Lamprey

Lampreys (sometimes inaccurately called lamprey eels) are an ancient, extant

lineage of jawless fish of the order Petromyzontiformes, placed in the superclass
Lamprey
Cyclostomata. The adult lamprey may be characterized by a toothed, funnel-like Temporal range:
Late Devonian–Holocene [1]
sucking mouth. The common name "lamprey" is probably derived from Latin
lampetra, which may mean "stone licker" (lambere "to lick" + petra "stone"), PreЄ Є O S D C P T J K PgN
though the etymology is uncertain.[3]

There are about 38 known extant species of lampreys and five known extinct
species.[4] "Parasitic" carnivorous species are the most well-known, and feed by
boring into the flesh of other fish to suck their blood;[5] but only 18 species of
lampreys engage in this lifestyle (more correctly known as carnivorous "micro-
predation"[6]).[7] Of the 18 carnivorous species, nine migrate from saltwater to
freshwater to breed (some of them also have freshwater populations), and nine
live exclusively in freshwater. All non-carnivorous forms are freshwater
A European river lamprey (Lampetra
species.[8] Adults of the non-carnivorous species do not feed; they live off
reserves acquired as ammocoetes (larvae), which they obtain through filter fluviatilis)
feeding. Scientific classification
Kingdom: Animalia
Phylum: Chordata
Contents
Class: Hyperoartia
Distribution
Biology
Order: Petromyzontiformes
Lifecycle Berg, 1940 [2]
Classification Subgroups
Fossil record
Use in research Geotriidae
In human culture
As food
Mordaciidae
As pests Petromyzontidae Risso, 1827
In folklore
(Northern lampreys)
In literature
† = extinct
References
Further reading
General
Research on pheromones for pest control
External links

Distribution
Lampreys live mostly in coastal and fresh waters and are found in most temperate regions except those in Africa. Some species (e.g.
Geotria australis, Petromyzon marinus, and Entosphenus tridentatus) travel significant distances in the open ocean,[9] as evidenced
by their lack of reproductive isolation between populations. Other species are found in land-locked lakes. Their larvae (ammocoetes)
have a low tolerance for high water temperatures, which may explain why they are not distributed in the tropics.
Lamprey distribution may be adversely affected by overfishing and pollution. In
Britain, at the time of the Conquest, lampreys were found as far upstream in the
River Thames as Petersham. The reduction of pollution in the Thames and River
Wear has led to recent sightings in London andChester-le-Street.[10][11]

Distribution of lampreys may also be adversely affected by dams and other

construction projects due to disruption of migration routes and obstruction of access
to spawning grounds. Conversely, the construction of artificial channels has exposed
new habitats for colonisation, notably in North America where sea lampreys have
become a significant introduced pest in the Great Lakes. Active control programs to Mouth of a sea lamprey, Petromyzon
control lampreys are undergoing modifications due to concerns of drinking water marinus
quality in some areas.[12]

Biology

Basic external anatomy of a lamprey

Adults superficially resemble eels in that they have scaleless, elongated bodies, and
can range from 13 to 100 cm (5 to 40 inches) in length. Lacking paired fins, adult
lampreys have large eyes, one nostril on the top of the head, and seven gill pores on
Microscopic cross section through
each side of the head. the pharynx of a larva from an
unknown lamprey species
The pharynx is subdivided; the ventral part forming a respiratory tube that is isolated
from the mouth by a valve called the velum. This is
an adaptation to how the adults feed, by preventing
the prey's body fluids from escaping through the gills
or interfering with gas exchange, which takes place
by pumping water in and out of the gill pouches
instead of taking it in through the mouth. Sea lamprey, Petromyzon marinus

Near the gills are the eyes, which are poorly

developed and buried under skin in the larvae. The eyes complete their development during metamorphosis, and are covered by a thin
[13]
and transparent layer of skin that becomes opaque in preservatives.

The unique morphological characteristics of lampreys, such as their cartilaginous skeleton, suggest they are the sister taxon (see
cladistics) of all living jawed vertebrates (gnathostomes), and are usually considered the most basal group of the Vertebrata. Instead
of true vertebrae, they have a series of cartilaginous structures called arcualia arranged above the notochord. Hagfish, which resemble
lampreys, have traditionally been considered the sister taxon of the true vertebrates (lampreys and gnathostomes)[14] but DNA
[15]
evidence suggests that they are in fact the sister taxon of lampreys.

Studies have shown that lampreys are amongst the most energy-efficient swimmers. Their swimming movements generate low-
.[16]
pressure zones around their body, which pull rather than push their bodies through the water
The last common ancestor of lampreys appears to have been specialized to feed on the blood and body fluids of other fish after
metamorphosis. They attach their mouthparts to the target animal's body, then use three horny plates (laminae) on the tip of their
piston-like tongue, one transversely and two longitudinally placed, to scrape through surface tissues until they reach body fluids. The
teeth on their oral disc are primary used to help the animal attach itself to its prey. Made of keratin and other proteins, lamprey teeth
have a hollow core to give room for replacement teeth growing under the old ones. Some of the original blood-feeding forms have
evolved into species that feed on both blood and flesh, and some who have become specialized to eat flesh and may even invade the
internal organs of the host. Tissue feeders can also involve the teeth on the oral disc in the excision of tissue.[17][18][19][20][21][22]
[23] As a result, the flesh-feeders have smaller buccal glands as they don't require to produce anticoagulant continuously and
mechanisms for preventing solid material entering the branchial pouches, which could otherwise potentially clogg the gills.[24] A
study of the stomach content of some lampreys has shown the remains of intestines, fins and vertebrae from their prey.[25] Although
attacks on humans do occur,[26] they will generally not attack humans unless starved.
[27][14]

Carnivorous forms have given rise to the non-carnivorous species,[28] and "giant" individuals amongst the otherwise small American
brook lamprey have occasionally been observed, leading to the hypothesis that sometimes individual members of non-carnivorous
[29]
forms return to the carnivorous lifestyle of their ancestors.

Research on sea lampreys has revealed that sexually mature males use a specialized heat-producing tissue in the form of a ridge of fat
cells near the anterior dorsal fin to stimulate females. After having attracted a female with pheromones, the heat detected by the
[30]
female through body contact will encourage spawning.

Lampreys provide valuable insight into adaptive immune systems, as they possess a convergently evolved adaptive immunity with
cells that function like the T cells and B cells seen in higher vertebrates. Lamprey leukocytes express surface variable lymphocyte
receptors (VLRs) generated from somatic recombination of leucine-rich repeat gene segments in a recombination activating gene-
independent manner.[31][32]

Northern lampreys (Petromyzontidae) have the highest number ofchromosomes (164–174) among vertebrates.[33]

Pouched lamprey (Geotria australis) larvae also have a very high tolerance for free iron in their bodies, and have well-developed
biochemical systems for detoxification of the large quantities of these metal ions.[34]

Lampreys are the only extant vertebrate to have four eyes.[35] Most lampreys have two additional parietal eyes: a pineal and
parapineal one (the exception is members ofMordacia).[36]

Lifecycle
The adults spawn in nests of sand, gravel and pebbles in clear streams, and after
hatching from the eggs, young larvae—called ammocoetes—will drift downstream
with the current till they reach soft and fine sediment in silt beds, where they will
burrow in silt, mud and detritus, taking up an existence as filter feeders, collecting
detritus, algae, and microorganisms.[37] The eyes of the larvae are underdeveloped,
but are capable of discriminating changes in illuminance.[38] Ammocoetes can grow
from 3–4 inches (8–10 cm) to about 8 inches (20 cm).[39][40] Many species change
color during a diurnal cycle, becoming dark at day and pale at night.[41] The skin Larva of an unknown lamprey
also has photoreceptors, light sensitive cells, most of them concentrated in the tail, species
which helps them to stay buried.[42] Lampreys may spend up to eight years as
ammocoetes,[43] while species such as the Arctic lamprey may only spend one to
two years as larvae,[44] prior to undergoing a metamorphosis which generally lasts 3–4 months, but can vary between species.[45]
While metamorphosing, they do not eat.[46]

The rate of water moving across the ammocoetes' feeding apparatus is the lowest recorded in any suspension feeding animal, and
they therefore require water rich in nutrients to fulfill their nutritional needs. While the majority of (invertebrate) suspension feeders
thrive in waters containing under 1 mg suspended organic solids per litre (>1 mg/l), ammocoetes demand minimum 4 mg/l, with
 [47]
concentrations in their habitats having been measured up to 40 mg/l.

During metamorphosis the lamprey loses both the gallbladder and the biliary tract,[48] and the endostyle turns into a thyroid
gland.[49]

Some species, including those that are not carnivorous and do not feed even following metamorphosis,[46] live in freshwater for their
entire lifecycle, spawning and dying shortly after metamorphosing.[50] In contrast, many species are anadromous and migrate to the
sea,[46] beginning to prey on other animals while still swimming downstream after their metamorphosis provides them with eyes,
teeth, and a sucking mouth.[51][50] Those that are anadromous are carnivorous, feeding on fishes or marine mammals.
[9][52][53]

Anadromous lampreys spend up to four years in the sea before migrating back to freshwater, where they spawn. Adults create nests
(called redds) by moving rocks, and females release thousands of eggs, sometimes up to 100,000. The male, intertwined with the
female, fertilizes the eggs simultaneously.Being semelparous, both adults die after the eggs are fertilized.[50]

Classification
Taxonomists place lampreys and hagfish in the subphylum Vertebrata of the phylum
Chordata, which also includes the invertebrate subphyla Tunicata (sea-squirts) and
the fish-like Cephalochordata (lancelets or Amphioxus). Recent molecular and
morphological phylogenetic studies place lampreys and hagfish in the superclass
Agnatha or Agnathostomata (both meaning without jaws). The other vertebrate
superclass is Gnathostomata (jawed mouths) and includes the classes
Chondrichthyes (sharks), Osteichthyes (bony fishes), Amphibia, Reptilia, Aves, and
Mammalia.
Ammocoetes larva of Lethenteron
Some researchers have classified lampreys as the sole surviving representatives of reissneri
the Linnean class Cephalaspidomorphi.[54] Cephalaspidomorpha is sometimes given
as a subclass of the Cephalaspidomorphi. Fossil evidence now suggests lampreys
and cephalaspids acquired their shared characters by convergent evolution.[55][56]
As such, many newer works, such as the fourth edition of Fishes of the World,
classify lampreys in a separate group called Hyperoartia or Petromyzontida,[54] but
whether this is actually a clade is disputed. Namely, it has been proposed that the
non-lamprey "Hyperoartia" are in fact closer to thejawed vertebrates.

The debate about their systematics notwithstanding, lampreys constitute a single

Several species of European
order Petromyzontiformes. Sometimes still seen is the alternative spelling lampreys
"Petromyzoniformes", based on the argument that the type genus is Petromyzon
and not "Petromyzonta" or similar. Throughout most of the 20th century, both names
were used pretty much indiscriminately, even by the same author in subsequent
publications. In the mid-1970s, the ICZN was called upon to fix one name or the
other, and after much debate had to resolve the issue by voting. Thus, in 1980, the
spelling with a "t" won out, and in 1981, it became official that all higher-level taxa
based on Petromyzon have to start with "Petromyzont-".

The following taxonomy is based upon the treatment by FishBase as of April 2012
with phylogeny compiled by Mikko Haaramo.[57] Within the order are 10 living Pouched lamprey
genera in three families. Two of the latter are monotypic at genus level today, and in
one of them a single living species is recognized (though it may be a cryptic species
complex):[58]

Petromyzontiformes Geotriidae Geotria Gray 1851 (pouched lamprey)

Mordaciidae
 Mordaciidae Mordacia Gray 1853 (southern topeyed lampreys)

Petromyzon Linnaeus 1758 (Sea lamprey)

Petromyzontinae
Ichthyomyzon Girard 1858

Caspiomyzon Berg 1906 (Caspian lamprey)

Tetrapleurodon Creaser & Hubbs 1922

Petromyzontidae Entosphenini Entosphenus Gill 1863

Lampetrinae
Lethenteron Creaser & Hubbs
1922

Lampetrini
Eudontomyzon Regan 1911

Lampetra Bonnaterre 1788

List of species, from FishBase in 2017.[59][60] Click "show" to [show]

expand.

Geotria australis Gray 1851 (Pouched lamprey)

Mordacia lapicida (Gray 1851) (Chilean lamprey)
Mordacia mordax (Richardson 1846) (Australian lamprey)
Mordacia praecox Potter 1968 (Non-parasitic/Australian brook lamprey)
Petromyzon marinus Linnaeus 1758 (Sea lamprey)
Ichthyomyzon bdellium(Jordan 1885) (Ohio lamprey)
Ichthyomyzon castaneusGirard 1858 (Chestnut lamprey)
Ichthyomyzon fossor Reighard & Cummins 1916(Northern brook lamprey)
Ichthyomyzon gagei Hubbs & Trautman 1937 (Southern brook lamprey)
Ichthyomyzon greeleyiHubbs & Trautman 1937 (Mountain brook lamprey)
Ichthyomyzon unicuspisHubbs & Trautman 1937 (Silver lamprey)
Caspiomyzon wagneri(Kessler 1870) Berg 1906(Caspian lamprey)
Caspiomyzon graecus(Renaud & Economidis 2010)(Ionian brook lamprey)
Caspiomyzon hellenicus(Vladykov et al. 1982) (Greek lamprey)
Tetrapleurodon geminisÁlvarez 1964 (Mexican brook lamprey)
Tetrapleurodon spadiceus(Bean 1887) (Mexican lamprey)
Entosphenus folletti Vladykov & Kott 1976 (Northern California brook lamprey)
Entosphenus lethophagus(Hubbs 1971) (Pit-Klamath brook lamprey)
Entosphenus macrostomus(Beamish 1982) (Lake lamprey)
Entosphenus minimus(Bond & Kan 1973) (Miller Lake lamprey)
Entosphenus similis Vladykov & Kott 1979 (Klamath river lamprey)
Entosphenus tridentatus(Richardson 1836) (Pacific lamprey)
Lethenteron alaskenseVladykov & Kott 1978 (Alaskan brook lamprey)
Lethenteron appendix (DeKay 1842) (American brook lamprey)
Lethenteron camtschaticum(Tilesius 1811) (Arctic lamprey)
Lethenteron kessleri (Anikin 1905) (Siberian brook lamprey)
Lethenteron ninae Naseka, Tuniyev & Renaud 2009(Western Transcaucasian lamprey)
Lethenteron reissneri (Dybowski 1869) (Far Eastern brook lamprey)
Lethenteron zanandreai(Vladykov 1955) (Lombardy lamprey)
Eudontomyzon stankokaramani(Karaman 1974) (Drin brook lamprey)
Eudontomyzon morii (Berg 1931) (Korean lamprey)
Eudontomyzon danfordiRegan 1911 (Carpathian brook lamprey)
 Eudontomyzon danfordiRegan 1911 (Carpathian brook lamprey)
Eudontomyzon mariae(Berg 1931) (Ukrainian brook lamprey)
Eudontomyzon vladykovi(Oliva & Zanandrea 1959)(Vladykov's lamprey)
Lampetra aepyptera (Abbott 1860) (Least brook lamprey)
Lampetra alavariensisMateus et al. 2013 (Portuguese lamprey)
Lampetra auremensis Mateus et al. 2013 (Qurem lamprey)
Lampetra ayresi (Günther 1870) (Western river lamprey)
Lampetra fluviatilis (Linnaeus 1758) (European river lamprey)
Lampetra hubbsi (Vladykov & Kott 1976)(Kern brook lamprey)
Lampetra lanceolata Kux & Steiner 1972 (Turkish brook lamprey)
Lampetra lusitanica Mateus et al. 2013 (lusitanic lamprey)
Lampetra pacifica Vladykov 1973 (Pacific brook lamprey)
Lampetra planeri (Bloch 1784) (European brook lamprey)
Lampetra richardsoni Vladykov & Follett 1965(Western brook lamprey)

Fossil record
Lamprey fossils are rare because
cartilage does not fossilize as
readily as bone. The first fossil
lampreys were originally found in Mayomyzon, one of the oldest known
lampreys
Early Carboniferous limestones,
marine sediments in North
America: Mayomyzon pieckoensis and Hardistiella montanensis, from the
Mississippian Mazon Creek lagerstätte and the Bear Gulch limestone sequence.
None of the fossil lampreys found to date have been longer than 10 cm (3,9
inches),[61] and all the Paleozoic forms have been found in marine deposits.
[62]

In the 22 June 2006 issue of Nature, Mee-mann Chang and colleagues reported on a
fossil lamprey from the Yixian Formation of Inner Mongolia. The new species,
morphologically similar to Carboniferous and other forms, was given the name
Jamoytius kerwoodi, a putative Mesomyzon mengae ("Meng Qingwen's Mesozoic lamprey").
lamprey relative from theSilurian
The exceedingly well-preserved fossil showed a well-developed sucking oral disk, a
relatively long branchial apparatus showing a branchial basket, seven gill pouches,
gill arches, and even the impressions of gill filaments, and about 80 myomeres of its musculature. Unlike the North American fossils,
its habitat was almost certainly fresh water.[63]

Months later, a fossil lamprey even older than the Mazon Creek genera was reported from Witteberg Group rocks near Grahamstown,
in the Eastern Cape of South Africa. This species,Priscomyzon riniensis, is very similar to lampreys found today.[64][65][66]

Use in research
The lamprey has been extensively studied because its relatively simple brain is thought in many respects to reflect the brain structure
of early vertebrate ancestors. Beginning in the 1970s, Sten Grillner and his colleagues at the Karolinska Institute in Stockholm
followed on from extensive work on the lamprey started by Carl Rovainen in the 1960s that used the lamprey as a model system to
[68]
work out the fundamental principles of motor control in vertebrates starting in the spinal cord and working toward the brain.

In a series of studies by Rovainen and his student James Buchanan, the cells that formed the neural circuits within the spinal cord
capable of generating the rhythmic motor patterns that underlie swimming were examined. Note that there are still missing details in
the network scheme despite claims by Grillner that the network is characterised (Parker 2006, 2010[69][70] ). Spinal cord circuits are
controlled by specific locomotor areas in the brainstem and midbrain, and these areas are in turn controlled by higher brain structures,
including the basal ganglia andtectum.
 In a study of the lamprey tectum published in 2007,[71] they found electrical
stimulation could elicit eye movements, lateral bending movements, or swimming
activity, and the type, amplitude, and direction of movement varied as a function of
the location within the tectum that was stimulated. These findings were interpreted
as consistent with the idea that the tectum generates goal-directed locomotion in the
lamprey.

Lampreys are used as a model organism in biomedical research, where their large
reticulospinal axons are used to investigate synaptic transmission.[72] The axons of
lamprey are particularly large and allow for microinjection of substances for
experimental manipulation.
Stimulation of the olfactory sensory
They are also capable of full functional recovery after complete spinal cord
neurons in the periphery activates
neurons in the olfactory bulb of a sea transection. Another trait is the ability to delete several genes from their somatic cell
lamprey[67] lineages, about 20% of their DNA, which are vital during development of the
embryo, but which in humans can cause problems such as cancer later in life, after
they have served their purpose. How the genes destined for deletion are targeted is
not yet known.[73][74]

In human culture

As food
Lampreys have long been used as food for humans.[75] They were highly
appreciated by the ancient Romans. During the Middle Ages, they were widely eaten
by the upper classes throughout Europe—especially during Lent when eating meat
was prohibited, on account of their meaty taste and texture. King Henry I of England
is claimed to have been so fond of lampreys that he often ate them late into life and
poor health against the advice of his physician concerning their richness, and is said
to have died from eating "a surfeit of lampreys". Whether or not his lamprey
.[76]
indulgence actually caused his death is unclear

Portuguese lamprey rice

On 4 March 1953, Queen Elizabeth II's coronation pie was made by the Royal Air
Force using lampreys.[77]

In southwestern Europe (Portugal, Spain, and France), the northern half of Finland
and in Latvia (where lamprey is routinely sold in supermarkets), larger lampreys are
still a highly prized delicacy. Sea lamprey is the most sought-after species in
Portugal and one of only two that can legally bear the commercial name "lamprey"
(lampreia): the other one being Lampetra fluviatilis, the European river lamprey,
both according to Portaria (Government regulation no. 587/2006, from 22 June).
Overfishing has reduced their number in those parts. Lampreys are also consumed in
Sweden, Russia, Lithuania, Estonia, Japan, and South Korea. In Finland, they are
commonly sold pickled in vinegar.[78] Yatsume kabayaki in Japan

The mucus and serum of several lamprey species, including the Caspian lamprey
(Caspiomyzon wagneri), river lampreys (Lampetra fluviatilis and L. planeri), and sea lamprey (Petromyzon marinus), are known to
[79][80]
be toxic, and require thorough cleaning before cooking and consumption.

In Britain, lampreys are commonly used asbait, normally as dead bait. Northern pike, perch, and chub all can be caught on lampreys.
Frozen lampreys can be bought from most bait andtackle shops.
As pests
Sea lampreys have become a major pest in the North American Great
Lakes. It is generally believed that they gained access to the lakes via
canals during the early 20th century,[81][82] but this theory is
controversial.[83] They are considered an invasive species, have no
natural enemies in the lakes, and prey on many species of
commercial value, such aslake trout.[81]

Lampreys are now found mostly in the streams that feed the lakes,
and controlled with special barriers to prevent the upstream
movement of adults, or by the application of toxicants called
lampricides, which are harmless to most other aquatic species;
however, these programs are complicated and expensive, and do not
eradicate the lampreys from the lakes, but merely keep them in Lampreys attached to alake trout.
check.[84]

New programs are being developed, including the use of chemically sterilized male lampreys in a method akin to the sterile insect
technique.[85] Finally, pheromones critical to lamprey migratory behaviour have been isolated, their chemical structures determined,
and their impact on lamprey behaviour studied, in the laboratory and in the wild, and active forts
ef are underway to chemically source
[86][87][88]
and to address regulatory considerations that might allow this strategy to proceed.

Control of sea lampreys in the Great Lakes is conducted by the U.S. Fish and Wildlife Service and the Canadian Department of
Fisheries and Oceans, and is coordinated by the Great Lakes Fishery Commission.[89] Lake Champlain, bordered by New York,
Vermont, and Quebec, and New York's Finger Lakes are also home to high populations of sea lampreys that warrant control.[90] Lake
Champlain's lamprey control program is managed by the New York State Department of Environmental Conservation, the Vermont
Department of Fish and Wildlife, and the U.S. Fish and Wildlife Service.[90] New York's Finger Lakes sea lamprey control program
is managed solely by the New York State Department of Environmental Conservation.[90]

In folklore
In folklore, lampreys are called "nine-eyed eels". The name is derived from the seven external gill slits that, along with one nostril
and one eye, line each side of a lamprey's head section. Likewise, the German word for lamprey is Neunauge, which means "nine-
eye",[91] and in Japanese they are called yatsume-unagi (八つ目鰻, "eight-eyed eels"), which excludes the nostrils from the count. In
British folklore, the monster known as the Lambton Worm may have been based on a lamprey, since it is described as an eel-like
creature with nine eyes.

In literature
Vedius Pollio kept a pool of lampreys into which slaves who incurred his displeasure would be thrown as food.[92] On one occasion,
Vedius was punished byAugustus for attempting to do so in his presence:

...one of his slaves had broken a crystal cup. Vedius ordered him to
be seized and then put to death, but in an unusual way. He ordered
him to be thrown to the huge lampreys which he had in his fish
pond. Who would not think he did this for display? Yet it was out of
cruelty. The boy slipped from the captor's hands and fled to
Augustus' feet asking nothing else other than a different way to die –
he did not want to be eaten. Augustus was moved by the novelty of
the cruelty and ordered him to be released, all the crystal cups to be
broken before his eyes, and the fish pond to be filled in...

[93]
 — Seneca, On Anger, III, 40[93]

This incident was incorporated into the plot of the 2003 novel Pompeii by Robert
Harris in the incident of Ampliatus feeding a slave to his lampreys.

Lucius Licinius Crassus was mocked by Gnaeus Domitius Ahenobarbus (cos. 54

BC) for weeping over the death of his pet lamprey:

So, when Domitius said to Crassus the orator, Did not you weep for
the death of the lamprey you kept in your fish pond? – Did not you,
said Crassus to him again, bury three wives without ever shedding a
tear? – Plutarch, On the Intelligence of Animals, 976a[94]

This story is also found in Aelian (Various Histories VII, 4) and Macrobius Illustration from an edition of
(Saturnalia III.15.3). It is included by Hugo von Hofmannsthal in the Chandos Tacuinum Sanitatis, 15th century
Letter:

And in my mind I compare myself from time to time with the orator Crassus, of whom it is reported that he grew so
excessively enamoured of a tame lamprey – a dumb, apathetic, red-eyed fish in his ornamental pond – that it became
the talk of the town; and when one day in the Senate Domitius reproached him for having shed tears over the death of
this fish, attempting thereby to make him appear a fool, Crassus answered, "Thus have I done over the death of my
fish as you have over the death of neither your first nor your second wife."

I know not how oft this Crassus with his lamprey enters my mind as a mirrored image of my Self, reflected across the
abyss of centuries.

— Philip, Lord Chandos, (fictional) younger son of the Earl of Bath, in a letter to Francis
Bacon[95]

In George R. R. Martin's novel series, A Song of Ice and Fire, Lord Wyman Manderly is mockingly called "Lord Lamprey" by his
subjects in reference to his rumored affinity to lamprey pie and his strikingobesity.[96]

Kurt Vonnegut, in his late short story "The Big Space Fuck", posits a future America so heavily polluted – "Everything had turned to
shit and beer cans", in his words – that the Great Lakes have been infested with a species of massive, ambulatory lampreys; all three
.[97]
of the main characters are eaten by lampreys on leaving a house near the end of the story

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Further reading

General
Renaud, C.B. (2011) Lampreys of the world. An annotated and illustrated catalogue of lamprey species known to
date FAO Species Catalogue for Fishery Purposes. No. 5. Rome. ISBN 978-92-5-106928-8.

Research on pheromones for pest control

Sorensen, Peter W.; Fine, Jared M.; Dvornikovs, Vadims; Jeffrey, Christopher S.; Shao, Feng; Wang, Jizhou; Vrieze,
Lance A.; Anderson, Kari R.; Hoye, Thomas R. (2005). "Mixture of new sulfated steroids functions as a migratory
pheromone in the sea lamprey".Nature Chem. Biol. 1 (6): 324–328. doi:10.1038/nchembio739. PMID 16408070.,
see [4], accessed 1 July 2015. [Primary source example.]
Dittman, Andrew (2005). "News and Views: Chemical cues for sea lamprey migration".Nature Chem. Biol.
(Submitted manuscript).1 (6): 316–317. doi:10.1038/nchembio1105-316. PMID 16408065., see [5], accessed 1 July
2015. [Lay summary of Sorensen, et al. (2005)]
Johnson, Nicholas S.; Yun, Sang-Seon; Thompson, Henry T.; Brant, Cory O.; Li, Weiming (2009). "A synthesized
pheromone induces upstream movement in female sea lamprey and summons them into traps" . Proc. Natl. Acad.
Sci. U.S.A. 106 (4): 1021–1026. doi:10.1073/pnas.0808530106. PMC 2633527. PMID 19164592. [Primary source
example.]
Richard Black, 2009, "Sex smell lures 'vampire' to doom,"BBC News (online), 20 January 2009, see[6], accessed 1
July 2015. [Lay summary of Johnson, et al. (2009); Subtitle: "A synthetic 'chemical sex smell' could help rid North
America's Great Lakes of a devastating pest, scientists say
."]

External links
Media related to Petromyzontiformes at Wikimedia Commons
"ITIS report on the lampreys". ITIS. Retrieved 27 September 2012.
"Lamprey". Inland Fisheries Ireland. Retrieved 27 September 2012.
"The Tree of Life". Retrieved 27 September 2012. A Tree of Life diagram showing the relation of Lampreys to other
organisms.

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