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For other uses, seeLeech (disambiguation).

Leech
Temporal range: MiddlePermian–recent,266–0MaPossibleVirgilianrecord
Hirudo medicinalissucking blood
Helobdellasp.
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Annelida
Clade: Pleistoannelida
Clade: Sedentaria
Class: Clitellata
Subclass: Hirudinea
Lamarck1818

Leechesare segmentedparasiticorpredatorywormsthat comprise thesubclassHirudineawithin thephylumAnnelida.They are closely related to theoligochaetes,which include theearthworm,and like them have soft, muscular segmented bodies that can lengthen and contract. Both groups arehermaphroditesand have aclitellum,but leeches typically differ from the oligochaetes in having suckers at both ends and in having ring markings that do not correspond with their internal segmentation. The body is muscular and relatively solid, and thecoelom,the spacious body cavity found in other annelids, is reduced to small channels.

The majority of leeches live in freshwater habitats, while some species can be found in terrestrial or marine environments. The best-known species, such as the medicinal leech,Hirudo medicinalis,arehematophagous,attaching themselves to a host with a sucker and feeding on blood, having first secreted the peptidehirudinto prevent the blood from clotting. The jaws used to pierce the skin are replaced in other species by aprobosciswhich is pushed into the skin. A minority of leech species are predatory, mostly preying on small invertebrates.

The eggs are enclosed in a cocoon, which in aquatic species is usually attached to an underwater surface; members of one family,Glossiphoniidae,exhibit parental care, the eggs being brooded by the parent. In terrestrial species, the cocoon is often concealed under a log, in a crevice or buried in damp soil. Almost seven hundred species of leech are currently recognised, of which some hundred are marine, ninety terrestrial and the remainder freshwater.

Leeches have been used inmedicinefrom ancient times until the 19th century todraw bloodfrom patients. In modern times, leeches find medical use in treatment of joint diseases such asepicondylitisand osteoarthritis, extremity vein diseases, and inmicrosurgery,while hirudin is used as ananticoagulantdrug to treat blood-clotting disorders.

The leech appears in the biblicalBook of Proverbsas an archetype of insatiablegreed.[1]The term "leech" is used to characterise a person who takes without giving, living at the expense of others.[2]

Diversity and phylogeny

A terrestrial leech, Haemadipsa zeylanica
Haemadipsa zeylanica,a terrestrial leech
Dorsal (upper) surface and ventral (lower) surface of Placobdelloides siamensis, ventral showing numerous young leeches
Placobdelloides siamensis,a parasite of turtles inThailand.The ventral face (right) shows many young leeches.[3]

Some 680 species of leech have been described, of which around 100 are marine, 480 freshwater and the remainder terrestrial.[4][5]AmongEuhirudinea,the true leeches, the smallest is about1 cm (1⁄2in) long, and the largest is the giant Amazonian leech,Haementeria ghilianii,which can reach 30 cm (12 in). Except for Antarctica,[4]leeches are found throughout the world but are at their most abundant in temperate lakes and ponds in the northern hemisphere. The majority of freshwater leeches are found in the shallow, vegetated areas on the edges of ponds, lakes and slow-moving streams; very few species tolerate fast-flowing water. In their preferred habitats, they may occur in very high densities; in a favourable environment with water high in organic pollutants, over 10,000 individuals were recorded per square metre (over 930 per square foot) under rocks inIllinois.Some speciesaestivateduring droughts, burying themselves in the sediment, and can lose up to 90% of their bodyweight and still survive.[6]Among the freshwater leeches are theGlossiphoniidae,dorso-ventrally flattened animals mostly parasitic on vertebrates such as turtles, and unique amongannelidsin both brooding their eggs and carrying their young on the underside of their bodies.[7]

The terrestrialHaemadipsidaeare mostly native to the tropics and subtropics,[8]while the aquaticHirudinidaehave a wider global range; both of these feed largely on mammals, including humans.[6]A distinctive family is thePiscicolidae,marine or freshwaterectoparasiteschiefly of fish, with cylindrical bodies and usually well-marked, bell-shaped, anterior suckers.[9]Not all leeches feed on blood; theErpobdelliformes,freshwater or amphibious, are carnivorous and equipped with a relatively large, toothless mouth to ingest insect larvae, molluscs, and other annelid worms, which are swallowed whole.[10]In turn, leeches are prey to fish, birds, and invertebrates.[11]

The name for the subclass, Hirudinea, comes from the Latinhirudo(genitivehirudinis), a leech; the element-bdellafound in many leech group names is from the Greek βδέλλαbdella,also meaning leech.[12]The nameLes hirudinéeswas given byJean-Baptiste Lamarckin 1818.[13]Leeches were traditionally divided into two infraclasses, theAcanthobdellidea(primitive leeches) and the Euhirudinea (true leeches).[14]The Euhirudinea are divided into the proboscis-bearingRhynchobdellidaand the rest, including some jawed species, the "Arhynchobdellida",without a proboscis.[15]

Thephylogenetic treeof the leeches and their annelid relatives is based onmolecular analysis(2019) of DNA sequences. Both the former classes "Polychaeta"(bristly marine worms) and"Oligochaeta"(including the earthworms) areparaphyletic:in each case the complete groups (clades) would include all the other groups shown below them in the tree. TheBranchiobdellidaare sister to the leech clade Hirudinida, which approximately corresponds to the traditional subclass Hirudinea. The main subdivision of leeches is into the Rhynchobdellida and the Arhynchobdellida, though theAcanthobdellaare sister to the clade that contains these two groups.[15]

Annelida

Evolution

Fossil of a possible leech found in Wisconsin
Fossil of a worm that was once considered as leech but denied, from theWaukesha Biota,in theSilurianofWisconsin

The most ancient annelid group consists of the free-living polychaetes that evolved in theCambrianperiod, being plentiful in theBurgess Shaleabout 500million years ago. Oligochaetes evolved from polychaetes and the leeches branched off from the oligochaetes.[16]The oldest leech fossils are from the middlePermianperiod around 266million years ago,[17]there is also unpublished study about possible leech fromVirgilian(LateCarboniferous) ofNew Mexico.[18]Although fossil with external ring markings found fromSilurianstrata inWisconsinis sometimes identified as leech,[19][20]but assignment of fossil is still putative and contentious,[21][17]and the animal was also alternatively interpreted as a member ofCycloneuralia.[22][18]

Anatomy and physiology

Leeches show a remarkable similarity to each other in morphology, very different from typical annelids which are cylindrical with a fluid-filled space, thecoelom(body cavity). In leeches, most of the coelom is filled with botryoidal tissue, a loose connective tissue composed of clusters of cells of mesodermal origin.[23]The remaining body cavity has been reduced to four slender longitudinal channels. Typically, the body isdorso-ventrallyflattened and tapers at both ends. Longitudinal and circular muscles in the body wall are supplemented by diagonal muscles, giving the leech the ability to adopt a large range of body shapes and show great flexibility. Most leeches have a sucker at both the anterior (front) and posterior (back) ends, but some primitive leeches have a single sucker at the back.[24][25]

Cross-section of a leech showing its anatomy
Leech anatomy in cross-section: the body is solid, thecoelom(body cavity) reduced to channels, with circular, longitudinal, and transverse muscles making the animal strong and flexible.[26]

Like most annelids, with a few exceptions likeSipuncula,EchiuraandDiurodrilus,[27]the leech is a segmented animal, but unlike other annelids, thesegmentationis masked by secondary external ring markings (annuli).[28]The number of annulations varies, both between different regions of the body and between species.[24]In one species, the body surface is divided into 102annuli.[29]All leech species, however, have 32 segments, called somites, (34 if two head segments, which have different organization, are counted).[30][31]Of these segments, the first five are designated as the head and include the anterior brain, severalocelli(eyespots) dorsally and the sucker ventrally. The following 21 mid-body segments each contain a nerveganglion,and between them contain two reproductive organs, a single femalegonoporeand nine pairs oftestes.The last seven segments contain the posterior brain and are fused to form the animal's tail sucker.[24]Theseptathat separates the body segments—and themesenterieswhich in turn separates each segment into a left and right half—in the majority of annelids, have been lost in leeches except for the primitive genusAcanthobdella,which still have some septa and mesenteries.[30][32]

The body wall consists of acuticle,anepidermisand a thick layer of fibrousconnective tissuein which are embedded the circular muscles, the diagonal muscles and the powerful longitudinal muscles. There are also dorso-ventral muscles. In leeches the originalblood vascular systemhas been lost and replaced by the modified coelom known as the haemocoelomic system, and the coelomic fluid, called the haemocoelomic fluid, has taken over the role as blood. The haemocoelomic channels run the full length of the body, the two main ones being on either side.[33]Part of the liningepitheliumconsists ofchloragogen cellswhich are used for the storage of nutrients and inexcretion.There are 10 to 17 pairs ofmetanephridia(excretory organs) in the mid-region of the leech. From these, ducts typically lead to aurinary bladder,which empties to the outside at anephridiopore.[26]

Reproduction and development

Further information:Leech embryogenesis

Leeches are hermaphrodites, with the malereproductive organs,thetestes,maturing first and theovarieslater. In hirudinids, a pair will line up with theclitellarregions in contact, with the anterior end of one leech pointing towards the posterior end of the other; this results in the male gonopore of one leech being in contact with the female gonopore of the other. The penis passes aspermatophoreinto the female gonopore and sperm is transferred to, and probably stored in, the vagina.[34]

Somejawless leeches(Rhynchobdellida) andproboscisless leeches(Arhynchobdellida) lack a penis, and in these, sperm is passed from one individual to another byhypodermic injection.The leeches intertwine and grasp each other with their suckers. A spermatophore is pushed by one through the integument of the other, usually into the clitellar region. The sperm is liberated and passes to the ovisacs, either through the coelomic channels or interstitially through specialist "target tissue" pathways.[34]

Some time after copulation, the small, relatively yolkless eggs are laid. In most species, analbumin-filled cocoon is secreted by the clitellum and receives one or more eggs as it passes over the female gonopore.[34]In the case of the North AmericanErpobdella punctata,the clutch size is about five eggs, and some ten cocoons are produced.[35]Each cocoon is fixed to a submerged object, or in the case of terrestrial leeches, deposited under a stone or buried in damp soil. The cocoon ofHemibdella soleaeis attached to a suitable fishhost.[34][36]The glossiphoniids brood their eggs, either by attaching the cocoon to the substrate and covering it with their ventral surface, or by securing the cocoon to their ventral surface, and even carrying the newly hatched young to their first meal.[37]

When breeding, most marine leeches leave their hosts and become free-living in estuaries. Here they produce their cocoons, after which the adults of most species die. When the eggs hatch, the juveniles seek out potential hosts when these approach the shore.[37]Leeches mostly have an annual or biannual life cycle.[34]

Feeding and digestion

About three quarters of leech species areparasitesthat feed on the blood of a host, while the remainder arepredators.Leeches either have a pharynx that they can protrude, commonly called a proboscis, or a pharynx that they cannot protrude, which in some groups is armed with jaws.[38]

In the proboscisless leeches, the jaws (if any) of Arhynchobdellids are at the front of the mouth, and have three blades set at an angle to each other. In feeding, these slice their way through the skin of the host, leaving a Y-shaped incision. Behind the blades is the mouth, located ventrally at the anterior end of the body. It leads successively into thepharynx,a short oesophagus, acrop(in some species), a stomach and a hindgut, which ends at an anus located just above the posterior sucker. The stomach may be a simple tube, but the crop, when present, is an enlarged part of the midgut with a number of pairs ofcecathat store ingested blood. The leech secretes an anticoagulant,hirudin,in itssalivawhich prevents the blood from clotting before ingestion.[38]A maturemedicinal leechmay feed only twice a year, taking months to digest a blood meal.[25]

Magnified reddish leech bites on a cow's udder
Leech bites on a cow's udder

The bodies of predatory leeches are similar, though instead of a jaw many have a protrusibleproboscis,which for most of the time they keep retracted into the mouth. Such leeches are oftenambush predatorsthat lie in wait until they can strike prey with the proboscises in a spear-like fashion.[39]Predatory leeches feed on small invertebrates such as snails, earthworms and insect larvae. The prey is usually sucked in and swallowed whole. Some Rhynchobdellida however suck the soft tissues from their prey, making them intermediate between predators and blood-suckers.[38]

A leech attacking a slug's underside
Leech attacking a slug

Blood-sucking leeches use their anterior suckers to connect to hosts for feeding. Once attached, they use a combination of mucus and suction to stay in place while they inject hirudin into the hosts'blood.In general,blood-feedingleeches are non host-specific, and do little harm to their host, dropping off after consuming a blood meal. Some marine species however remain attached until it is time to reproduce. If present in great numbers on a host, these can be debilitating, and in extreme cases, cause death.[37]

Leeches are unusual in that they do not produce certaindigestive enzymessuch asamylases,lipasesorendopeptidases.[38]A deficiency of these enzymes and ofB complex vitaminsis compensated for by enzymes and vitamins produced byendosymbioticmicroflora. InHirudo medicinalis,these supplementary factors are produced by an obligatorymutualistic relationshipwith the bacterial species,Aeromonas veronii.Non-bloodsucking leeches, such asErpobdella octoculata,are host to more bacterial symbionts.[40]In addition, leeches produce intestinalexopeptidaseswhich remove amino acids from the long protein molecules one by one, possibly aided byproteasesfrom endosymbiotic bacteria in the hindgut.[41]This evolutionary choice of exopeptic digestion in Hirudinea distinguishes these carnivorous clitellates from oligochaetes, and may explain why digestion in leeches is so slow.[38]

Nervous system

A leech's nervous system is formed of a few largenerve cells.Their large size makes leeches convenient asmodel organismsfor the study of invertebrate nervous systems. The main nerve centre consists of the cerebral ganglion above the gut and another ganglion beneath it, with connecting nerves forming a ring around the pharynx a little way behind the mouth. A nerve cord runs backwards from this in the ventral coelomic channel, with 21 pairs ofganglia in segmentssix to 26. In segments 27 to 33, other paired ganglia fuse to form the caudal ganglion.[42]Several sensory nerves connect directly to the cerebral ganglion; there are sensory and motor nerve cells connected to the ventral nerve cord ganglia in each segment.[25]

Leeches have between two and tenpigment spot ocelli,arranged in pairs towards the front of the body. There are also sensory papillae arranged in a lateral row in one annulation of each segment. Each papilla contains many sensory cells. Some rhynchobdellids have the ability to change colour dramatically by moving pigment inchromatophorecells; this process is under the control of the nervous system but its function is unclear as the change in hue seems unrelated to the colour of the surroundings.[42]

Leeches can detect touch, vibration, movement of nearby objects, and chemicals secreted by their hosts; freshwater leeches crawl or swim towards a potential host standing in their pond within a few seconds. Species that feed on warm-blooded hosts move towards warmer objects. Many leeches avoid light, though some blood feeders move towards light when they are ready to feed, presumably increasing the chances of finding a host.[25]

Gas exchange

Leeches live in damp surroundings and in general respire through their body wall. The exception to this is in the Piscicolidae, where branching or leaf-like lateral outgrowths from the body wall formgills.Some rhynchobdellid leeches have an extracellularhaemoglobinpigment, but this only provides for about half of the leech's oxygen transportation needs, the rest occurring by diffusion.[26]

Movement

Leeches move using their longitudinal and circular muscles in a modification of the locomotion byperistalsis,self-propulsion by alternately contracting and lengthening parts of the body, seen in other annelids such as earthworms. They use their posterior and anterior suckers (one on each end of the body) to enable them toprogress by loopingor inching along, in the manner ofgeometer mothcaterpillars. The posterior end is attached to the substrate, and the anterior end is projected forward peristaltically by the circular muscles until it touches down, as far as it can reach, and the anterior end is attached. Then the posterior end is released, pulled forward by the longitudinal muscles, and reattached; then the anterior end is released, and the cycle repeats.[43][25]Leeches explore their environment with head movements and body waving.[44]The Hirudinidae andErpobdellidaecan swim rapidly with up-and-down or sideways undulations of the body; the Glossiphoniidae in contrast are poor swimmers and curl up and fall to the sediment below when disturbed.[45]

Interactions with humans

Hand preparing to grip a leech and remove it from the top of a foot
Leeches can be removed by hand, since they do not burrow into the skin or leave the head in the wound.[46][47]

Bites

Leech bites are generally alarming rather than dangerous, though a small percentage of people have severe allergic oranaphylactic reactionsand require urgent medical care. Symptoms of these reactions include red blotches or an itchy rash over the body, swelling around the lips or eyes, a feeling of faintness or dizziness, and difficulty in breathing.[48]An externally attached leech will detach and fall off on its own accord when it is satiated on blood, which may take from twenty minutes to a few hours; bleeding from the wound may continue for some time.[48]Internal attachments, such as inside the nose, are more likely to require medical intervention.[49]

Bacteria, viruses, and protozoan parasites from previous blood sources can survive within a leech for months, so leeches could potentially act as vectors of pathogens. Nevertheless, only a few cases of leeches transmitting pathogens to humans have been reported.[50][51]

Leech saliva is commonly believed to containanaestheticcompounds to numb the bite area, but some authorities disagree.[52][53][54]Althoughmorphine-like substances have been found in leeches, they have been found in theneural tissues,not the salivary tissues. They are used by the leeches in modulating their ownimmunocytesand not for anaesthetising bite areas on their hosts.[55][52]Depending on the species and size, leech bites can be barely noticeable or they can be fairly painful.[56][57]

Medical use

Further information:HirudotherapyandHirudo medicinalis § Current

The medicinal leechHirudo medicinalis,and some otherspecies,have been used for clinicalbloodlettingfor at least 2,500 years:Ayurvedictexts describe their use for bloodletting in ancient India. Inancient Greece,bloodletting was practised according to thetheory of humoursfound in theHippocratic Corpusof the fifth centuryBC, which maintained that health depended on a balance of the four humours: blood,phlegm,black bileandyellow bile.Bloodletting using leeches enabled physicians to restore balance if they considered blood was present in excess.[58][59]

Pliny the Elderreported in hisNatural Historythat thehorse leechcould drive elephants mad by climbing up inside their trunks to drink blood.[60]Pliny also noted the medicinal use of leeches inancient Rome,stating that they were often used forgout,and that patients became addicted to the treatment.[61]InOld English,lǣcewas the name for a physician as well as for the animal, though the words had different origins, andlǣcecraft,leechcraft, was the art of healing.[62]

William Wordsworth's 1802 poem "Resolution and Independence"describes one of the last of the leech-gatherers, people who travelled Britain catching leeches from the wild, and causing a sharp decline in their abundance, though they remain numerous inRomney Marsh.By 1863, British hospitals had switched to imported leeches, some seven million being imported to hospitals in London that year.[60]

In the nineteenth century, demand for leeches was sufficient for hirudiculture, the farming of leeches, to become commercially viable.[63]Leech usage declined with the demise of humoral theory,[64]but made a small-scale comeback in the 1980s after years of decline, with the advent ofmicrosurgery,where venous congestion can arise due to inefficient venous drainage. Leeches can reduce swelling in the tissues and promote healing, helping in particular to restore circulation after microsurgery to reattach body parts.[65][66]Other clinical applications includevaricose veins,muscle cramps,thrombophlebitis,and joint diseases such as epicondylitis andosteoarthritis.[67][68][69][70]

Leech secretions contain several bioactive substances withanti-inflammatory,anticoagulantand antimicrobial effects.[69]One active component of leech saliva is a small protein, hirudin.[71]It is widely used as an anticoagulant drug to treat blood-clotting disorders, and manufactured byrecombinant DNAtechnology.[72][73]

In 2012 and 2018, Ida Schnell and colleagues trialled the use ofHaemadipsaleeches to gather data on the biodiversity of theirmammalianhosts in thetropical rainforestof Vietnam, where it is hard to obtain reliable data on rare andcrypticmammals. They showed that mammalmitochondrial DNA,amplified by thepolymerase chain reaction,can be identified from a leech's blood meal for at least four months after feeding. They detectedAnnamite striped rabbit,small-toothed ferret-badger,Truong Son muntjac,andserowin this way.[74][75]

Water pollution

Exposure tosynthetic estrogenas used in contraceptive medicines, which may enter freshwater ecosystems from municipalwastewater,can affect leeches' reproductive systems. Although not as sensitive to these compounds as fish, leeches showed physiological changes after exposure, including longer sperm sacs andvaginal bulbs,and decreasedepididymisweight.[76]

Notes

  1. ^The caption below the lithograph reads "There's redundancy of blood andhumours,we'll bleed you to-morrow, till then, very little food. "

References

  1. ^"Proverbs 30:15 | Ellicott's Commentary for English Readers".BibleHub.Retrieved27 April2018.
  2. ^"Leech".Merriam-Webster.Retrieved27 April2018.
  3. ^Chiangkul, Krittiya; Trivalairat, Poramad; Purivirojkul, Watchariya (2018)."Redescription of the Siamese shield leechPlacobdelloides siamensiswith new host species and geographic range ".Parasite.25:56.doi:10.1051/parasite/2018056.ISSN1776-1042.PMC6254108.PMID30474597.
  4. ^abSket, Boris; Trontelj, Peter (2008). "Global diversity of leeches (Hirudinea) in freshwater".Hydrobiologia.595(1): 129–137.doi:10.1007/s10750-007-9010-8.S2CID46339662.
  5. ^Fogden, S.; Proctor, J. (1985). "Notes on the Feeding of Land Leeches (Haemadipsa zeylanicaMoore andH. pictaMoore) in Gunung Mulu National Park, Sarawak ".Biotropica.17(2): 172–174.Bibcode:1985Biotr..17..172F.doi:10.2307/2388511.JSTOR2388511.
  6. ^abRuppert, Fox & Barnes 2004,p. 471
  7. ^Siddall, Mark E. (1998)."Glossiphoniidae".American Museum of Natural History.Archived fromthe originalon 9 August 2019.Retrieved1 May2018.
  8. ^Ruppert, Fox & Barnes 2004,p. 480
  9. ^Meyer, Marvin C. (July 1940). "A Revision of the Leeches (Piscicolidae) Living on Fresh-Water Fishes of North America".Transactions of the American Microscopical Society.59(3): 354–376.doi:10.2307/3222552.JSTOR3222552.
  10. ^Oceguera, A.; Leon, V.; Siddall, M. (2005)."Phylogeny and revision of Erpobdelliformes (Annelida, Arhynchobdellida) from Mexico based on nuclear and mitochondrial gene sequences".Revista Mexicana de Biodiversidad.76(2): 191–198.doi:10.22201/ib.20078706e.2005.002.307.Retrieved23 October2020.
  11. ^"Leeches".Australian Museum. 14 November 2019.Retrieved3 June2020.
  12. ^Scarborough, John (1992).Medical and Biological Terminologies: Classical Origins.University of Oklahoma Press. p. 58.ISBN978-0-8061-3029-3.
  13. ^Lamarck, Jean-Baptiste(1818).Histoire naturelle des animaux sans vertèbres... précédée d'une introduction offrant la détermination des caractères essentiels de l'animal, sa distinction du végétal et des autres corps naturels, enfin, l'exposition des principes fondamentaux de la zoologie. Volume 5.Vol. 5. Paris: Verdière.
  14. ^Kolb, Jürgen (2018)."Hirudinea".WoRMS.World Register of Marine Species.Retrieved7 May2018.
  15. ^abPhillips, Anna J.; Dornburg, Alex; Zapfe, Katerina L.; et al. (2019)."Phylogenomic Analysis of a Putative Missing Link Sparks Reinterpretation of Leech Evolution".Genome Biology and Evolution.11(11): 3082–3093.doi:10.1093/gbe/evz120.ISSN1759-6653.PMC6598468.PMID31214691.
  16. ^Margulis, Lynn;Chapman, Michael J. (2009).Kingdoms and Domains: An Illustrated Guide to the Phyla of Life on Earth.Academic Press.p. 308.ISBN978-0-08-092014-6.
  17. ^abPrevec, Rosemary; Nel, André; Day, Michael O.; et al. (30 October 2022)."South African Lagerstätte reveals middle Permian Gondwanan lakeshore ecosystem in exquisite detail".Communications Biology.5(1): 1154.doi:10.1038/s42003-022-04132-y.ISSN2399-3642.PMC9618562.PMID36310243.
  18. ^abBraddy, Simon J.; Gass, Kenneth C.; Tessler, Michael (2023)."Not the first leech: An unusual worm from the early Silurian of Wisconsin".Journal of Paleontology.97(4): 799–804.Bibcode:2023JPal...97..799B.doi:10.1017/jpa.2023.47.
  19. ^Thorp, James H.; Covich, Alan P. (2001).Ecology and Classification of North American Freshwater Invertebrates.Academic Press.p. 466.ISBN978-0-12-690647-9.
  20. ^Mikulic, D. G.; Briggs, D. E. G.; Kluessendorf, J. (1985). "A new exceptionally preserved biota from the Lower Silurian of Wisconsin, U.S.A.".Philosophical Transactions of the Royal Society of London B.311(1148): 75–85.Bibcode:1985RSPTB.311...75M.doi:10.1098/rstb.1985.0140.
  21. ^Wendruff, Andrew J.; Babcock, Loren E.; Kluessendorf, Joanne; et al. (15 May 2020)."Paleobiology and taphonomy of exceptionally preserved organisms from the Waukesha Biota (Silurian), Wisconsin, USA".Palaeogeography, Palaeoclimatology, Palaeoecology.546:109631.Bibcode:2020PPP...54609631W.doi:10.1016/j.palaeo.2020.109631.ISSN0031-0182.S2CID212824469.
  22. ^Shcherbakov, Dmitry; Tarmo, Timm; Tzetlin, Alexander B.; Vinn, Olev; Zhuravlev, Andrey (2020)."A probable oligochaete from an Early Triassic Lagerstätte of the southern Cis-Urals and its evolutionary implications".Acta Palaeontologica Polonica.65(2): 219–233.doi:10.4202/app.00704.2019.S2CID219097612.
  23. ^Ultrastructure and functional versatility of hirudinean botryoidal tissue
  24. ^abcRuppert, Fox & Barnes 2004,pp. 471–472
  25. ^abcdeBrusca, Richard (2016). "Hirudinoidea: Leeches and Their Relatives".Invertebrates.Sinauer Associates.pp. 591–597.ISBN978-1-60535-375-3.
  26. ^abcRuppert, Fox & Barnes 2004,pp. 474–475
  27. ^Laumer, Christopher E.; Bekkouche, Nicolas; Kerbl, Alexandra; Goetz, Freya; Neves, Ricardo C.; Sørensen, Martin V.; Kristensen, Reinhardt M.; Hejnol, Andreas; Dunn, Casey W.; Giribet, Gonzalo; Worsaae, Katrine (August 2015)."Spiralian Phylogeny Informs the Evolution of Microscopic Lineages".Current Biology.25(15): 2000–2006.Bibcode:2015CBio...25.2000L.doi:10.1016/j.cub.2015.06.068.
  28. ^Buchsbaum, Ralph; Buchsbaum, Mildred; Pearse, John; et al. (1987).Animals Without Backbones(3rd ed.). TheUniversity of Chicago Press.pp. 312–317.ISBN978-0-226-07874-8.
  29. ^Payton, Brian (1981). Muller, Kenneth; Nicholls, John; Stent, Gunther (eds.).Neurobiology of the Leech.Cold Spring Harbor Laboratory.pp. 35–50.ISBN978-0-87969-146-2.
  30. ^abKuo, Dian-Han; Lai, Yi-Te (4 November 2018)."On the origin of leeches by evolution of development".Development, Growth & Differentiation.61(1): 43–57.doi:10.1111/dgd.12573.PMID30393850.S2CID53218704.
  31. ^Castle, W. E. (1900)."The Metamerism of the Hirudinea".Proceedings of the American Academy of Arts and Sciences.35(15): 285–303.doi:10.2307/25129933.ISSN0199-9818.JSTOR25129933.
  32. ^Reproductive Strategies and Developmental Patterns in Annelids
  33. ^Modern Text Book of Zoology: Invertebrates
  34. ^abcdeRuppert, Fox & Barnes 2004,pp. 477–478
  35. ^Sawyer, R. T. (1970). "Observations on the Natural History and Behavior ofErpobdella punctata(Leidy) (Annelida: Hirudinea) ".The American Midland Naturalist.83(1): 65–80.doi:10.2307/2424006.JSTOR2424006.
  36. ^Gelder, Stuart R.; Gagnon, Nicole L.; Nelson, Kerri (2002). "Taxonomic Considerations and Distribution of the Branchiobdellida (Annelida: Clitellata) on the North American Continent".Northeastern Naturalist.9(4): 451–468.doi:10.1656/1092-6194(2002)009[0451:TCADOT]2.0.CO;2.JSTOR3858556.S2CID85774943.
  37. ^abcRohde, Klaus (2005).Marine Parasitology.CSIROPublishing. p. 185.ISBN978-0-643-09927-2.
  38. ^abcdeRuppert, Fox & Barnes 2004,pp. 475–477
  39. ^Govedich, Fredric R.; Bain, Bonnie A. (14 March 2005)."All about leeches"(PDF).Archived fromthe original(PDF)on 21 August 2010.Retrieved19 January2010.
  40. ^Sawyer, Roy T."Leech biology and behaviour"(PDF).biopharm-leeches.com.Archived fromthe original(PDF)on 10 September 2011.
  41. ^Dziekońska-Rynko, Janina; Bielecki, Aleksander; Palińska, Katarzyna (2009)."Activity of selected hydrolytic enzymes from leeches (Clitellata: Hirudinida) with different feeding strategies".Biologia.64(2): 370–376.Bibcode:2009Biolg..64..370D.doi:10.2478/s11756-009-0048-0.
  42. ^abRuppert, Fox & Barnes 2004,pp. 472–474
  43. ^abElder, H. Y. (1980). Elder, H. Y.; Trueman, E. R. (eds.).Peristaltic Mechanisms.CUP Archive. pp. 84–85.ISBN978-0-521-29795-0.{{cite book}}:|work=ignored (help)
  44. ^Sawyer, Roy (1981). Kenneth, Muller; Nicholls, John; Stent, Gunther (eds.).Neurobiology of the Leech.Cold Spring Harbor Laboratory.pp. 7–26.ISBN978-0-87969-146-2.
  45. ^Smith, Douglas Grant (2001).Pennak's Freshwater Invertebrates of the United States: Porifera to Crustacea.John Wiley & Sons.pp. 304–305.ISBN978-0-471-35837-4.
  46. ^Burke, Don (2005).The complete Burke's backyard: the ultimate book of fact sheets.Murdoch Books.ISBN978-1-74045-739-2.
  47. ^Fujimoto, Gary; Robin, Marc; Dessery, Bradford (2003).The Traveler's Medical Guide.Prairie Smoke Press.ISBN978-0-9704482-5-5.
  48. ^abVictorian Poisons Information Centre: LeechesVictorian Poisons Information Centre. Retrieved 28 July 2007
  49. ^Chow, C. K.; Wong, S. S.; Ho, A. C.; Lau, S. K. (2005)."Unilateral epistaxis after swimming in a stream".Hong Kong Medical Journal.11(2): 110–112.PMID15815064.See also:lay summaryfrom Reuters, 11 April 2005.
  50. ^Ahl-Khleif, A.; Roth, M.; Menge, C.; et al. (2011)."Tenacity of mammalian viruses in the gut of leeches fed with porcine blood".Journal of Medical Microbiology.60(6): 787–792.doi:10.1099/jmm.0.027250-0.PMID21372183.
  51. ^Nehili, Malika; Ilk, Christoph; Mehlhorn, Heinz; et al. (1994). "Experiments on the possible role of leeches as vectors of animal and human pathogens: a light and electron microscopy study".Parasitology Research.80(4): 277–290.doi:10.1007/bf02351867.ISSN0044-3255.PMID8073013.S2CID19770060.
  52. ^abMeir, Rigbi; Levy, Haim; Eldor, Amiram; et al. (1987). "The saliva of the medicinal leech Hirudo medicinalis—II. Inhibition of platelet aggregation and of leukocyte activity and examination of reputed anaesthetic effects".Comparative Biochemistry and Physiology C.88(1): 95–98.doi:10.1016/0742-8413(87)90052-1.PMID2890494.
  53. ^Siddall, Mark (7 July 2008)."Myth Busters: Leech Anaesthetic".BdellaNea.
  54. ^Singh (2011)."Efficacy of leech therapy in the management of osteoarthritis (Sandhivata)".Ayu.32(2): 213–217.doi:10.4103/0974-8520.92589.PMC3296343.PMID22408305.
  55. ^Laurent, V.; Salzet, B.; Verger-Bocquet, M.; Bernet, F.; Salzet, M. (2000)."Morphine-like substance in leech ganglia. Evidence and immune modulation".European Journal of Biochemistry.267(8): 2354–2361.doi:10.1046/j.1432-1327.2000.01239.x.PMID10759861.
  56. ^Siddall, Mark; Borda, Liz; Burreson, Gene; et al."Blood Lust II".Laboratory of Phylohirudinology,American Museum of Natural History.Archived fromthe originalon 7 June 2020.Retrieved15 December2013.
  57. ^Yi-Te Lai; Jiun-Hong Chen (2010).Đài loan điệt loại động vật chí: Leech Fauna of Taiwan-Biota Taiwanica(in Chinese). Quốc lập đài loan đại học xuất bản trung tâm. p. 89.ISBN978-986-02-2760-4.
  58. ^Payton, Brian (1981). Muller, Kenneth; Nicholls, John; Stent, Gunther (eds.).Neurobiology of the Leech.Cold Spring Harbor Laboratory.pp. 27–34.ISBN978-0-87969-146-2.
  59. ^Mory, Robert N.; Mindell, David; Bloom, David A. (2014). "The Leech and the Physician: Biology, Etymology, and Medical Practice withHirudinea medicinalis".World Journal of Surgery.24(7): 878–883.doi:10.1007/s002680010141.hdl:2027.42/42411.PMID10833259.S2CID18166996.
  60. ^abMarren, Peter;Mabey, Richard(2010).Bugs Britannica.Chatto & Windus.pp. 45–48.ISBN978-0-7011-8180-2.
  61. ^Pliny(1991).Natural History: A Selection.Translated by Healy, John F.Penguin Books.p. 283.ISBN978-0-14-044413-1.
  62. ^Mory, Robert N.; Mindell, David; Bloom, David A. (2014). "The Leech and the Physician: Biology, Etymology, and Medical Practice with Hirudinea medicinalis".World Journal of Surgery.24(7): 878–883.doi:10.1007/s002680010141.hdl:2027.42/42411.ISSN0364-2313.PMID10833259.S2CID18166996.
  63. ^Jourdier, August; Coste, M. (March 1859)."Hirudiculture (Leech-Culture) (fromLa Pisciculture et la Production des Sanguesues(Fish farming and leech production). Paris: Hachette et Cie ".The Journal of Agriculture.New Series. 8 (July 1857–March 1859 ).William Blackwood and Sons:641–648.
  64. ^anon (2016).Medicine: The Definitive Illustrated History.Dorling Kindersley. p. 35.ISBN978-0-241-28715-6.
  65. ^Cho, Joohee (4 March 2008)."Some Docs Latching Onto Leeches".ABC News.Retrieved27 April2018.
  66. ^Adams, Stephen L. (1988). "The Medicinal Leech: A Page from the Annelids of Internal Medicine".Annals of Internal Medicine.109(5): 399–405.doi:10.7326/0003-4819-109-5-399.PMID3044211.
  67. ^Teut, M.; Warning, A. (2008). "Leeches, phytotherapy and physiotherapy in osteo-arthrosis of the knee—a geriatric case study".Forsch Komplementärmed.15(5): 269–272.doi:10.1159/000158875.PMID19001824.S2CID196365336.
  68. ^Michalsen, A.; Moebus, S.; Spahn, G.; Esch, T.; Langhorst, J.; Dobos, G.J. (2002). "Leech therapy for symptomatic treatment of knee osteoarthritis: Results and implications of a pilot study".Alternative Therapies in Health and Medicine.8(5): 84–88.PMID12233807.
  69. ^abSig, A. K.; Guney, M.; Uskudar Guclu, A.; et al. (2017)."Medicinal leech therapy—an overall perspective".Integrative Medicine Research.6(4): 337–343.doi:10.1016/j.imr.2017.08.001.PMC5741396.PMID29296560.
  70. ^Abdualkader, A. M.; Ghawi, A. M.; Alaama, M.; Awang, M.; Merzouk, A. (2013)."Leech Therapeutic Applications".Indian Journal of Pharmacological Science.75(2 (March–April)): 127–137.PMC3757849.PMID24019559.
  71. ^Haycraft, John B. (1883). "IV. On the action of a secretion obtained from the medicinal leech on the coagulation of the blood".Proceedings of the Royal Society of London.36(228–231): 478–487.doi:10.1098/rspl.1883.0135.S2CID83910684.
  72. ^Fischer, Karl-Georg; Van de Loo, Andreas; Bohler, Joachim (1999)."Recombinant hirudin (lepirudin) as anticoagulant in intensive care patients treated with continuous hemodialysis".Kidney International.56(Suppl. 72): S46–S50.doi:10.1046/j.1523-1755.56.s72.2.x.PMID10560805.
  73. ^Sohn, J.; Kang, H.; Rao, K.; Kim, C.; Choi, E.; Chung, B.; Rhee, S. (2001). "Current status of the anticoagulant hirudin: its biotechnological production and clinical practice".Applied Microbiology and Biotechnology.57(5–6): 606–613.doi:10.1007/s00253-001-0856-9.PMID11778867.S2CID19304703.
  74. ^Schnell, Ida Bærholm; Thomsen, Philip Francis; Wilkinson, Nicholas; et al. (2012)."Screening mammal biodiversity using DNA from leeches".Current Biology.22(8): R262–R263.Bibcode:2012CBio...22.R262S.doi:10.1016/j.cub.2012.02.058.ISSN0960-9822.PMID22537625.
  75. ^Schnell, Ida Bærholm; Bohmann, Kristine; Schultze, Sebastian E.; et al. (2018)."Debugging diversity – a pan-continental exploration of the potential of terrestrial blood-feeding leeches as a vertebrate monitoring tool"(PDF).Molecular Ecology Resources.18(6): 1282–1298.doi:10.1111/1755-0998.12912.PMID29877042.S2CID46972335.
  76. ^Kidd, Karen A.; Graves, Stephanie D.; McKee, Graydon I.; et al. (2020)."Effects of Whole-Lake Additions of Ethynylestradiol on Leech Populations".Environmental Toxicology and Chemistry.39(8): 1608–1619.doi:10.1002/etc.4789.PMID32692460.S2CID220669536.

General bibliography

  • Ruppert, Edward E.; Fox, Richard S.; Barnes, Robert D. (2004).Invertebrate Zoology, 7th Edition.Cengage Learning.ISBN978-81-315-0104-7.

External links

  • Media related toHirudineaat Wikimedia Commons
  • Data related toHirudineaat Wikispecies
  • The dictionary definition ofleechat Wiktionary

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