Soil , also commonly referred to as earth , is a mixture of organic matter , minerals , gases , liquids , and organisms that together support the life of plants and soil organisms . Some scientific definitions distinguish dirt from soil by restricting the former term specifically to displaced soil.
127-403: An earthworm is a soil -dwelling terrestrial invertebrate that belongs to the phylum Annelida . The term is the common name for the largest members of the class (or subclass , depending on the author) Oligochaeta . In classical systems, they were in the order of Opisthopora since the male pores opened posterior to the female pores, although the internal male segments are anterior to
254-418: 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. Currently, over 6,000 species of terrestrial earthworms are named, as provided in a species name database, but the number of synonyms
381-438: 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
508-466: 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 contract. This causes the writhing movements observed when a human picks up an earthworm. This behaviour is a reflex and does not require the CNS; it occurs even if
635-435: A bisected specimen in certain species. Gates's reports included: An unidentified Tasmanian earthworm shown growing a replacement head has been reported. 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
762-461: 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 (see Complementation ). 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 colour. Sometime after copulation, long after
889-437: A double transport system made of coelomic fluid that moves within the fluid-filled coelom and a simple, closed circulatory system , and respire (breathe) via cutaneous respiration . As soft-bodied invertebrates, they lack a true skeleton , but their structure is maintained by fluid-filled coelom chambers that function as a hydrostatic skeleton . Earthworms have a central nervous system consisting of two ganglia above
1016-643: A dry bulk density (density of soil taking into account voids when dry) between 1.1 and 1.6 g/cm , though the soil particle density is much higher, in the range of 2.6 to 2.7 g/cm . Little of the soil of planet Earth is older than the Pleistocene and none is older than the Cenozoic , although fossilized soils are preserved from as far back as the Archean . Collectively the Earth's body of soil
1143-451: A medium for plant growth , making it a critically important provider of ecosystem services . Since soil has a tremendous range of available niches and habitats , it contains a prominent part of the Earth's genetic diversity . A gram of soil can contain billions of organisms, belonging to thousands of species, mostly microbial and largely still unexplored. Soil has a mean prokaryotic density of roughly 10 organisms per gram, whereas
1270-418: A mouth, 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. Food enters at 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
1397-419: A net absorption of oxygen and methane and undergo a net release of carbon dioxide and nitrous oxide . Soils offer plants physical support, air, water, temperature moderation, nutrients, and protection from toxins. Soils provide readily available nutrients to plants and animals by converting dead organic matter into various nutrient forms. Components of a silt loam soil by percent volume A typical soil
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#17327800128821524-549: A pH of 9, plant growth is reduced. High pH results in low micro-nutrient mobility, but water-soluble chelates of those nutrients can correct the deficit. Sodium can be reduced by the addition of gypsum (calcium sulphate) as calcium adheres to clay more tightly than does sodium causing sodium to be pushed into the soil water solution where it can be washed out by an abundance of water. There are acid-forming cations (e.g. hydronium, aluminium, iron) and there are base-forming cations (e.g. calcium, magnesium, sodium). The fraction of
1651-431: A pair of commissurals running along the posterior surface of the septum. The pumping action on 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 ring 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 haemoglobin dissolved in
1778-540: 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; the Australian Didymogaster sylvaticus (known as the "blue squirter earthworm") can squirt fluid as high as 30 cm (12 in). The CNS consists of a bilobed brain (cerebral ganglia , or supra-pharyngeal ganglion), sub-pharyngeal ganglia, circum-pharyngeal connectives and
1905-731: A result of a trade-off between toxicity and requirement most nutrients are better available to plants at moderate pH, although most minerals are more soluble in acid soils. Soil organisms are hindered by high acidity, and most agricultural crops do best with mineral soils of pH 6.5 and organic soils of pH 5.5. Given that at low pH toxic metals (e.g. cadmium, zinc, lead) are positively charged as cations and organic pollutants are in non-ionic form, thus both made more available to organisms, it has been suggested that plants, animals and microbes commonly living in acid soils are pre-adapted to every kind of pollution, whether of natural or human origin. In high rainfall areas, soils tend to acidify as
2032-401: A result, layers (horizons) form in the soil profile. The alteration and movement of materials within a soil causes the formation of distinctive soil horizons . However, more recent definitions of soil embrace soils without any organic matter, such as those regoliths that formed on Mars and analogous conditions in planet Earth deserts. An example of the development of a soil would begin with
2159-412: A soil can be said to be developed, and can be described further in terms of color, porosity, consistency, reaction ( acidity ), etc. Water is a critical agent in soil development due to its involvement in the dissolution, precipitation, erosion, transport, and deposition of the materials of which a soil is composed. The mixture of water and dissolved or suspended materials that occupy the soil pore space
2286-399: A soil determines its ability to supply available plant nutrients and affects its physical properties and the health of its living population. In addition, a soil's chemistry also determines its corrosivity , stability, and ability to absorb pollutants and to filter water. It is the surface chemistry of mineral and organic colloids that determines soil's chemical properties. A colloid
2413-472: A valence of two, converts to (40 ÷ 2) × 1 milliequivalent = 20 milliequivalents of hydrogen ion per 100 grams of dry soil or 20 meq/100 g. The modern measure of CEC is expressed as centimoles of positive charge per kilogram (cmol/kg) of oven-dry soil. Most of the soil's CEC occurs on clay and humus colloids, and the lack of those in hot, humid, wet climates (such as tropical rainforests ), due to leaching and decomposition, respectively, explains
2540-553: Is 0.04%, but in the soil pore space it may range from 10 to 100 times that level, thus potentially contributing to the inhibition of root respiration. Calcareous soils regulate CO 2 concentration by carbonate buffering , contrary to acid soils in which all CO 2 respired accumulates in the soil pore system. At extreme levels, CO 2 is toxic. This suggests a possible negative feedback control of soil CO 2 concentration through its inhibitory effects on root and microbial respiration (also called soil respiration ). In addition,
2667-603: Is a measure of the acidity or alkalinity of the soil. More precisely, it is a measure of hydronium concentration in an aqueous solution and ranges in values from 0 to 14 (acidic to basic) but practically speaking for soils, pH ranges from 3.5 to 9.5, as pH values beyond those extremes are toxic to life forms. At 25 °C an aqueous solution that has a pH of 3.5 has 10 moles H 3 O (hydronium ions) per litre of solution (and also 10 moles per litre OH ). A pH of 7, defined as neutral, has 10 moles of hydronium ions per litre of solution and also 10 moles of OH per litre; since
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#17327800128822794-488: Is a product of several factors: the influence of climate , relief (elevation, orientation, and slope of terrain), organisms, and the soil's parent materials (original minerals) interacting over time. It continually undergoes development by way of numerous physical, chemical and biological processes, which include weathering with associated erosion . Given its complexity and strong internal connectedness , soil ecologists regard soil as an ecosystem . Most soils have
2921-509: Is a small, insoluble particle ranging in size from 1 nanometer to 1 micrometer , thus small enough to remain suspended by Brownian motion in a fluid medium without settling. Most soils contain organic colloidal particles called humus as well as the inorganic colloidal particles of clays . The very high specific surface area of colloids and their net electrical charges give soil its ability to hold and release ions . Negatively charged sites on colloids attract and release cations in what
3048-411: Is about 50% solids (45% mineral and 5% organic matter), and 50% voids (or pores) of which half is occupied by water and half by gas. The percent soil mineral and organic content can be treated as a constant (in the short term), while the percent soil water and gas content is considered highly variable whereby a rise in one is simultaneously balanced by a reduction in the other. The pore space allows for
3175-424: Is also a net sink of methane (CH 4 ) but a net producer of methane (a strong heat-absorbing greenhouse gas ) when soils are depleted of oxygen and subject to elevated temperatures. Soil atmosphere is also the seat of emissions of volatiles other than carbon and nitrogen oxides from various soil organisms, e.g. roots, bacteria, fungi, animals. These volatiles are used as chemical cues, making soil atmosphere
3302-437: 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-shaped 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
3429-556: Is an important characteristic of soil. This ventilation can be accomplished via networks of interconnected soil pores , which also absorb and hold rainwater making it readily available for uptake by plants. Since plants require a nearly continuous supply of water, but most regions receive sporadic rainfall, the water-holding capacity of soils is vital for plant survival. Soils can effectively remove impurities, kill disease agents, and degrade contaminants , this latter property being called natural attenuation . Typically, soils maintain
3556-464: Is assumed acid-forming cations). Base saturation is almost in direct proportion to pH (it increases with increasing pH). It is of use in calculating the amount of lime needed to neutralise an acid soil (lime requirement). The amount of lime needed to neutralize a soil must take account of the amount of acid forming ions on the colloids (exchangeable acidity), not just those in the soil water solution (free acidity). The addition of enough lime to neutralize
3683-414: Is called the pedosphere . The pedosphere interfaces with the lithosphere , the hydrosphere , the atmosphere , and the biosphere . Soil has four important functions : All of these functions, in their turn, modify the soil and its properties. Soil science has two basic branches of study: edaphology and pedology . Edaphology studies the influence of soils on living things. Pedology focuses on
3810-401: Is called the soil solution. Since soil water is never pure water, but contains hundreds of dissolved organic and mineral substances, it may be more accurately called the soil solution. Water is central to the dissolution , precipitation and leaching of minerals from the soil profile . Finally, water affects the type of vegetation that grows in a soil, which in turn affects the development of
3937-421: Is dependent on the type of parent material , the processes that modify those parent materials, and the soil-forming factors that influence those processes. The biological influences on soil properties are strongest near the surface, though the geochemical influences on soil properties increase with depth. Mature soil profiles typically include three basic master horizons: A, B, and C. The solum normally includes
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4064-401: Is induced. Capillary action is responsible for moving groundwater from wet regions of the soil to dry areas. Subirrigation designs (e.g., wicking beds , sub-irrigated planters ) rely on capillarity to supply water to plant roots. Capillary action can result in an evaporative concentration of salts, causing land degradation through salination . Soil moisture measurement —measuring
4191-531: Is likely home to 59 ± 15% of the species on Earth. Enchytraeidae (worms) have the greatest percentage of species in soil (98.6%), followed by fungi (90%), plants (85.5%), and termites ( Isoptera ) (84.2%). Many other groups of animals have substantial fractions of species living in soil, e.g. about 30% of insects , and close to 50% of arachnids . While most vertebrates live above ground (ignoring aquatic species), many species are fossorial , that is, they live in soil, such as most blind snakes . The chemistry of
4318-477: Is more advanced. Most plant nutrients, with the exception of nitrogen , originate from the minerals that make up the soil parent material. Some nitrogen originates from rain as dilute nitric acid and ammonia , but most of the nitrogen is available in soils as a result of nitrogen fixation by bacteria . Once in the soil-plant system, most nutrients are recycled through living organisms, plant and microbial residues (soil organic matter), mineral-bound forms, and
4445-427: Is most commonly cylindrical like the rest of the body, but depending on the species, it 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. The exterior of an individual segment is a thin cuticle over the skin, commonly pigmented red to brown, which has specialized cells that secrete mucus over
4572-440: Is necessary to stimulate reproduction, even though no male genetic material passes to the offspring. Earthworm 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
4699-409: Is referred to as cation exchange . Cation-exchange capacity is the amount of exchangeable cations per unit weight of dry soil and is expressed in terms of milliequivalents of positively charged ions per 100 grams of soil (or centimoles of positive charge per kilogram of soil; cmol c /kg ). Similarly, positively charged sites on colloids can attract and release anions in the soil, giving
4826-540: Is unknown. The families, with their known distributions or origins: 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. Of the 182 taxa of earthworms found in the United States and Canada, 60 (33%) are introduced species. Earthworms are classified into three main ecophysiological categories: (1) leaf litter - or compost-dwelling worms that are nonburrowing, live at
4953-545: Is very little organic material. Basaltic minerals commonly weather relatively quickly, according to the Goldich dissolution series . The plants are supported by the porous rock as it is filled with nutrient-bearing water that carries minerals dissolved from the rocks. Crevasses and pockets, local topography of the rocks, would hold fine materials and harbour plant roots. The developing plant roots are associated with mineral-weathering mycorrhizal fungi that assist in breaking up
5080-594: The Mekong worm or Mekong giant earthworm , previously known as Megascolex mekongianus , is a species of earthworm in the family Megascolecidae . It is native to the vicinity of the River Mekong in southeastern Asia and may have more than 500 segments and grow to a length of 2.9 m (10 ft). The Mekong giant earthworm may grow to a length of up to 2.9 m (10 ft). Compared to their great length, these worms are relatively slender. The type specimen
5207-490: The aeration of the soil and the ability of water to infiltrate and to be held within the soil. Soil water content can be measured as volume or weight . Soil moisture levels, in order of decreasing water content, are saturation, field capacity , wilting point , air dry, and oven dry. Field capacity describes a drained wet soil at the point water content reaches equilibrium with gravity. Irrigating soil above field capacity risks percolation losses. Wilting point describes
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5334-561: The humus form ), the copedon (in intermediary position, where most weathering of minerals takes place) and the lithopedon (in contact with the subsoil). The soil texture is determined by the relative proportions of the individual particles of sand , silt , and clay that make up the soil. The interaction of the individual mineral particles with organic matter, water, gases via biotic and abiotic processes causes those particles to flocculate (stick together) to form aggregates or peds . Where these aggregates can be identified,
5461-496: The megascolecid earthworms is in chaos." Over the years, many scientists have 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 which was developed by Blakemore (2000), is a modern reversion to the Classical System that is historically proven and widely accepted. Categorization of
5588-425: The mineralogy of those particles can strongly modify those properties. The mineralogy of the finest soil particles, clay, is especially important. Large numbers of microbes , animals , plants and fungi are living in soil. However, biodiversity in soil is much harder to study as most of this life is invisible, hence estimates about soil biodiversity have been unsatisfactory. A recent study suggested that soil
5715-618: The mouth , one on either side, connected to an axial nerve running along its length to motor neurons and sensory cells in each segment. Large numbers of chemoreceptors concentrate near its mouth. Circumferential and longitudinal muscles edging each segment let the worm move. Similar sets of muscles line the gut tube , and their actions propel digested food toward the worm's anus . Earthworms are hermaphrodites : each worm carries male and female reproductive organs and genital pores . When mating, two individual earthworms will exchange sperm and fertilize each other's ova . Depending on
5842-488: The semiaquatic families Tubificidae , Lumbricidae and Enchytraeidae . The megadriles are characterized by a distinct clitellum (more extensive than that of microdriles) and a vascular system with true capillaries . Earthworms are commonly found in moist, compost -rich soil, eating a wide variety of organic matters , which include detritus , living protozoa , rotifers , nematodes , bacteria , fungi and other microorganisms . An earthworm's digestive system runs
5969-448: The A and B horizons. The living component of the soil is largely confined to the solum, and is generally more prominent in the A horizon. It has been suggested that the pedon , a column of soil extending vertically from the surface to the underlying parent material and large enough to show the characteristics of all its horizons, could be subdivided in the humipedon (the living part, where most soil organisms are dwelling, corresponding to
6096-625: The CEC increases. Hence, pure sand has almost no buffering ability, though soils high in colloids (whether mineral or organic) have high buffering capacity . Buffering occurs by cation exchange and neutralisation . However, colloids are not the only regulators of soil pH. The role of carbonates should be underlined, too. More generally, according to pH levels, several buffer systems take precedence over each other, from calcium carbonate buffer range to iron buffer range. Amynthas mekongianus Megascolex mekongianus Amynthas mekongianus ,
6223-447: 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) published only a few of his findings. These nevertheless show it is theoretically possible to grow two whole worms from
6350-406: The acronym CROPT. The physical properties of soils, in order of decreasing importance for ecosystem services such as crop production , are texture , structure , bulk density , porosity , consistency, temperature , colour and resistivity . Soil texture is determined by the relative proportion of the three kinds of soil mineral particles, called soil separates: sand , silt , and clay . At
6477-446: The addition of any more hydronium ions or aluminum hydroxyl cations drives the pH even lower (more acidic) as the soil has been left with no buffering capacity. In areas of extreme rainfall and high temperatures, the clay and humus may be washed out, further reducing the buffering capacity of the soil. In low rainfall areas, unleached calcium pushes pH to 8.5 and with the addition of exchangeable sodium, soils may reach pH 10. Beyond
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#17327800128826604-452: The apparent sterility of tropical soils. Live plant roots also have some CEC, linked to their specific surface area. Anion exchange capacity is the soil's ability to remove anions (such as nitrate , phosphate ) from the soil water solution and sequester those for later exchange as the plant roots release carbonate anions to the soil water solution. Those colloids which have low CEC tend to have some AEC. Amorphous and sesquioxide clays have
6731-437: The atmosphere due to increased biological activity at higher temperatures, a positive feedback (amplification). This prediction has, however, been questioned on consideration of more recent knowledge on soil carbon turnover. Soil acts as an engineering medium, a habitat for soil organisms , a recycling system for nutrients and organic wastes , a regulator of water quality , a modifier of atmospheric composition , and
6858-551: 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 , meaning that growth and development of embryos happens without fertilization . Among lumbricid earthworms, parthenogenesis arose from sexual relatives many times. Parthenogenesis in some Aporrectodea trapezoides lineages arose 6.4 to 1.1 million years ago from sexual ancestors. A few species exhibit pseudogamous parthogenesis, meaning that mating
6985-471: 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-in-a-tube, divided into a series of segments (called metameres ) that compartmentalize the body. Furrows are generally externally visible on the body demarking the segments; dorsal pores and nephridiopores exude a fluid that moistens and protects
7112-426: The basic cations are forced off the soil colloids by the mass action of hydronium ions from usual or unusual rain acidity against those attached to the colloids. High rainfall rates can then wash the nutrients out, leaving the soil inhabited only by those organisms which are particularly efficient to uptake nutrients in very acid conditions, like in tropical rainforests . Once the colloids are saturated with H 3 O ,
7239-563: 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. Earthworms use, in addition to
7366-410: The bodies of their mates. 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
7493-757: The body ( peristalsis ). The shortened part is anchored to the surrounding soil by tiny clawlike 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. As a result of their movement through their lubricated tunnels, worms can make gurgling noises underground when disturbed. 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. Earthworms have
7620-433: The body and are therefore considered one giant axon. The sympathetic nervous system consists of nerve plexuses in the epidermis and alimentary canal. (A plexus is a web of connected nerve cells.) 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
7747-413: The body, but in reverse order—an inner circular layer within an outer longitudinal layer. Earthworms have 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
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#17327800128827874-401: 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 fibers) 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
8001-443: 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. Earthworms have no special respiratory organs. Gases are exchanged through
8128-431: The coelomic fluid to pass between segments. 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
8255-429: The cricopharyngeal connective. 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. Touching an earthworm, which causes
8382-506: The cuticle to keep the body moist and ease movement through the 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. Interior to the muscle layer is a fluid-filled chamber called a coelom 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") which are perforated transverse walls, allowing
8509-427: The digestive proteins, a class of surface active compounds called drilodefensins , which help digest plant material. Instead of being coiled like a mammalian intestine, in the earthworm's intestine a large mid-dorsal, tongue-like fold is present, called a typhlosole , with many folds running along its length, increasing its surface area to increase nutrient absorption. The intestine has its own pair of muscle layers like
8636-437: The digestive tract; the subneural vessel, which runs below the ventral nerve cord; and two lateroneural vessels on either side of the nerve cord. The dorsal vessel is mainly a collecting structure in the intestinal region. It receives a pair commissural and dorsal intestines in each segment. The ventral vessel branches off to a pair of ventro-tegumentaries and ventro-intestinals in each segment. The subneural vessel also gives out
8763-436: The dry limit for growing plants. During growing season, soil moisture is unaffected by functional groups or specie richness. Available water capacity is the amount of water held in a soil profile available to plants. As water content drops, plants have to work against increasing forces of adhesion and sorptivity to withdraw water. Irrigation scheduling avoids moisture stress by replenishing depleted water before stress
8890-475: 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. Enkephalin and α-endorphin -like substances have been found in earthworms. Injections of naloxone (an opioid antagonist) inhibit
9017-430: The epithelium of the buccal chamber. These receptors are gustatory and olfactory (related to taste and smell). They also respond to chemical stimuli. (Chemoreceptor) The gut of the earthworm is a straight tube that extends from the worm's mouth to its anus . It is differentiated into an alimentary canal and associated glands which are embedded in the wall of the alimentary canal itself. The alimentary canal consists of
9144-402: The escape responses of earthworms. This indicates that opioid substances play a role in sensory modulation, similar to that found in many vertebrates. Although some worms have eyes , earthworms do not. However, they do have specialized photosensitive cells called "light cells of Hess". These photoreceptor cells have a central intracellular cavity ( phaosome ) filled with microvilli . As well as
9271-422: 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 to 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
9398-424: The female. Theoretical cladistic studies have placed them in the suborder Lumbricina of the order Haplotaxida , but this may change. Other slang names for earthworms include "dew-worm", "rainworm", "nightcrawler", and "angleworm" (from its use as angling hookbaits ). Larger terrestrial earthworms are also called megadriles (which translates to "big worms") as opposed to the microdriles ("small worms") in
9525-482: 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 narrower 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
9652-399: The first three and the last ones. 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 the fourth, fifth and sixth segments. The waste in
9779-527: The formation, description (morphology), and classification of soils in their natural environment. In engineering terms, soil is included in the broader concept of regolith , which also includes other loose material that lies above the bedrock, as can be found on the Moon and other celestial objects . Soil is a major component of the Earth 's ecosystem . The world's ecosystems are impacted in far-reaching ways by
9906-419: The highest AEC, followed by the iron oxides. Levels of AEC are much lower than for CEC, because of the generally higher rate of positively (versus negatively) charged surfaces on soil colloids, to the exception of variable-charge soils. Phosphates tend to be held at anion exchange sites. Iron and aluminum hydroxide clays are able to exchange their hydroxide anions (OH ) for other anions. The order reflecting
10033-632: The infiltration and movement of air and water, both of which are critical for life existing in soil. Compaction , a common problem with soils, reduces this space, preventing air and water from reaching plant roots and soil organisms. Given sufficient time, an undifferentiated soil will evolve a soil profile that consists of two or more layers, referred to as soil horizons. These differ in one or more properties such as in their texture , structure , density , porosity, consistency, temperature, color, and reactivity . The horizons differ greatly in thickness and generally lack sharp boundaries; their development
10160-442: The length of its body. They are one of nature's most important detritivores and coprophages , and also serve as food for many low-level consumers within the ecosystems. Earthworms exhibit an externally segmented tube-within-a-tube body plan with corresponding internal segmentations, and usually have setae on all segments. They have a cosmopolitan distribution wherever soil, water and temperature conditions allow. They have
10287-448: The microvilli, there are several sensory cilia in the phaosome which are structurally independent of the microvilli. 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 first segment. They are most numerous in the prostomium, and reduce in density in the first three segments; they are very few in number past
10414-415: 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. 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." Also, while, as
10541-414: The moist skin and capillaries, where the oxygen is picked up by the haemoglobin 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. At birth, earthworms emerge small but fully formed, lacking only their sex structures which develop in about 60 to 90 days. They attain full size in about one year. Scientists predict that
10668-590: The name suggests, the main habitat of earthworms is in soil, they are not restricted to this habitat. 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
10795-429: The negatively charged colloids resist being washed downward by water and are out of reach of plant roots, thereby preserving the soil fertility in areas of moderate rainfall and low temperatures. There is a hierarchy in the process of cation exchange on colloids, as cations differ in the strength of adsorption by the colloid and hence their ability to replace one another ( ion exchange ). If present in equal amounts in
10922-412: The negatively-charged soil colloid exchange sites (CEC) that are occupied by base-forming cations is called base saturation . If a soil has a CEC of 20 meq and 5 meq are aluminium and hydronium cations (acid-forming), the remainder of positions on the colloids ( 20 − 5 = 15 meq ) are assumed occupied by base-forming cations, so that the base saturation is 15 ÷ 20 × 100% = 75% (the compliment 25%
11049-419: 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. 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
11176-419: The next larger scale, soil structures called peds or more commonly soil aggregates are created from the soil separates when iron oxides , carbonates , clay, silica and humus , coat particles and cause them to adhere into larger, relatively stable secondary structures. Soil bulk density , when determined at standardized moisture conditions, is an estimate of soil compaction . Soil porosity consists of
11303-612: The numbers of worms that go into diapause . The more acidic the soil, the sooner worms go into diapause, and remain in diapause the longest time at a pH of 6.4. Earthworms are preyed upon by many species of birds (e.g. robins, starlings , thrushes , gulls , crows ), snakes, wood turtles, mammals (e.g. bears , boars, foxes , hedgehogs , pigs , moles ) and invertebrates (e.g. ants , flatworms , ground beetles and other beetles , snails , spiders , and slugs ). Earthworms have many internal parasites , including protozoa , platyhelminthes, mites, and nematodes ; they can be found in
11430-472: The ocean has no more than 10 prokaryotic organisms per milliliter (gram) of seawater. Organic carbon held in soil is eventually returned to the atmosphere through the process of respiration carried out by heterotrophic organisms, but a substantial part is retained in the soil in the form of soil organic matter; tillage usually increases the rate of soil respiration , leading to the depletion of soil organic matter. Since plant roots need oxygen, aeration
11557-635: The other cations more weakly bound to colloids are pushed into solution as hydrogen ions occupy exchange sites ( protonation ). A low pH may cause the hydrogen of hydroxyl groups to be pulled into solution, leaving charged sites on the colloid available to be occupied by other cations. This ionisation of hydroxy groups on the surface of soil colloids creates what is described as pH-dependent surface charges. Unlike permanent charges developed by isomorphous substitution , pH-dependent charges are variable and increase with increasing pH. Freed cations can be made available to plants but are also prone to be leached from
11684-492: The plant roots release hydrogen ions to the solution. CEC is the amount of exchangeable hydrogen cation (H ) that will combine with 100 grams dry weight of soil and whose measure is one milliequivalents per 100 grams of soil (1 meq/100 g). Hydrogen ions have a single charge and one-thousandth of a gram of hydrogen ions per 100 grams dry soil gives a measure of one milliequivalent of hydrogen ion. Calcium, with an atomic weight 40 times that of hydrogen and with
11811-432: 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. The excretory system contains a pair of nephridia in every segment, except for
11938-486: The porous lava, and by these means organic matter and a finer mineral soil accumulate with time. Such initial stages of soil development have been described on volcanoes, inselbergs, and glacial moraines. How soil formation proceeds is influenced by at least five classic factors that are intertwined in the evolution of a soil: parent material, climate, topography (relief), organisms, and time. When reordered to climate, relief, organisms, parent material, and time, they form
12065-414: The processes carried out in the soil, with effects ranging from ozone depletion and global warming to rainforest destruction and water pollution . With respect to Earth's carbon cycle , soil acts as an important carbon reservoir , and it is potentially one of the most reactive to human disturbance and climate change . As the planet warms, it has been predicted that soils will add carbon dioxide to
12192-414: The rate of diffusion of gases into and out of soil. Platy soil structure and soil compaction (low porosity) impede gas flow, and a deficiency of oxygen may encourage anaerobic bacteria to reduce (strip oxygen) from nitrate NO 3 to the gases N 2 , N 2 O, and NO, which are then lost to the atmosphere, thereby depleting the soil of nitrogen, a detrimental process called denitrification . Aerated soil
12319-788: The rest. As the worm slips out of the ring, the ends of the cocoon seal to form a vaguely onion-shaped incubator ( cocoon ) in which the embryonic worms develop. Hence fertilization is external. The cocoon is then deposited in the soil. After three weeks, 2 to 20 offspring hatch with an average of four. Development is direct i.e. without formation of any larva. Exposure of the earthworm Eisenia fetida to ionizing radiation induced DNA strand breaks and oxidized DNA bases . These DNA damages could then be repaired in somatic and spermatogenic cells. Earthworms testis cells are also capable of repairing hydrogen peroxide induced oxidative DNA adducts. Earthworms travel underground by means of waves of muscular contractions which alternately shorten and lengthen
12446-453: 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. Soil Soil consists of a solid phase of minerals and organic matter (the soil matrix), as well as a porous phase that holds gases (the soil atmosphere) and water (the soil solution). Accordingly, soil is a three- state system of solids, liquids, and gases. Soil
12573-442: The seat of interaction networks playing a decisive role in the stability, dynamics and evolution of soil ecosystems. Biogenic soil volatile organic compounds are exchanged with the aboveground atmosphere, in which they are just 1–2 orders of magnitude lower than those from aboveground vegetation. Humans can get some idea of the soil atmosphere through the well-known 'after-the-rain' scent, when infiltering rainwater flushes out
12700-441: The soil anion exchange capacity. The cation exchange, that takes place between colloids and soil water, buffers (moderates) soil pH, alters soil structure, and purifies percolating water by adsorbing cations of all types, both useful and harmful. The negative or positive charges on colloid particles make them able to hold cations or anions, respectively, to their surfaces. The charges result from four sources. Cations held to
12827-538: The soil by volatilisation (loss to the atmosphere as gases) or leaching. Soil is said to be formed when organic matter has accumulated and colloids are washed downward, leaving deposits of clay, humus , iron oxide , carbonate , and gypsum , producing a distinct layer called the B horizon. This is a somewhat arbitrary definition as mixtures of sand, silt, clay and humus will support biological and agricultural activity before that time. These constituents are moved from one level to another by water and animal activity. As
12954-420: The soil solution composition (attenuate changes in the soil solution) as soils wet up or dry out, as plants take up nutrients, as salts are leached, or as acids or alkalis are added. Plant nutrient availability is affected by soil pH , which is a measure of the hydrogen ion activity in the soil solution. Soil pH is a function of many soil forming factors, and is generally lower (more acidic) where weathering
13081-402: The soil solution. Both living soil organisms (microbes, animals and plant roots) and soil organic matter are of critical importance to this recycling, and thereby to soil formation and soil fertility . Microbial soil enzymes may release nutrients from minerals or organic matter for use by plants and other microorganisms, sequester (incorporate) them into living cells, or cause their loss from
13208-473: The soil voids are saturated with water vapour, at least until the point of maximal hygroscopicity , beyond which a vapour-pressure deficit occurs in the soil pore space. Adequate porosity is necessary, not just to allow the penetration of water, but also to allow gases to diffuse in and out. Movement of gases is by diffusion from high concentrations to lower, the diffusion coefficient decreasing with soil compaction . Oxygen from above atmosphere diffuses in
13335-430: The soil water solution will be insufficient to change the pH, as the acid forming cations stored on the soil colloids will tend to restore the original pH condition as they are pushed off those colloids by the calcium of the added lime. The resistance of soil to change in pH, as a result of the addition of acid or basic material, is a measure of the buffering capacity of a soil and (for a particular soil type) increases as
13462-433: The soil water solution: Al replaces H replaces Ca replaces Mg replaces K same as NH 4 replaces Na If one cation is added in large amounts, it may replace the others by the sheer force of its numbers. This is called law of mass action . This is largely what occurs with the addition of cationic fertilisers ( potash , lime ). As the soil solution becomes more acidic (low pH , meaning an abundance of H ),
13589-462: The soil where it is consumed and levels of carbon dioxide in excess of above atmosphere diffuse out with other gases (including greenhouse gases ) as well as water. Soil texture and structure strongly affect soil porosity and gas diffusion. It is the total pore space ( porosity ) of soil, not the pore size, and the degree of pore interconnection (or conversely pore sealing), together with water content, air turbulence and temperature, that determine
13716-502: The soil, a complex feedback which is exemplified in the dynamics of banded vegetation patterns in semi-arid regions. Soils supply plants with nutrients , most of which are held in place by particles of clay and organic matter ( colloids ) The nutrients may be adsorbed on clay mineral surfaces, bound within clay minerals ( absorbed ), or bound within organic compounds as part of the living organisms or dead soil organic matter. These bound nutrients interact with soil water to buffer
13843-414: The soil, possibly making the soil less fertile. Plants are able to excrete H into the soil through the synthesis of organic acids and by that means, change the pH of the soil near the root and push cations off the colloids, thus making those available to the plant. Cation exchange capacity is the soil's ability to remove cations from the soil water solution and sequester those to be exchanged later as
13970-500: 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 favour neutral to slightly acidic soils. Lumbricus terrestris is still present in a pH of 5.4, Dendrobaena octaedra at a pH of 4.3 and some Megascolecidae are present in extremely acidic humic soils. Soil pH may also influence
14097-548: The soil-litter interface and eat decomposing organic matter ( epigeic ) e.g. Eisenia fetida ; (2) topsoil- or subsoil-dwelling worms that feed (on soil), burrow and cast within the soil, creating horizontal burrows in upper 10–30 cm of soil ( endogeic ); 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 ( anecic , meaning "reaching up"), e.g. Lumbricus terrestris . Earthworm populations depend on both physical and chemical properties of
14224-402: 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. Probably the longest worm on confirmed records is Amynthas mekongianus that extends up to 3 m (10 ft) in the mud along
14351-837: 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 . Sex pheromones are probably important in earthworms because they live in an environment where chemical signaling may play
14478-409: The strength of anion adhesion is as follows: The amount of exchangeable anions is of a magnitude of tenths to a few milliequivalents per 100 g dry soil. As pH rises, there are relatively more hydroxyls, which will displace anions from the colloids and force them into solution and out of storage; hence AEC decreases with increasing pH (alkalinity). Soil reactivity is expressed in terms of pH and
14605-564: The third segment. These receptors are abundant and distributed all over the epidermis . Each receptor shows a slightly elevated cuticle which covers a group of tall, slender and columnar receptor cells. These cells bear small hairlike processes at their outer ends and their inner ends are connected with nerve fibres. The epidermal receptors are tactile in function. They are also concerned with changes in temperature and respond to chemical stimuli. Earthworms are extremely sensitive to touch and mechanical vibration. These receptors are located only in
14732-491: The two concentrations are equal, they are said to neutralise each other. A pH of 9.5 has 10 moles hydronium ions per litre of solution (and also 10 moles per litre OH ). A pH of 3.5 has one million times more hydronium ions per litre than a solution with pH of 9.5 ( 9.5 − 3.5 = 6 or 10 ) and is more acidic. The effect of pH on a soil is to remove from the soil or to make available certain ions. Soils with high acidity tend to have toxic amounts of aluminium and manganese . As
14859-473: The void part of the soil volume and is occupied by gases or water. Soil consistency is the ability of soil materials to stick together. Soil temperature and colour are self-defining. Resistivity refers to the resistance to conduction of electric currents and affects the rate of corrosion of metal and concrete structures which are buried in soil. These properties vary through the depth of a soil profile, i.e. through soil horizons . Most of these properties determine
14986-553: The water content of the soil, as can be expressed in terms of volume or weight—can be based on in situ probes (e.g., capacitance probes , neutron probes ), or remote sensing methods. Soil moisture measurement is an important factor in determining changes in soil activity. The atmosphere of soil, or soil gas , is very different from the atmosphere above. The consumption of oxygen by microbes and plant roots, and their release of carbon dioxide, decreases oxygen and increases carbon dioxide concentration. Atmospheric CO 2 concentration
15113-461: The weathering of lava flow bedrock, which would produce the purely mineral-based parent material from which the soil texture forms. Soil development would proceed most rapidly from bare rock of recent flows in a warm climate, under heavy and frequent rainfall. Under such conditions, plants (in a first stage nitrogen-fixing lichens and cyanobacteria then epilithic higher plants ) become established very quickly on basaltic lava, even though there
15240-476: The whole soil atmosphere after a drought period, or when soil is excavated, a bulk property attributed in a reductionist manner to particular biochemical compounds such as petrichor or geosmin . Soil particles can be classified by their chemical composition ( mineralogy ) as well as their size. The particle size distribution of a soil, its texture, determines many of the properties of that soil, in particular hydraulic conductivity and water potential , but
15367-405: The worm's sides; usually, two nephridia (sometimes more) are found in most segments. At the centre 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
15494-410: The worm's surface, allowing it to breathe. Except for the mouth and anal segments, each segment carries bristlelike hairs called lateral setae used to anchor parts of the body during movement; species may have four pairs of setae on each segment or more than eight sometimes forming a complete circle of setae per segment. Special ventral setae are used to anchor mating earthworms by their penetration into
15621-420: 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. Thus each worm becomes the genetic father of some of their offspring (due to its own sperm transferred to other earthworm) and the genetic mother (offsprings from its own egg cells) of
15748-503: The worms live. Pigs forage and fishermen dig worms for bait there. Unlike most other species of giant earthworm, which generally inhabit pastureland, the Mekong worm burrows into the muddy banks of the River Mekong in Laos . The worm forms complex networks of tunnels and brings large quantities of ingested soil to the surface in the form of worm castings . The worms may be collected from
15875-431: The worms' blood , seminal vesicles , coelom , or intestine , or in their cocoons (e.g. the mite Histiostoma murchiei is a parasite of earthworm cocoons). The 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
16002-657: Was immature and lacked a clitellum . The type specimen was recorded as being from "Ban Leum on Mekong River, Annam". This was mistakenly believed to refer to a village in Central Vietnam well away from the River Mekong, but it has been found that "Ban leum" means "[I] forget the place" in the local dialect, and the type locality is thought to be in Thailand or Laos. The river forms a delta at that point and contains extensive mud and sand flats and embankments where
16129-436: Was one metre long and 8 mm (0.3 in) wide at the broadest point (segment 5). It had 370 segments and was a greyish colour, rather paler on the ventral surface. The prostomium was poorly preserved. The second segment had 46 setae (bristles) in an incomplete ring with a gap on the ventral surface. Segments 3 to 25 bore about 100 setae each, arranged in a complete ring. The male pores were on segment 17, but this specimen
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