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The European medical leech, ''[[Hirudo medicinalis]]'', and some [[congener]]s, as well as some other [[species]], have been used for clinical [[bloodletting]] for thousands of years, although most leeches do not feed on human blood, but instead prey on small [[invertebrate]]s, which they eat whole. Haemophagic leeches attach to their hosts and remain there until they become full, at which point they fall off to digest. To feed on its host, leeches use their anterior suckers to connect to a host for feeding, and also release an anesthetic to prevent the host from feeling the leech. Once attached, leeches use a combination of mucus and suction to stay attached and secrete an anti-clotting enzyme, [[hirudin]], into the host's [[Blood|blood stream]].
The European medical leech, ''[[Hirudo medicinalis]]'', and some [[congener]]s, as well as some other [[species]], have been used for clinical [[bloodletting]] for thousands of years, although most leeches do not feed on human blood, but instead prey on small [[invertebrate]]s, which they eat whole. Haemophagic leeches attach to their hosts and remain there until they become full, at which point they fall off to digest. To feed on its host, leeches use their anterior suckers to connect to a host for feeding, and also release an anesthetic to prevent the host from feeling the leech. Once attached, leeches use a combination of mucus and suction to stay attached and secrete an anti-clotting enzyme, [[hirudin]], into the host's [[Blood|blood stream]].



==Systematics and taxonomy==
==Systematics and taxonomy==
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There is some dispute as to whether Hirudinea should be a class itself, or a subclass of the [[Clitellata]]. The resolution mainly depends on the eventual fate of the oligochaetes, which as noted above do not form a natural group as traditionally circumscribed. Another possibility would be to include the leeches in the [[taxon]] Oligochaeta, which would then be ranked as a class and contain most of the clitellates. The [[Branchiobdellida]] are leechlike clitellates which were formerly included in the Hirudinea but are apparently just really close relatives.
There is some dispute as to whether Hirudinea should be a class itself, or a subclass of the [[Clitellata]]. The resolution mainly depends on the eventual fate of the oligochaetes, which as noted above do not form a natural group as traditionally circumscribed. Another possibility would be to include the leeches in the [[taxon]] Oligochaeta, which would then be ranked as a class and contain most of the clitellates. The [[Branchiobdellida]] are leechlike clitellates which were formerly included in the Hirudinea but are apparently just really close relatives.
[[File:Americobdella.jpg|thumb|right|This giant ''[[Americobdella]]'' leech from southern Chile is an ancient arhynchobdellid. It is a [[predator]], feeding on [[earthworm]]s which it swallows whole.]]
[[File:Americobdella.jpg|thumb|right|This giant ''[[Americobdella]]'' leech from southern Chile is an ancient arhynchobdellid. It is a [[predator]], feeding on [[earthworm]]s which it swallows whole.]]
[[File:HaemopisSanguisuga.jpg|thumb|The [[Horse leech]], ''Haemopis sanguisuga'', does not feed on blood, but prey on small invertebrates and [[tadpole]]s]]
[[File:20100214 Leech climbing door at Lake Leake, Tasmania.ogg|thumb|right|Leech climbing a door by [[Lake Leake, Tasmania]]]]
[[File:20100214 Leech climbing door at Lake Leake, Tasmania.ogg|thumb|right|Leech climbing a door by [[Lake Leake, Tasmania]]]]



Revision as of 11:34, 30 July 2011

Leech
Hirudo medicinalis
Scientific classification
Kingdom:
Phylum:
Class:
Subclass:
Lamarck, 1818
Infraclasses

Acanthobdellidea
Euhirudinea
(but see below)

Haemadipsa zeylanica, a terrestrial leech found in the mountains of Japan.

Leeches are annelids comprising the subclass Hirudinea. There are freshwater, terrestrial[1], and marine leeches. Like the Oligochaeta, they share the presence of a clitellum. Like earthworms, leeches are hermaphrodites. Some, but not all, leeches are hematophagous. Some species of leech will nurture their young, while providing food, transport, and protection, which is unusual behavior amongst annelids.

Unlike other annelids, there is no correspondance between the external segmentation of a leech's body surface with the segmentation of its internal organs.[2] The body surface of the animal can be divided into 102 annuli whereas its internal structures are divided into 32 segments.[3] Of the 32 segments within the body, the first four anterior segments are designated as head segments, which include an anterior brain and sucker. This is followed by 21 midbody segments. The midbody segments include 21 neuronal ganglia, two reproductive organs, and 9 pairs of testes. Finally, the last 7 segments are fused to form the animal's tail sucker as well as its posterior brain.

The European medical leech, Hirudo medicinalis, and some congeners, as well as some other species, have been used for clinical bloodletting for thousands of years, although most leeches do not feed on human blood, but instead prey on small invertebrates, which they eat whole. Haemophagic leeches attach to their hosts and remain there until they become full, at which point they fall off to digest. To feed on its host, leeches use their anterior suckers to connect to a host for feeding, and also release an anesthetic to prevent the host from feeling the leech. Once attached, leeches use a combination of mucus and suction to stay attached and secrete an anti-clotting enzyme, hirudin, into the host's blood stream.

Systematics and taxonomy

Leeches are presumed to have evolved from certain Oligochaeta, most of which feed on detritus. However, some species in the Lumbriculidae are predatory and have similar adaptations as found in leeches. Consequently, the systematics and taxonomy of leeches is in need of review. While leeches form a clade, the remaining oligochetes are not their sister taxon but a diverse paraphyletic group containing some lineages that are closely related to leeches, and others that are far more distant.

There is some dispute as to whether Hirudinea should be a class itself, or a subclass of the Clitellata. The resolution mainly depends on the eventual fate of the oligochaetes, which as noted above do not form a natural group as traditionally circumscribed. Another possibility would be to include the leeches in the taxon Oligochaeta, which would then be ranked as a class and contain most of the clitellates. The Branchiobdellida are leechlike clitellates which were formerly included in the Hirudinea but are apparently just really close relatives.

This giant Americobdella leech from southern Chile is an ancient arhynchobdellid. It is a predator, feeding on earthworms which it swallows whole.
The Horse leech, Haemopis sanguisuga, does not feed on blood, but prey on small invertebrates and tadpoles
Leech climbing a door by Lake Leake, Tasmania

The more primitive Acanthobdellidea are often included with the leeches, but some authors treat them as a separate clitellate group. True leeches of the infraclass Euhirudinea have both anterior and posterior suckers. They are divided into two groups: Arhynchobdellida and Rhynchobdellida

  • Rhynchobdellida): "Jawless" leeches, armed with a muscular straw-like proboscis puncturing organ in a retractable sheath. The Rhynchobdellae consist of two families:
    • Glossiphoniidae: Flattened leeches with a poorly defined anterior sucker
    • Piscicolida: have cylindrical bodies and a usually well-marked, bell-shaped, anterior sucker. The Glossiphoniidae live in fresh-water habitats; the Pisciolidae are found in seawater habitats.
  • Arhynchobdellida: Leeches which lack a proboscis and which may or may not have jaws armed with teeth. Arhynchobellids are divided into two orders:
    • Gnathobdela: In this order of "jawed" leeches, armed with teeth, is found the quintessential leech: the European medical (bloodsucking) leech, Hirudo medicinalis. It has a tripartite-jaw filled with hundreds of tiny sharp teeth. The incision mark left on the skin by the European medical leech is an inverted Y inside a circle. Its North American counterpart is Macrobdela decora, a much less efficient medical leech.[4] Within this order, the family Hirudidae is characterized by aquatic leeches and the family Haemadipsidae by terrestrial leeches. In the latter are Haemadipsa sylvestris, the Indian leech and Haemadipsa zeylanica (yamabiru), the Japanese mountain or land leech.
    • Pharyngobdella: These so called worm-leeches consist of freshwater or amphibious leeches that have lost the ability to penetrate a host's tissue and suck blood. They are carnivorous and equipped with a relatively large, toothless, mouth to ingest worms or insect larvae, which are swallowed whole.
      The Pharyngobdella have six to eight pairs of eyes, as compared with five pairs in Gnathobdelliform leeches, and include three related families. The Erpobdellidae are some species from freshwater habitats.

Reproduction

Leeches are hermaphrodites, meaning each one of them has both female and male reproductive organs (ovaries and testes respectively). Leeches reproduce by reciprocal fertilization, and sperm transfer occurs during copulation. Similarly to the earthworms, leeches also use a clitellum to hold their eggs and secrete the cocoon.

During reproduction leeches utilize hyperdermic injection of their sperm. They use a spermatophore, which is a structure containing the sperm. Once next to another leech, the two will line up with their anterior side opposite the other's posterior. The leech then shoots the spermatophore into the clitellur region of the opposing leech where its sperm will make its way to the female reproductive parts.

Nutrition

Mouthparts and sucker
A leech swollen with blood
Leech attacking a slug

For leeches, the digestive system starts with the jaw which is located ventrally on the anterior side of the body. It is attached to the pharynx, then the esophagus extending to the crop, then to the gizzard, which leads to the intestinum, where it ends at the posterior sucker. The crop is a type of stomach that works like an expandable storage compartment. The crop allows a leech to store blood up to five times its body size; and because the leech produces an anti-coagulant, the stored blood remains in a liquid state; because of this ability to hold blood without the blood decaying, due to bacteria living inside the crop, medicinal leeches only need to feed two times a year.

The body of predatory leeches are similar, though some may also have a protrusible proboscis which is retracted in their mouth. Such leeches are often ambush predators, which lie in wait, and strike their prey using their proboscis in a spear-like fashion.[5]

It was long thought that bacteria in the gut carried on digestion for the leech instead of endogenous enzymes which are very low or absent in the intestine. Relatively recently it has been discovered that all leeches and leech species studied do produce endogenous intestinal exopeptidases,[6] which can unlink free terminal-end amino acids, one amino acid monomer at a time, from a gradually unwinding and degrading protein polymer. However, unzipping of the protein can start from either the amino (tail) or carboxyl (head) terminal-end of the protein molecule. It just so happens that the leech exopeptidase (arylamidases), possibly aided by proteases from endosymbiotic bacteria in the intestine, starts from the tail or amino protein, free-end, slowly but progressively removing many hundreds of individual terminal amino acids for resynthesis into proteins that constitute the leech. Since leeches lack endopeptidases, the mechanism of protein digestion can not follow the same sequence as it would in all other animals where exopeptidases act sequentially on peptides produced by the action of endopeptidases.[6] Exopeptidases are especially prominent in the common North American worm-leech Erpobdella punctata. This evolutionary choice of exopeptic digestion in Hirudinea distinguishes these carnivorous clitellates from Oligochaeta.

Deficiency of digestive enzymes (except exopeptidases) but more importantly deficiency of vitamins, B complex for example, in leeches is compensated for by enzymes and vitamins produced by endosymbiotic microflora. In Hirudo medicinalis these supplementary factors are produced by an obligatory symbiotic relationship with two bacterial species, Aeromonas veronii and a still uncharacterized Rikenella species. Non-bloodsucking leeches such as Erpobdella punctata are host to three bacterial symbionts, Pseudomonas sp., Aeromonas sp., and Klebsiella sp. (a slime producer). The bacteria are passed from parent to offspring in the cocoon as it is formed.

Leech bites

Effects

Though certain species of leeches feed on blood, not all species can bite; 90% of them solely feed off decomposing bodies and open wounds of amphibians, reptiles, waterfowl, fish, and mammals (including, but not limited to, humans). A leech attaches itself when it bites, and it will stay attached until it has had its fill of blood. Due to an anticoagulant (hirudin) that leeches secrete, bites may bleed more than a normal wound after the leech is removed. The effect of the anticoagulant will wear off several hours after the leech is removed and the wound is cleaned.

Leeches normally carry parasites in their digestive tract which cannot survive in humans and do not pose a threat. However, bacteria, viruses, and parasites from previous blood sources can survive within a leech for months, and may be retransmitted to humans.[citation needed] A study found both HIV and hepatitis B in African leeches from Cameroon.[7]

Removal

Hand removing a land leech—since they do not burrow into the skin nor leave the head in the wound.[8] A sore develops and lasts for about a week.[9] Grande Ronde River, Oregon (U.S.)

One recommended method of removal is using a fingernail or other flat, blunt object to break the seal of the oral sucker at the anterior end (the smaller, thinner end) of the leech, repeating with the posterior end, then flicking the leech away. As the fingernail is pushed along the person's skin against the leech, the suction of the sucker's seal is broken, at which point the leech will detach its jaws.[10][11]

Common but medically inadvisable techniques to remove a leech are to apply a flame, a lit cigarette, salt, soap, or a chemical such as alcohol, vinegar, lemon juice, insect repellent, heat rub, or certain carbonated drinks. These cause the leech to regurgitate its stomach contents into the wound and quickly detach. However, the vomit may carry disease, and thus increase the risk of infection.[10][11][12]

An externally attached leech will detach and fall off on its own when it is satiated on blood, which may be anywhere from 20 min. to 2 hours or more. Usually an hour is needed. After finishing feeding, the leech will detach and move away or just roll off that portion of the body.[12] Internal attachments, such as nasal passage or vaginal attachments, are more likely to require medical intervention.[13][14]

Treatment

After removal or detachment, the wound should be cleaned with soap and water, and bandaged. Bleeding may continue for some time, due to the leech's anti-clotting enzyme. Bleeding time will vary with location (some areas of the back bleed less than areas on the calves or thighs for example). Bleeding time may vary from just a few hours to perhaps as much as three days. This is a function of not only the hirudin but of other compounds(unspecified) that reduce the surface tension of the blood. Another factor would be if the patient were on anti-clotting medications at the time of leech therapy. Applying pressure can reduce bleeding, although blood loss from a single bite is not dangerous. The wound normally itches as it heals, but should not be scratched as this may complicate healing and introduce other infections. An antihistamine can reduce itching, and applying a cold pack can reduce pain or swelling.

Some people suffer severe allergic or anaphylactic reactions from leech bites, and require urgent medical care. Symptoms include red blotches or an itchy rash over the body, swelling away from the bitten area (especially around the lips or eyes), feeling faint or dizzy, and difficulty breathing.[12]

Prevention

There is no guaranteed method of preventing leech bites in leech-infested areas. The most reliable method is to cover exposed skin. The effect of insect repellents is disputed, but it is generally accepted that strong (maximum strength or tropical) insect repellents do help prevent bites.[citation needed]

Leech socks can be helpful in preventing bites when the full body will not be at risk of contact with leeches. Leech socks are pulled over the wearer's trousers to prevent leeches reaching the exposed skin of the legs and attaching there or climbing towards the torso. The socks are generally a light color that also makes it easier to spot leeches climbing up from the feet and looking for skin to attach to.[citation needed]

There are many home remedies to help prevent leech bites. Many people have a great deal of faith in these methods, but none of them have been proven effective. Some home remedies include: a dried residue of bath soap, tobacco leaves between the toes, pastes of salt or baking soda, citrus juice, Neem oil and eucalyptus oil. Diluted calcium hydroxide may also be used as a repellent, but may be damaging or irritating to the skin. One other remedy commonly practiced in the western ghats of southern India is castor oil mixed with snuff powder, or powdered tobacco.[citation needed]

Hirudotherapy

The term refers to the use of leeches in medicine.

The use of leeches in medicine dates as far back as 2,500 years ago when they were used for bloodletting in ancient India. Leech therapy is explained in ancient Ayurvedic texts. Many ancient civilizations practiced bloodletting including Indian and Greek civilizations. In ancient Greek history, bloodletting was practiced according to the humoral theory, which proposed that when the four humors, blood, phlegm, black and yellow bile in the human body were in balance, good health was guaranteed. An imbalance in the proportions of these humors was believed to be the cause of ill health. Records of this theory were found in the Greek philosopher Hippocrates' collection in the fifth century B.C. Bloodletting using leeches was one method used by physicians to balance the humors and to rid the body of the plethora.

The use of leeches in modern medicine made its comeback in the 1980s after years of decline, with the advent of microsurgery such as plastic and reconstructive surgeries. In operations such as these, one problem that arises is venous congestion due to inefficient venous drainage. Sometimes because of the technical difficulties in forming an anastomosis of a vein, no attempt is made to re-attach a venous supply to a flap at all. This condition is known as venous insufficiency. If this congestion is not cleared up quickly, the blood will clot, arteries that bring the tissues their necessary nourishment will become plugged and the tissues will die. To prevent this leeches are applied to a congested flap and a certain amount of excess blood is consumed before the leech falls away. The wound will also continue to bleed for a while due to the anticoagulant (hirudin) in the leeches' saliva. The combined effect is to reduce the swelling in the tissues and promoting healing by allowing fresh, oxygenated blood to reach the area.[15]

The active anticoagulant principle of leech saliva, is a small protein, hirudin. Discovery and isolation of this protein led to a method of producing it by recombinant technology. Recombinant hirudin is available to physicians as an intravenous anticoagulant preparation for injection, particularly useful for patients who are allergic to or cannot tolerate heparin.

Embryonic development

The first cleavage during early embryonic development in leech occurs at stage 2. This cleavage gives rise to an AB and a CD blastomere and is in the interphase of this cell division when a yolk-free cytoplasm called teloplasm is formed.[16] The teloplasm is known to be a determinant for the specification of the D cell fate.[17] In stage 3, during the second cleavage, an unequal division occurs in the CD blastomere. As a consequence, it creates a big D cell on the left and a smaller C cell to the right. This unequal division process is dependent on actinomycin,[18] and by the end of stage 3 the AB cell divides. On stage 4 of development, the micromeres and teloblast stem cells are formed and subsequently, the D quadrant divides to form the DM and the DNOPQ teloblast precursor cells. By the end stage 6, the zygote contains a set of 25 micromeres, 3 macromeres (A, B and C) and 10 teloblasts derived from the D quadrant.[19]

The teloblast are pairs of five different types (M, N, O, P and Q) of embryonic stem cells that form segmented columns of cells (germinal-band) in the surface of the embryo.[20] The M-derived cells make mesoderm and some small set of neurons, N results in neural tissues and some ventral ectoderm, Q contributes to the dorsal ectoderm and O and P in the leech are equipotent cells (same developmental potential) that produce lateral ectoderm; however the difference between the two of them is that P creates bigger batches of dorso-lateral epidermis than O.[17] The sludgeworm Tubifex, unlike the leech, specifies the O and P lineages early in development and therefore, these two cells are not equipotent.[21] Each segment of the body of the leech is generated from one M, O, P cell types and two N and two Q cells types.[17]

The ectoderm and mesoderm of the body trunk is exclusively derived from the teloblast cells in a region called posterior progress zone.[22][23] The head of the leech that come from a non-segmented region, is formed by the first set of micromeres derived from A, B, C and D cells, keeping the bilateral symmetry between the AD and BC cells.[23]

References

  1. ^ Fogden, S.; Proctor, J. (1985). "Notes on the Feeding of Land Leeches (Haemadipsa zeylanica Moore and H. picta Moore) in Gunung Mulu National Park, Sarawak". Biotropica. 17 (2): 172–174.
  2. ^ Buchsbaum, Ralph; Buchsbaum, Mildred; Pearse, John; Pearse, Vicki (1987). Animals Without Backbone (3rd ed.). Chicago: The University of Chicago Press. pp. 312–317. ISBN 0-226-07874-4.
  3. ^ Payton, Brian (1981). Kenneth, Muller; Nicholls, John; Stent, Gunther (eds.). Neurobiology of the Leech. New York: Cold Spring Harbor Laboratory. pp. 35–50. ISBN 0-87969-146-8.
  4. ^ http://www.fcps.edu/islandcreekes/ecology/freshwater_leech.htm
  5. ^ Govedich, Fredric R.; Bain, Bonnie A. (March 14, 2005). "All about leeches" (PDF). Retrieved January 19, 2010.
  6. ^ a b Sawyer, Roy T. "Leech biology and behaviour" (PDF).
  7. ^ Nehili M, Ilk C, Mehlhorn H, Ruhnau K, Dick W, Njayou M (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): 277–90. doi:10.1007/BF02351867. PMID 8073013.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  8. ^ Burke, Don (2005). The complete Burke's backyard: the ultimate book of fact sheets. Murdoch Books. ISBN 1740457390. Retrieved 2009-09-11.
  9. ^ Fujimoto, Gary (2003). The Traveler's Medical Guide. Prairie Smoke Press. ISBN 0970448252. Retrieved 2009-09-11. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  10. ^ a b "The Knowledge: Removing a leech". Times Online. October 15, 2006. Retrieved July 28, 2007.
  11. ^ a b Scenario Archive, Travel Survival: How to Remove a Leech Worst Case Scenarios. Retrieved on 2007-07-28.
  12. ^ a b c Victorian Poisons Information Centre: Leeches Victorian Poisons Information Centre. Retrieved on 2007-07-28.
  13. ^ Ibrahim, Adibah; Gharib, Hakim Bilal; Bidin, Mohd. Nizar (2003). "An Unusual Cause Of Vaginal Bleeding: A Case Report". The Internet Journal of Gynecology and Obstetrics. 2 (2). ISSN 1528-8439.
  14. ^ Blood-sucker gets up woman's nose Reuters via ABC News. 2005-04-11. Retrieved on 2007-07-28.
  15. ^ Calling Doctors Leech and Maggot to the O.R. Islamonline.net
  16. ^ J. Fernandez, N. Olea, V. Tellez & C. Matte (1990). "Structure and development of the egg of the glossiphoniid leech Theromyzon rude: reorganization of the fertilized egg during completion of the first meiotic division". Developmental Biology. 137 (1): 142–154. doi:10.1016/0012-1606(90)90015-B. PMID 2295361.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  17. ^ a b c D. A. Weisblat & M. Shankland (1985). "Cell lineage and segmentation in the leech". Philosophical Transactions of the Royal Society B: Biological Sciences. 312 (1153): 39–56. doi:10.1098/rstb.1985.0176. PMID 2869529.
  18. ^ D. C. Lyons & D. A. Weisblat (2009). "D quadrant specification in the leech Helobdella: actomyosin contractility controls the unequal cleavage of the CD blastomere". Developmental Biology. 334 (1): 46–58. doi:10.1016/j.ydbio.2009.07.007. PMID 19607823.
  19. ^ M. Sandig & W. Dohle (1988). "The cleavage pattern in the leech Theromyzon tessulatum (Hirudinea, Glossiphoniidae)". Journal of Morphology. 196 (2): 217–252. doi:10.1002/jmor.1051960210. PMID 3385778.
  20. ^ V. K. Berezovskii & M. Shankland (1996). "Segmental diversification of an identified leech neuron correlates with the segmental domain in which it expresses Lox2, a member of the Hox gene family". Journal of Neurobiology. 29 (3): 319–329. doi:10.1002/(SICI)1097-4695(199603)29:3<319::AID-NEU4>3.0.CO;2-C. PMID 8907161.
  21. ^ A. Arai, A. Nakamoto & T. Shimizu (2001). "Specification of ectodermal teloblast lineages in embryos of the oligochaete annelid Tubifex: involvement of novel cell-cell interactions". Development. 128 (7): 1211–1219. PMID 11245587.
  22. ^ D. Nardelli-Haefliger & M. Shankland (1993). "Lox10, a member of the NK-2 homeobox gene class, is expressed in a segmental pattern in the endoderm and in the cephalic nervous system of the leech Helobdella". Development. 118 (3): 877–892. PMID 7915671.
  23. ^ a b M. Shankland & A. E. Bruce (1998). "Axial patterning in the leech: developmental mechanisms and evolutionary implications" (PDF). Biological Bulletin. 195 (3): 370–372. doi:10.2307/1543150. JSTOR 1543150. PMID 9924777.