matter, preferably as a surface mulch, on a regular basis will provide earthworms with their food and nutrient requirements, and will create the optimum conditions of temperature and moisture that will stimulate their activity.

Economic impact[edit]

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Earthworms being raised at "La Chonita Hacienda in Mexico

Various species of worms are used in "vermiculture, the practice of feeding organic waste to earthworms to decompose food waste. These are usually "Eisenia fetida (or its close relative "Eisenia andrei) or the Brandling worm, commonly known as the tiger worm or red wiggler. They are distinct from soil-dwelling earthworms. In the tropics, the African nightcrawler "Eudrilus eugeniae and the Indian blue "Perionyx excavatus are used.

Earthworms are sold all over the world; the market is sizable. According to Doug Collicut, "In 1980, 370 million worms were exported from Canada, with a Canadian export value of $13 million and an American retail value of $54 million."

Earthworms are also sold as food for human consumption. "Noke is a culinary term used by the "Māori of "New Zealand, and refers to earthworms which are considered delicacies for their chiefs.

Taxonomy and distribution[edit]

Within the world of taxonomy, the stable 'Classical System' of Michaelsen (1900) and Stephenson (1930) was gradually eroded by the controversy over how to classify earthworms, such that Fender and McKey-Fender (1990) went so far as to say, "The family-level classification of the megascolecid earthworms is in chaos."[47] Over the years, many scientists developed their own classification systems for earthworms, which led to confusion, and these systems have been and still continue to be revised and updated. The classification system used here, developed by Blakemore (2000), is a modern reversion to the Classical System that is historically proven and widely accepted.[48]

Categorization of a megadrile earthworm into one of its taxonomic families under suborders Lumbricina and Moniligastrida is based on such features as the makeup of the clitellum, the location and disposition of the sex features (pores, prostatic glands, etc.), number of gizzards, and body shape.[48] Currently, over 6,000 species of terrestrial earthworms are named, as provided in a species name database,[49] but the number of synonyms is unknown.

The families, with their known distributions or origins:[48]

See also[edit]

References[edit]

  1. ^ "ITIS Report for Lumbricina, Taxonomic Serial No.: 69069". ITIS. Retrieved May 14, 2012. 
  2. ^ a b Cleveland P. Hickman Jr., Larry S. Roberts, Frances M Hickman (1984). Integrated Principles of Zoology (7th ed.). Times Mirror/Mosby College Publishing. p. 344. "ISBN "0-8016-2173-9. 
  3. ^ a b c d e Nyle C. Brady; Ray R. Weil (2009). Elements of the Nature and Properties of Soils (3rd Edition). Prentice Hall. "ISBN "978-0-13-501433-2. 
  4. ^ Blakemore 2012, p. xl.
  5. ^ R.J. Blakemore et al. (2007). "Megascolex (Promegascolex) mekongianus Cognetti, 1922 - its extent, ecology and allocation to Amynthas (Clitellata/Oligochaeta: Megascolecidae)" ("PDF). Opuscula Zoologica. 
  6. ^ Edwards & Bohlen 1996, p. 11.
  7. ^ a b c Sims & Gerard 1985, pp. 3–6.
  8. ^ Edwards & Bholen 1996, p. 3.
  9. ^ Edwards & Bohlen 1996, p. 8-9.
  10. ^ Edwards & Bohlen 1996, p. 1.
  11. ^ Sims & Gerard 1985, p. 8.
  12. ^ a b Edwards & Bohlen 1996, p. 12.
  13. ^ Edwards & Bohlen 1996, p. 6.
  14. ^ "Brain – Invertebrate Brain – Ganglia, Nervous, System, and Head". HSBE. Retrieved April 3, 2015. 
  15. ^ Priyadarshini, S. "Different parts of nervous system of earthworm (with diagram)". Biology Discussion. Retrieved April 3, 2015. 
  16. ^ "Experiment: Comparing speeds of two nerve fiber sizes". BackyardBrains. Retrieved April 4, 2015. 
  17. ^ Drewes, C.D., Landa, K.B. and McFall, J.L. (1978). "Giant nerve fibre activity in intact, freely moving earthworms". The Journal of Experimental Biology. 72: 217–227. "PMID 624897. 
  18. ^ Quillin, K.J. (1999). "Kinematic scaling of locomotion by hydrostatic animals: ontogeny of peristaltic crawling by the earthworm lumbricus terrestris". Journal of Experimental Biology. 202: 661–674. "PMID 10021320. 
  19. ^ a b "Earthworm-nervous system". Cronodon. Retrieved April 3, 2015. 
  20. ^ Elwood, R.W. (2011). "Pain and suffering in invertebrates?". ILAR Journal. 52 (2): 175–84. "PMID 21709310. 
  21. ^ Smith, J.A. "A question of pain in invertebrates". ILAR Journal. 33 (1–2): 25–31. Retrieved April 3, 2015. 
  22. ^ Röhlich, P., Aros, B. and Virágh, Sz. (1970). "Fine structure of photoreceptor cells in the earthworm, Lumbricus terrestris". Zeitschrift für Zellforschung und Mikroskopische Anatomie. 104 (3): 345–357. "doi:10.1007/BF00335687. 
  23. ^ Edwards & Bohlen 1996, p. 13.
  24. ^ Liebeke, Manuel; Strittmatter, Nicole; Fearn, Sarah; Morgan, A. John; Kille, Peter; Fuchser, Jens; Wallis, David; Palchykov, Vitalii; Robertson, Jeremy (2015-08-04). "Unique metabolites protect earthworms against plant polyphenols". Nature Communications. 6: 7869. "doi:10.1038/ncomms8869. "PMC 4532835Freely accessible. "PMID 26241769. 
  25. ^ Edwards & Bohlen 1996, pp. 13–15.
  26. ^ Sims & Gerard 1985, p. 10.
  27. ^ Cleveland P. Hickman Jr., Larry S. Roberts, Frances M Hickman (1984). Integrated Principles of Zoology (7th ed.). Times Mirror/Mosby College Publishing. pp. 344–345. "ISBN "0-8016-2173-9. 
  28. ^ a b Farabee, H.J. "Excretory System". Archived from the original on 30 July 2012. Retrieved 29 July 2012. 
  29. ^ Cleveland P. Hickman Jr., Larry S. Roberts, Frances M Hickman (1984). Integrated Principles of Zoology (7th ed.). Times Mirror/Mosby College Publishing. pp. 345–346. "ISBN "0-8016-2173-9. 
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Further reading[edit]

External links[edit]

General[edit]

Academic[edit]

Agriculture and ecology[edit]

Worm farming[edit]

For children[edit]

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