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Equisetum

Equisetum (/kwsitm/; horsetail, snake grass, puzzlegrass) is the only


Equisetum
living genus in Equisetaceae, a family of vascular plants that reproduce by spores
rather than seeds.[2] Temporal range:
Callovian [1]-Holocene
Equisetum is a "living fossil" as it is the only living genus of the entire class
Pre OS D C P T J K Pg N
Equisetopsida, which for over one hundred million years was much more diverse
and dominated the understory of late Paleozoic forests. Some Equisetopsida were
large trees reaching to 30 meters tall.[3] The genus Calamites of the family
Calamitaceae, for example, is abundant in coal deposits from the Carboniferous
period.

A superficially similar but entirely unrelated flowering plant genus, mare's tail
(Hippuris), is occasionally referred to as "horsetail", and adding to confusion, the
name mare's tail is sometimes applied toEquisetum.[4]

The pattern of spacing of nodes in horsetails, wherein those toward the apex of
the shoot are increasingly close together, inspired John Napier to invent
logarithms.[5]

Contents
1 Etymology "Candocks" of the great horsetail
2 Description (Equisetum telmateia subsp.
2.1 Spores
telmateia), showing whorls of
2.2 Equisetum cell walls
branches and the tiny dark-tipped
3 Taxonomy
3.1 Species leaves
3.2 Named hybrids Scientific classification
4 Distribution, ecology and uses
Kingdom: Plantae
5 Medicinal uses
6 See also
Division: Pteridophyta
7 References Class: Equisetopsida
8 Further reading Order: Equisetales
9 External links
Family: Equisetaceae
Genus: Equisetum
Etymology L.

The name "horsetail", often used for the entire group, arose because the branched Type species
species somewhat resemble ahorse's tail. Similarly, the scientific name Equisetum Equisetum arvense
is derived from the Latin equus ("horse") + seta ("bristle"). L.

Other names include candock for branching individuals, and snake grass or Species
scouring-rush for unbranched or sparsely branched individuals. The latter name
See text
refers to the rush-like appearance of the plants, and to the fact that the stems are
coated with abrasive silicates, making them useful for scouring (cleaning) metal
items such as cooking pots or drinking mugs, particularly those made of tin. In German, the corresponding name is Zinnkraut ("tin-
herb"). Rough horsetail E. hyemale is still boiled and then dried in Japan, to be used for the final polishing process on woodcraft to
produce a smoother finish than any sandpaper. In Spanish-speaking countries, these plants are known as "cola de caballo," meaning
"horsetail".

Description
In these plants the leaves are greatly reduced and usually non-photosynthetic. They contain a single, non-branching vascular trace,
which is the defining feature of microphylls. However, it has recently been recognised that horsetail microphylls are probably not
ancestral as in Lycopodiophyta (clubmosses and relatives), but rather derived adaptations, evolved by reduction of megaphylls.[6]
They are, therefore, sometimes referred to as megaphylls to reflect thishomology.

The leaves of horsetails are arranged in whorls fused into nodal sheaths. The stems are usually green and photosynthetic, and are
distinctive in being hollow, jointed and ridged (with sometimes 3 but usually 6-40 ridges). There may or may not be whorls of
branches at the nodes.

Spores
The spores are borne under sporangiophores in strobili, cone-like structures at the tips of some of the
stems. In many species the cone-bearing shoots are unbranched, and in some (e.g. field horsetail, E.
arvense) they are non-photosynthetic, produced early in spring. In some other species (e.g. marsh
horsetail, E. palustre) they are very similar to sterile shoots, photosynthetic and with whorls of
branches.

Horsetails are mostly homosporous, though in the field horsetail smaller spores give rise to male
prothalli. The spores have four elaters that act as moisture-sensitive springs, assisting spore dispersal
through crawling and hopping motions after thesporangia have split open longitudinally.[7]

Equisetum cell walls


The crude cell extracts of all Equisetum species tested contain mixed-linkage glucan : Xyloglucan
endotransglucosylase (MXE) activity.[8] This is a novel enzyme and is not known to occur in any other
plants. In addition, the cell walls of all Equisetum species tested contain mixed-linkage glucan (MLG),
a polysaccharide which, until recently, was thought to be confined to the Poales.[9][10] The evolutionary
distance between Equisetum and the Poales suggests that each evolved MLG independently. The Vegetative stem:
presence of MXE activity in Equisetum suggests that they have evolved MLG along with some B = branch in whorl
mechanism of cell wall modification. The lack of MXE in the Poales suggests that there it must play I = internode
L = leaves
some other, currently unknown, role. Due to the correlation between MXE activity and cell age, MXE
N = node
has been proposed to promote the cessation of cell expansion.

Taxonomy

Species
The living members of the genus Equisetum are divided into two distinct lineages, which are usually treated as subgenera. The name
of the type subgenus, Equisetum, means "horse hair" inLatin, while the name of the other subgenus,Hippochaete, means "horse hair"
in Greek. Hybrids are common, but hybridization has only been recorded between members of the same subgenus.[11] While plants
of subgenus Equisetum are usually referred to as horsetails, those of subgenus Hippochaete are often called scouring rushes,
especially when unbranched.
Two Equisetum plants are sold commercially under the names Equisetum japonicum
(barred horsetail) and Equisetum camtschatcense (Kamchatka horsetail). These are
both types of E. hyemale var. hyemale, although they may also be listed as varieties
of E. hyemale.

Subgenus Equisetum

Equisetum arvense L. field horsetail, common horsetail or mare's tail;


circumboreal down through temperate zones
Equisetum bogotense Kunth Andean horsetail; upland South America
up to Costa Rica; includesE. rinihuense, sometimes treated as a
separate species
Equisetum diffusum L. Himalayan horsetail; Himalayan India and
China and adjacent nations above about 1500 feet (450 m)
Equisetum fluviatile L. water horsetail; circumboreal down through
temperate zones
Equisetum palustre L. marsh horsetail; circumboreal down through
temperate zones
Equisetum pratense Ehrh. meadow horsetail, shade horsetail, shady
horsetail; circumboreal except for tundra down through cool temperate
zones
Equisetum sylvaticum L. wood horsetail; circumboreal down through
cool temperate zones, more restricted in east Asia
Strobilus of Northern giant horsetail
Equisetum telmateia Ehrh. great horsetail, northern giant horsetail;
Europe to Asia Minor and north Africa, also west coast of North America (Equisetum telmateia subsp. braunii),
terminal on an unbranched stem.
Subgenus Hippochaete

Equisetum giganteum L. southern giant horsetail or giant horsetail;


temperate to tropical South America and Central America north to
southern Mexico
Equisetum hyemale L. rough horsetail, rough scouring rush; most of
non-tropical northern hemisphere
Equisetum laevigatumA.Braun smooth horsetail, smooth scouring rush;
western 3/4 of North America down into northwestern Mexico; also
sometimes known as Equisetum kansanum
Equisetum myriochaetumSchltdl. & Cham. Mexican giant horsetail; from
central Mexico south to Peru
Equisetum ramosissimumDesf. (including E. debile) branched
horsetail; Asia, Europe, Africa, southwest Pacific islands
Equisetum scirpoides Michx. dwarf horsetail, dwarf scouring rush;
northern (cool temperate) zones worldwide
Equisetum variegatumSchleich. ex Weber & Mohr variegated horsetail,
variegated scouring rush; northern (cool temperate) zones worldwide,
except for northeasternmost Asia Microscopic view of rough horsetail,
Equisetum hyemale (2-1-0-1-2 is one
unplaced to subgenus millimetre with 1/20th graduation).
The small white protuberances are
Equisetum dimorphum- Lower Jurassic, Argentina
accumulated silicates on cells.
Equisetum thermale - Middle to Late Jurassic, Argentina

Named hybrids

Hybrids between species in subgenus Equisetum

Equisetum bowmanii C.N.Page (Equisetum sylvaticum Equisetum telmateia)


Equisetum dycei C.N.Page (Equisetum fluviatile Equisetum palustre)
Equisetum font-queri Rothm. (Equisetum palustre Equisetum telmateia)
Equisetum litorale Khlew ex Rupr. (Equisetum arvense Equisetum fluviatile)
Equisetum mchaffieae C.N.Page (Equisetum fluviatile Equisetum
pratense)
Equisetum mildeanum Rothm. (Equisetum pratense Equisetum
sylvaticum)
Equisetum robertsii Dines (Equisetum arvense Equisetum telmateia)
Equisetum rothmaleri C.N.Page (Equisetum arvense Equisetum
palustre)
Equisetum willmotii C.N.Page (Equisetum fluviatile Equisetum
telmateia)

Hybrids between species in subgenus Hippochaete

Equisetum ferrissii Clute (Equisetum hyemale Equisetum laevigatum)


Equisetum moorei Newman (Equisetum hyemale Equisetum
ramosissimum)
Equisetum nelsonii (A.A.Eaton) Schaffn. (Equisetum laevigatum
Equisetum variegatum)
Equisetum schaffneri Milde (Equisetum giganteum Equisetum
myriochaetum) Rough horsetail in Parc floral de
Equisetum trachyodon (A.Braun) W.D.J.Koch (Equisetum hyemale Paris
Equisetum variegatum)

Distribution, ecology and uses


The genus Equisetum as a whole, while concentrated in the non-tropical northern
hemisphere, is near-cosmopolitan, being absent only from Antarctica, though they
are not known to be native to Australia, New Zealand nor the islands of the Pacific.
They are most common in northern North America (Canada and the northernmost
United States), where the genus is represented by nine species (arvense, fluviatile,
palustre, pratense, sylvaticum, hyemale, laevigatum, scirpoides, and variegatum).
Only four (bogotense, giganteum, myriochaetum, and ramosissimum) of the fifteen
Branched horsetail (E.
species are known to be native south of the Equator. They are perennial plants, ramosissimum)
herbaceous and dying back in winter as most temperate species, or evergreen as
most tropical species and the temperate species rough horsetail (E. hyemale),
branched horsetail (E. ramosissimum), dwarf horsetail (E. scirpoides) and variegated
horsetail (E. variegatum). They typically grow 0.2-1.5 m tall, though the "giant
horsetails" are recorded to grow as high as 2.5 m (northern giant horsetail, E.
telmateia), 5 m (southern giant horsetail, E. giganteum) or 8 m (Mexican giant
horsetail, E. myriochaetum), and allegedly even more.[12]

Many species in this genus prefer wet sandy soils, while others are adapted to wet
clay soils. One species, Equisetum fluviatile, is an emergent aquatic, rooted in water
with shoots growing into the air. The stalks arise from rhizomes that are deep
underground and difficult to dig out. The field horsetail (E. arvense) can be a
nuisance weed, readily regrowing from the rhizome after being pulled out. It is
unaffected by many herbicides designed to kill seed plants. However, as E. arvense
prefers an acid soil, lime may be used to assist in eradication efforts to bring the soil
pH to 7 or 8.[13] Members of the genus have been declared noxious weeds in
Australia and in the US state ofOregon.[14][15]
Equisetum moorei (Rough
All the Equisetum are classed as "unwanted organisms" in New Zealand and are Horsetail Branched Horsetail)
listed on the National Pest Plant Accord.
If eaten over a long enough period of time, some species of horsetail can be poisonous to grazing animals, including horses.[16] The
toxicity appears to be due to thiaminase enzymes, which can cause thiamine (vitamin B1) deficiency.[17][18][19][20] People have
regularly consumed horsetails. The young plants are eaten cooked or raw, but considerable care must be taken.[21] For example, the
fertile stems bearing strobili of some species are cooked and eaten like asparagus (a dish called tsukushi[22] ) in Japan.[23] The people
of ancient Rome would eat meadow horsetail in a similar manner, and they also used it to make tea as well as a thickening
powder.[24] Native Americans in the Pacific Northwest eat the young shoots of this plant raw.[25] The plants are used as a dye and
give a soft green colour. An extract is often used to provide silica for supplementation. Horsetail was often used by Indians to polish
wooden tools. Equisetum species are often used to analyze gold concentrations in an area due to their ability to take up the metal
when it is in a solution.[24]

Medicinal uses
Extracts and other preparations of E. arvense have served as herbal remedies, with records dating to ancient Greek and Roman
medical sources.[21] Reliable modern alternative medicine sources include cautions with regard to its use.[21] In 2009, the European
Food Safety Authority issued a report assessing some specific health claims for E. arvensee.g., for invigoration, weight control,
and skin, hair, and bone healthconcluding that none could be substantiated.[26] There is currently insufficient scientific evidence to
draw conclusions regarding its effectiveness as a medicine for any human conditions described.[21] Even so, E. giganteum
preparations are widely used in South America as an orally administered diuretic to reduce swelling caused by excess fluid retention
and for urinary infections, bladder and kidney disorders. Horsetail preparations contain silicon, so they are sometimes suggested as a
treatment for osteoporosis (brittle bone disorders).[21][27]

See also
List of plants poisonous to equines

References
1. "Equisetum thermale sp. nov. (Equisetales) from the Jurassic SanAgustn hot spring deposit, Patagonia: anatomy ,
paleoecology, and inferred paleoecophysiology" (http://www.amjbot.org/cgi/content/abstract/98/4/680). American
Journal of Botany. 98 (4): 68097. April 2011.doi:10.3732/ajb.1000211 (https://doi.org/10.3732%2Fajb.1000211).
PMID 21613167 (https://www.ncbi.nlm.nih.gov/pubmed/21613167).
2. Sunset Western Garden Book, 1995:606607
3. "An Introduction to the GenusEquisetum and the Class Sphenopsida as a whole"(https://web.archive.org/web/2009
0714050011/http://www.fiu.edu/~chusb001/GiantEquisetum/Intro_Equisetum.html). Florida International University.
Archived from the original (http://www.fiu.edu/~chusb001/GiantEquisetum/Intro_Equisetum.html)on 2009-07-14.
Retrieved 2009-07-22.
4. Oxford English Dictionary.
5. Sacks, Oliver (August 2011). "Field Trip: Hunting Horsetails". The New Yorker.
6. Rutishauser, R (November 1999). "Polymerous leaf whorls in vascular plants: Developmental morphology and
fuzziness of organ identities".International Journal of Plant Sciences. 160 (S6): S81S103. doi:10.1086/314221 (http
s://doi.org/10.1086%2F314221). PMID 10572024 (https://www.ncbi.nlm.nih.gov/pubmed/10572024).
7. "Horsetail plant spores use 'legs' to walk and jump - BBC News"(http://www.bbc.com/news/science-environment-24
025365). BBC News. Retrieved 2015-11-30.
8. Fry, S. C.; Mohler, K. E.; Nesselrode, B. H. W. A.; Frankov, L. (2008). "Mixed-linkage -glucan:xyloglucan
endotransglucosylase, a novel wall-remodelling enzyme fromEquisetum (horsetails) and charophytic algae".The
Plant Journal. 55 (2): 240252. doi:10.1111/j.1365-313X.2008.03504.x(https://doi.org/10.1111%2Fj.1365-313X.200
8.03504.x). PMID 18397375 (https://www.ncbi.nlm.nih.gov/pubmed/18397375).
9. Fry, Stephen C.; Nesselrode, Bertram H. W. A.; Miller, Janice G.; Mewburn, Ben R. (2008). "Mixed-linkage
(13,14)--d-glucan is a major hemicellulose ofEquisetum (horsetail) cell walls". New Phytologist. 179 (1): 104
15. doi:10.1111/j.1469-8137.2008.02435.x(https://doi.org/10.1111%2Fj.1469-8137.2008.02435.x) . PMID 18393951
(https://www.ncbi.nlm.nih.gov/pubmed/18393951).
10. Srensen, Iben; Pettolino, Filomena A.; Wilson, Sarah M.; Doblin, Monika S.; Johansen, Bo; Bacic, Antony; Willats,
William G. T. (2008). "Mixed-linkage (13),(14)--d-glucan is not unique to the Poales and is an abundant
component of Equisetum arvense cell walls". The Plant Journal. 54 (3): 51021. doi:10.1111/j.1365-
313X.2008.03453.x (https://doi.org/10.1111%2Fj.1365-313X.2008.03453.x) . PMID 18284587 (https://www.ncbi.nlm.
nih.gov/pubmed/18284587).
11. Pigott, Anthony (4 October 2001)."Summary of Equisetum Taxonomy" (https://web.archive.org/web/2012102120193
9/http://www.btinternet.com/~pigott/equisetum/taxonsum.html). National Collection of Equisetum. Archived fromthe
original (http://www.btinternet.com/~pigott/equisetum/taxonsum.html) on 21 October 2012. Retrieved 17 June 2013.
12. Husby, Chad E. (2003): How large are the giant horsetails(http://www.fiu.edu/~chusb001/giant_equisetum.html)?
Version of 2003-03-19. Retrieved 2008-11-20. Archived (https://web.archive.org/web/20040404033635/http://www .fi
u.edu/~chusb001/giant_equisetum.html)April 4, 2004, at the Wayback Machine.
13. Kress, Henriette, Getting rid of horsetail (http://www.henriettesherbal.com/blog/getting-rid-horsetail.html), Henriette's
Herbal Homepage, April 7th, 2005. Retrieved May 19, 2010.
14. William Thomas Parsons; Eric George Cuthbertson (2001).Noxious weeds of Australia(https://books.google.com/bo
oks?id=sRCrNAQQrpwC&pg=PA14&dq=Equisetum+australia&q=Equisetum%20australia). CSIRO Publishing. p. 14.
ISBN 978-0-643-06514-7.
15. "Equisetum telmateia Ehrh. giant horsetail" (http://plants.usda.gov/java/profile?symbol=EQTE)
. USDA. Retrieved
2010-05-18.
16. Israelsen, Clark E.; McKendrick, Scott S. & Bagley
, Clell V. (2006): Poisonous Plants and Equine.PDF fulltext (http://
www.msuextension.org/ruralliving/Dream/PDF/Equine_poison.pdf) Archived (https://web.archive.org/web/201101120
23355/http://www.msuextension.org/ruralliving/Dream/PDF/Equine_poison.pdf)January 12, 2011, at theWayback
Machine.
17. Henderson JA, Evans EV, McIntosh RA (June 1952). "The antithiamine action ofEquisetum". Journal of the
American Veterinary Medical Association. 120 (903): 3758. PMID 14927511 (https://www.ncbi.nlm.nih.gov/pubmed/
14927511).
18. Fabre, B; Geay, B.; Beaufils, P. (1993). "Thiaminase activity in Equisetum arvense and its extracts". Plant Med
Phytother. 26: 1907.
19. "Horsetail" (http://www.nlm.nih.gov/medlineplus/druginfo/natural/843.html). National Library of Medicine. Retrieved
3 February 2013.
20. Pohl, Richard (1955). "Toxicity of ferns and Equisetum". American Fern Journal. 45 (3): 9597. doi:10.2307/1544850
(https://doi.org/10.2307%2F1544850).
21. Steven D. Ehrlich (2011) "Horsetail", University of Maryland Medical Center
Complementary and Alternative
Medicine Guide, 5 March 2011 review date, see[1] (http://umm.edu/health/medical/altmed/herb/horsetail)
, accessed
31 July 2014.
22. Michael Ashkenazi, Jeanne Jacob. 2003. Food culture in Japan. Greenwood Publishing Group. 232 p.
23. Plants For A Future Database.(http://www.pfaf.org/user/DatabaseSearhResult.aspx)
24. Plants of the Pacific Northwest Coast: Washington, Oregon, British Columbia & Alaska, W
ritten by Paul Alaback,
ISBN 978-1-55105-530-5
25. Erna Gunther. 1973. Ethnobotany of western Washington: The knowledge and use of indigenous plants by Native
Americans.
26. "Scientific opinion on the substantiation of health claims related toEquisetum arvense L. and invigoration of the body
(ID 2437), maintenance of skin (ID 2438), maintenance of hair (ID 2438), maintenance of bone (ID 2439), and
maintenance or achievement of a normal body weight (ID 2783) pursuant to Article 13 of Regulation (EC) No
1924/2006" (http://www.efsa.europa.eu/en/scdocs/scdoc/1289.htm). European Food Safety Authority. Retrieved
2013-10-09.
27. "Horsetail" (https://www.nlm.nih.gov/medlineplus/druginfo/natural/843.html). US National Institute of Health.
Retrieved 14 November 2013.

Further reading
Walkowiak, Radoslaw (2008):IEA Equisetum Taxonomy. Version of 2008-10-04. Retrieved 2011-11-07.
Pryer, K.M.; Schuettpelz, E.; Wolf, P.G.; Schneider, H.; Smith, A.R. & Cranfill, R. (2004): Phylogeny and evolution of
ferns (monilophytes) with a focus on the early leptosporangiate divergences. Am. J. Bot. 91(10): 1582-1598. PDF
fulltext
Rutishauser, R. (1999). "Polymerous leaf whorls in vascular plants: developmental morphology and fuzziness of
organ identities". International Journal of Plant Sciences. 160 (S6): S81S103. doi:10.1086/314221.
PMID 10572024.
Weber, Reinhard (2005): Equisetites aequecaliginosussp. nov., ein Riesenschachtelhalm aus der spttriassischen
Formation Santa Clara, Sonora, Mexiko Equisetites
[ aequecaliginosussp. nov., a tall horsetail from the Late Triassic
Santa Clara Formation, Sonora, Mexico].Revue de Palobiologie24(1): 331-364 [German with English abstract].
PDf fulltext

External links
Equisetum at the Tree of Life Web Project
National Collection of Equisetum
The Wonderful World of Equisetum
International Equisetological Association
"Horsetail". Encyclopdia Britannica(11th ed.). 1911.

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