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The Lateral Line in Fish: Structure, Function, and Role in Behavior

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Journal of Ichthyology, Vol. 43, Suppl. 2, 2003, pp. S175–S213.
Original Russian Text Copyright © 2003 by Kasumyan.
English Translation Copyright © 2003 by MAIK “Nauka /Interperiodica” (Russia).

The Lateral Line in Fish: Structure, Function,

and Role in Behavior
A. O. Kasumyan
Moscow State University, Vorob’evy gory, Moscow, 119899 Russia
Received April 9, 2003

Abstract—This paper reviews the recent studies of the lateral line in fishes. The structure of free canal neuro-
masts, their receptor units, lateral line canals, vesicle of Savi, innervation, size, and the distribution of the lateral
line organs in fishes with various life modes and systematics. The functional parameters of the lateral line, such
as the spectral and differential sensitivity of free and canal neuromasts are analyzed along with their functional
specialization, the nature and sources of adequate stimuli for the receptors of the lateral line, morphological
adaptations directed to increase the sensitivity and widening the spectrum of the stimuli perceived (the size of
the cupula, papillae, links between the lateral line canals and the swimming bladder, intra-canal septa and nar-
rowing of the canals near the neuromasts zones, elastic membranes closing the canals, etc.). The data are present
on the ability of fish to determine the source, of hydrodynamic fluctuations located in the water column, at the
water surface or hidden within the bottom substrate, and the distance to the source, the distance of lateral line
sensitivity in different fish species. The role of the lateral line in the feeding, defensive, schooling, reproductive,
and parental behavior of fish is considered in detail. We also review the responses of fish to the water current,
their orientation, and ways of hydrodynamic camouflage. The dynamics of the neuromasts appearance during
the ontogeny is traced, along with the development of the lateral line canals, and the development of its function
in the ontogeny, effects of pollutants on the behavior of fish, managed by the lateral line.

The lateral line, or the seismosensory system, is a
highly specialized mechanoreceptor system found in
most lower vertebrates (anamnia)—cyclostomata, all
classes of fishes, and most amphibians (Northcutt,
1989; Coombs et al., 1992). According to many inves-
tigators, the lateral line is the evolutionary precursor of
more complex mechanoreceptor systems, auditory and
vestibular (van Bergeijk, 1967; Jørgensen, 1989),
although this view is doubted (Popper et al., 1992; Pop-
per and Platt, 1993). All these systems have a common
origin because they develop from the same anlage
(ectodermic swelling): lateral line from the cells of pre-
and postauditory placodes, and the sensory structures
of the labyrinth, from cells of the central auditory pla-
code (Sand, 1984). Receptor elements of the lateral
line, as well as the auditory and vestibular system,
include typical hair cells. Because of the common ori-
gin and the fact that the peripheral region is formed by
the same type of receptor cells, these systems are often
combined under the same title: acoustic-lateral system
or octavo-lateral system. Electoreception was also for a
long time considered a part of a specialized system of
the lateral line (Dijkgraaf, 1963). Only recently, the
electroreception, the sensory elements of which
involves specialized hair cells, was considered as a
unique sensory system. Morphological studies con-
ducted during recent years allow us to think that elec-
troreception and the lateral line existed as individual
sensory systems in primitive forms of vertebrates,
ancestral to fish and fish-like organisms (Fishelson and
Baranes, 1998).
The lateral line is a well studied sensory system in
fish. Studies in this area are conducted in many coun-
tries. In Russia, detailed studies of the lateral line were
started in the 19th century by P.I. Mitrofanov (1892),
who used a large amount of material in lampreys,
sharks and teleost fish to show that the lateral line has a
common origin with the auditory system. In the begin-
ning of the 20th century, these studies were continued
by N.K. Koltsov (1901), D.K. Tretyakov (1915, 1944),
V. Vladykov (1926, cit. in Disler, 1960). N.N. Disler
(1960, 1977) who contributed very much in the studies
of the development of the lateral line organ. Studies of
V.M. Makushok (1958, 1961), A.V. Neelov (1979),
V.G. Sideleva (1982), S.A. Smirnov (1969; Disler and
Smirnov, 1977), N.V. Parin and D.A. Astakhov (Asta-
khov, 1980; Parin and Astakhov, 1982), A.V. Balushkin
(1996), S.A. Evseenko (2000) and others were made in
the field of morphology of the lateral line. Works by
G.A. Malyukina (1955, 1956, 1958), Yu.N. Andriyanov
and O.B. Il’inskii (1983), D.S. Pavlov and
S.N. Tyuryukov (Pavlov, 1979; Pavlov and Tyuryukov,
1988, 1993, 1995) were directed to analysis of the func-
tional parameters of the lateral line. However, during
recent decades, the intensity of the studies of the lateral
line, especially its functional characteristics, became
sporadic in Russia. Reviews are rare (Pavlov and
Kasumyan, 1994). In this paper we describe the basic
results of modern studies of the lateral line, character-
istics and diversity of its structural organization, func-
tional parameters, and its role in different kinds of
behavior.

S175
S176 KASUMYAN

(‡) (b)

10 µm
stereocilia kinocilium

nucleus

cupula

afferent efferent
nerve nerve

hair kinocilium
cells

stereocilia

foot synaps
cells

Fig. 1. The basic structural components of the neuromast (a) (following the Iwai, 1967, scheme) and the scheme of the hair cell (b)
(based on Blaxter, 1987).

STRUCTURAL ORGANIZATION kynocyle is similar to motile flagella, looking like

OF THE LATERAL LINE
Characteristic epidermal structures, neuromasts,
form the peripheral region of the lateral line (Fig. 1a).
Neuromasts (literally sensitive tubercles using outdated
terminology) consist of two basic cell types: hair cells
and supporting cells. Hair cells are the secondary-sen-
sitivity receptor cells having an epidermal origin (Fig.
1b). Each hair cell bears one high kynocyle (5–10 µm)
and 30–150 shorter stereocilia (2–3 µm) at its free sur-
face. The height of stereocilia continuously reduces
from the kynocyle, which is always located excentri-
cally. Such a position of the kynocyle makes the hair
cell morphologically polarized. The structure of the
micro tubules (9+2). Stereocilia lack such structures
(Flock, 1965).
The number of hair cells in neuromasts depends on
its size and differs widely, from several dozen to hun-
dreds and thousands. The average density of hair cells
in neuromasts varies less. For example, in the common
ruffe Gymnocephalus cernuus it is about 38000 per mm2,
in neuromasts of Cotus bairdi, about 50000 per mm2,
and in neuromasts of the australian shark Mustelus ant-
arcticus (Triakidae), 40000–52500 per mm2 (Gray and
Best, 1989; Janssen et al., 1987; Peach and Rouse,
2000). Hair cells in neuromasts are oriented in two
opposite directions. Zones with similarly oriented
receptor cells are absent in neuromasts and, therefore,

JOURNAL OF ICHTHYOLOGY Vol. 43 Suppl. 2 2003

 THE LATERAL LINE IN FISH: STRUCTURE, FUNCTION, AND ROLE IN BEHAVIOR
A V
Fig. 2. Capillary network of canal neuromast in Spicara smaris (Centracanthidae): A, arteries; V, veins (based on Jakubowski,
1966b).
even adjacent hair cells in neuromasts may be oriented in
opposite directions. The ratio of the cells with the opposite
orientation in canal neuromasts is close to 1 : 1.
Each hair cell is surrounded by supporting cells. At
the periphery of the neuromast, where receptor cells are
absent, there are so-called cover cells, separating the
sensory field of the neuromast from the epithelium
(Münz, 1979; Coombs et al., 1988). The surface of the
sensory zone of the neuromast ranges from 40 to 80%
of the overall area of the neuromast base (Jakubowski,
1967). In the basal part, the hair cell has synaptic con-
tacts with afferent and efferent nerve fibers. The affer-
ent fibers, which transmit signals to the neural centers
of the lateral line, ramify at the base of the neuromast
and in many fish species loose their myeline cover.
Each of them contacts with several similarly oriented
hair cells. The central regulation of hair cells is via
efferent fibers (Fig. 1a).
Kynocyle and stereocilia of hair cells are submerged
into transparent and flexible cupula, vertically dominat-
ing above the surface of the sensory epithelium and
having the shape of a cylinder hemisphere or blade. A
gel-like media, creating the cupula, is secreted by non-
receptor cells of the neuromast: the internal part of the
cupula, by the foot cells, the outer layer, by the cover
cells (Münz, 1979). There exist data that the cupula has
a certain net-like internal structure, formed from fila-
ments of high density (Iwai, 1967). In different species
of fish, depending on the location of the neuromast, the
depth of the cupula ranges from 0.1 to 0.7 mm and is
usually well discernible even at low magnification.
Neuromasts are supplied with blood due to a well-
developed network of blood capillaries (Fig. 2)
(Jakubowski, 1966a, 1966b).
Two types of neuromasts are usually distinguished:
free or superficial neuromasts and canal neuromasts.
Free neuromasts are located freely at the surface of the
body, their cupules are washed by the water and are
affected by environmental agents. Among the free neu-
romasts, researchers usually distinguish primary or
paedomorphic neuromasts, which develop from pla-
JOURNAL OF ICHTHYOLOGY Vol. 43 Suppl. 2
S177
V
cods, and secondary or neomorphic neuromasts, which
either bud off from the primary neuromasts or develop
from epidermal cells due to the inducing effects of
afferent nerves (Disler, 1960; O’Connel, 1981; Coombs
et al., 1992). All canal neuromasts are primary neuro-
masts. They are hidden inside epidermal or bony
canals, which are found on the body or on the head of
the fish, and open with special pores. Free and canal
neuromasts differ in structural and functional charac-
teristics, and the role which they play in the behavior of
the fish. Neuromasts, in various forms, can be found in
all species of fish.
Free Neuromasts
Free neuromasts are usually small. They commonly
have a cylindrical cupula and round base, the diameter
of which rarely reach 100 µkm (Münz, 1979; Coombs
et al., 1988). In certain species (Phrinichthys, Neocer-
atias, Astyanax, Gobius) the bases of free neuromasts
are stretched and the cupula is compressed laterally,
coinciding in its section with the neuromast base
(Montgomery et al., 2001). The number of hair cells in
free neuromasts is relatively small, from several dozens
to several hundreds.
Free neuromasts can be found in lampreys
Petromyzontes (in Mixini, typical neuromasts are
absent), they are very numerous in teleost fishes Chon-
drichthyes and in several bony fishes, Clupeidae, Cyp-
rinidae, and in certain Characinidae (Astyanax)
(Schemmel, 1967; Puzdrowski, 1989). Free neuromasts
are located openly on the head and the body of the fish,
in certain species at the caudal fin, most often at the
same level with the surface of the adjacent epithelium
(Fig. 3). Sometimes, e.g., in lampreys, free neuromasts
occur on small tubercles. In gobies Gobius, Poma-
toschistus (Gobiidae), deep-sea anglerfish Neoceratias
(Neoceratidae), Phrinichthys (Diceratiidae) and other
Ceratoidea (Lophiiformes), the tubercles, bearing free
neuromasts, are quite large and look like wide papillae
(Fig. 4) (Marshall, 1986, 1996). In Chondrichthyes and
2003
S178
1 2 3 4
Fig. 3. Basic types of free neuromast positioning: (1) neu-
romast and its cupula are completely hidden in the epider-
mal pit; (2) the base of the neuromast is in the pit, but the
cupula tip is outside; (3) the base of the neuromast is at the
same level with the outer surface of the fish body; (4) neu-
romast is located at an epidermal tubercle or papillae.
C bus-like cupula of the neuromast protect it from the dis-
75 µm
HC
250 µm P
250 µm
Fig. 4. Free neuromast of the black goby Gobius niger at the
top of epidermal papillae. C, cupula; HC, hair cell; P, papil-
lae (based on Marshall, 1986).
certain other fishes (e.g., in the American pike Esox
americanus, channel catfish Ictaluridae, Amia calva,
carp larvae Cyprinus carpio, etc.), free neuromasts may
be located in depresions of the epithelium. In sharks,
rays, chimaeras, dipnoans (Lepidosiren, Neoceratodus,
and Protopterus), Polypteridae and others, neuromasts
are located on small tubercles at the bottom of epithelial
pits or depressions. Free neuromasts in these fish are
called pit organs (Webb and Northcutt, 1997; Maruska,
2001). They have earlier been erroneously considered
electroreceptor structures, but recent detailed studies of
their morphology showed that the pit organs are just
typical neuromasts (Tester and Nelson, 1969; Webb and
Northcutt, 1997; Maruska and Tricas, 1998; Peach and
Rouse, 2000). In lower sharks, chimaeras, and certain
teleosts, pit organs are located on the body in rows,
sometimes submerged in grooves. This was the basis
for the hypothesis that such rows of pit organs represent
analogues of the lateral line canals and that they
appeared during the evolution from typical canals of the
KASUMYAN
lateral line found already in primitive ancestral forms of
vertebrates (Northcutt, 1989). It is hypothesized that, in
fish, the formation of the canals is not terminated by the
development of the epidermal roof, and that the pit
organs represent modified canal neuromasts (Webb and
Northcutt, 1997). Interestingly, in the coelacanth
Latimeria chalumnae, the grooves with pit organs are
covered by a thin, skinny membrane (Hensel and
Balon, 2001).
The system of free neuromasts the blind side of the
European sole Solea vulgaris is very characteristic. The
density of neuromasts here ranges from five to ten per
mm2, they are located orderly at the same distance from
each other and surrounded by high (up to 3 mm in
adults) papillae (Fig. 5). It is considered that the papil-
lae rising over the relatively high (about 100 µm) rhom-
turbances during the movement and feeding of these
bottom fish (Appelbaum and Schemmel, 1983). Similar
papillae surround free neuromasts on the blind side of
the body in New Zeland sole-like flounder Peltorham-
phus novazeelandiae (Roper, 1981). Due to the great
diversity of free neuromasts, a system for their classifi-
cation is suggested (Srivastava and Srivastava, 1968;
Coombs et al., 1988).
Free neuromasts may form groups, either short and
not connecting with each other as in gobies (Gobiidae)
or deep-sea anglers Phyrinchthys (Diceratiidae)
(Fig. 6) (Marshall, 1986). The orientation of the neuro-
masts combined in the same row usually coincides and
is usually perpendicular to the row direction. On the
surface of the body and on the head of the fish, free neu-
romasts are usually found in proximity to lateral line
canals (Coombs et al., 1988). In the latter case, neuro-
masts are oriented parallel or perpendicularly to the
canal direction (paedomorphic or primary free neuro-
masts, see Coombs et al., 1992). Free neuromasts of the
neomorphic type (secondary neuromasts) may have
very different orientation independent of the direction
of the canal (Dijkgraaf, 1963; Münz, 1979; Coombs
et al., 1988; Visher, 1989; Webb and Northcutt, 1997).
Due to the ability to proliferation, their number, espe-
cially in early ontogeny, rapidly increases. The number
and topography of free neuromasts on the head varies
less than on the body.
It was counted that in the goldfish, Carassius aura-
tus, the number of free neuromasts each side of the
body is about 1000, whereas the number of canal neu-
romasts is significantly smaller and ranges from 52 to
60 (Puzdrowski, 1989). The number of free neuromasts
is higher than canal neuromasts in cods (Gadidae)
(Table 1). However, in many fish, the number of free
neuromasts is small or they are almost absent, as, for
example, in Siluridae and Mormyridae (Lekander,
1949; Harder, 1968), which is usually linked with the
presence of numerous ampular electroreceptors in these
freshwater fishes, which are homologous to free neuro-
masts (Coombs et al., 1988). In fish living in running
JOURNAL OF ICHTHYOLOGY Vol. 43 Suppl. 2 2003
 THE LATERAL LINE IN FISH: STRUCTURE, FUNCTION, AND ROLE IN BEHAVIOR S179

waters with high turbulence, free neuromasts are usu-

ally not numerous or almost completely absent. In fish (a)
living in standing or slowly moving waters, free neuro-
masts are usually more numerous. The life-style and the
role of the lateral line in the behavior may determine
not only the number but also many structural character-
istics of free neuromasts. For example, in the blind cave
form of Astyanax fasciatus, the capsule of free neuro-
masts is two times deeper than in the nonblind form.
(earlier considered a separate species A. mexicanus)
(Teyke, 1990). Similarly, the depth of the cupula differs
in the species differently adapted to life in underground
water bodies (Poulson, 1963). In blind fish, the deepest
cupules are found on the neuromasts located on the
head (up to 300 µm), they are about three times smaller
on the body, and their depth homogeneously reduces in
the rostro-caudal direction (Teyke, 1990).

Lateral Line Canals

25 µm
Lateral line canals have been known in fish at least
since the 17th century. They were at that time consid-
ered as organs responsible for the secretion of skin
mucus (Stenonis, 1664, cit in Maruska, 2001). Most (b)
fish species have lateral line canals on the head and
body. The body canal usually stretches in the rostro-
caudal direction at the middle of the side part of the
body (mediolateral position) from the temporal zone to
the caudal fin and, in halibuts Hippoglossus, Reinhard-
tius and Atheresthes, even on the fin. In some fish, the
lateral line is shifted in the dorsal (Uranoscopidae, Tra-
chiidae) or ventral (Exocoetidae) direction, could curve
in a species-specific way (Chaetodontidae, Pentocer-
atidae, etc.), usually in proximity to pectoral fins, could
sharply change its direction (Balistidae) and be broken
(Scaridae, Cichlidae) or terminate not reaching the cau-
dal peduncle or the caudal fin (in certain cyprinids and 1 mm
smelts, such as verkhovka Leucaspius delineatus, bit-
terling Rhodeus sericeus amarus, European smelt
Osmerus eperlangus, etc.) (Figs. 7, 8 and 9). Fig. 5. Free neuromasts at the blind surface of the head of
Most species have only one canal on the side surface the European sole Solea vulgaris. (a) Individual neuromast
of the body, but some fish, e.g., greenlings Hexagram- surrounded by numerous epidermal papillae; (b) scheme of
the free neuromast (dark circles) and epidermal papillae
midae, Xiphister (Stichaeidae) and Notothenioidei have positioning (light circles) (based on Appelbaum and
several parallel canals on each side of the body, four– Schemmel, 1983).
six and more. Many ancient fossil fish had several
canals of the lateral line (Northcutt, 1989). Among the
living species, many body canals of the lateral line have first case, canals have a relatively large internal open-
been documented in 13 families (Webb, 1989). Addi- ing. They are called the wide canals. In such canals, the
tional branches of the lateral line were found on the bony fornix may be only above the place of the neuro-
caudal fin of halibuts Hippoglossus and Reinhardtius mast location. Canals in bones are much smaller in
(Evseenko, 2000). Dorsal, medial, anal, infraanal, and diameter. On the whole, the width of canals in the fish
other body canals are distinguished, depending on their head ranges widely, from 0.1 to 2–3 mm. The largest
position on the body (Balushkin, 1996). In Clupeidae, canals, up to 7 mm in width, are found in a small bat-
deep-water endemic to Lake Baikal Abyssocottidae, hypelagic fish Poromitra (Melamphaeidae) (Denton
and deep-sea anglers Ceratoidea, body canals are com- and Gray, 1988; Coombs et al., 1988). In Coryphae-
pletely absent (Fig. 9) (Blaxter et al., 1983; Marshall, niodes rupestris, the internal volume of wide canals
1996; Janssen et al., 1999). may be at least 3.5% of the total body volume (Mar-
The head of most fish species have several canals, shall, 1965). Wide canals on the body are characteristic
which could be in the skin and in the cover bones. In the of Brotulidae, Halosauridae, Myctophyidae, Melam-

JOURNAL OF ICHTHYOLOGY Vol. 43 Suppl. 2 2003

S180
(‡)
10 mm
(b)
10 mm
(c)
Fig. 6. Rows of free neuromasts in fish: (a) on the head and the body of Phrynichthys wedli (based on Marshall, 1996); (b) at the
head of Gobius niger; (c) polarization of free nevromasts forming rows at the head of Gobius niger (based on Marshall, 1986).
phaeidae, Evermannellidae, Platytroctidae, and other
inhabitants of oceanic depths. In Holocephali, wide
canals on the head are surrounded by a supporting car-
tilage cover. In most fish species, canals on the head
form a well-developed complex branching system,
which in many cases may occupy almost the whole area
of the head or a significant part (Fig. 10). In certain
Cobitidae, Gobiidae, Cyprinodontidae, and Hemirham-
phidae, canals are absent on the head, as well as on the
body and therefore the lateral line in these fish is repre-
sented only by free neuromasts.
According to location and attribution to certain
bones of the head, lateral line canals are separated into
infraorbital, supraorbital, temporal, mandibular, oper-
cular, preopercular, preoperculo–mandibular, and sev-
eral others. Canals on different sides of the head are
KASUMYAN
linked into a unitary network by the supratemporal
canal or supratemporal commissure (Fig. 11). This pat-
tern is the most general; in many fish some of these
canals may be weakly developed or completely reduced
(infraorbital canal on the eye side of the body in flat-
fishes of the genus Liopsetta, see Voronina and
Evseenko, 2001). There may be other canals, substitut-
ing for or in addition to the above listed, such as the eth-
moidal canal in herrings, hyomandibular canal in rays,
coronal commissure in antarctic Trematomus (Notothe-
niidae) (Jakubowski, 1970). In Clupeidae, Carangidae,
Scombridae, and Istiophoridae, the basic canals are
branching (ramification), creating a complex organized
network of canaliculi on the body (Chubukov et al.,
1987; Blaxter, 1987) (Fig. 12).
JOURNAL OF ICHTHYOLOGY Vol. 43 Suppl. 2 2003
 THE LATERAL LINE IN FISH: STRUCTURE, FUNCTION, AND ROLE IN BEHAVIOR S181

Table 1. The number of free and canal neuromasts on the head and the body in cods (based on Halama, 1977)
Number of neuromasts
Species free canal
on head on body on head on body
Whiting, Odontodagus merlangus 54–66 87–101 32 26–31
Pollack, Pollachius virens 61–85 67–89 32 38–44
Atlantic cod, Gadus morhua 76–91 78–101 32 37–44
Atlantic tomcod, Microgadus tomcod 95–117 90–110 32 45–51
Navaga, Eleginus navaga 113–115 106 32 –
Far Eastern navaga, E. glacilis 118–122 105–111 32 52–54
Arctic cod, Boreogadus saida 121–147 72–81 32 29–35
Siberian cod, Arctogadus borisovi – – 32 –
Polar cod, A. glacialis 168–173 84–85 32 32–34
Wall-eyed pollack, Theragra chalcogramma 110–128 – 32 25–31
Blue whiting, Micromesistius poutassou – – 32 31

In the typical case, canals on the head or the body distance, however there are certain exclusions. For
are connected with the environment by short connect- example, burbot (Lota lota) have pores in the body
ing pipes, leading to pores on the body surface. Pores canal only in the rostral and caudal regions and no
are remote from each other approximately to the same pores, in the middle part of the canal (Hyrtl, 1866, cit.

Salmonidae Clariidae

Esocidae

Trachiidae Uranoscopidae

Exocoetidae Trichiuridae
Fig. 7. Examples of fish having direct mediolateral body canal of the lateral line (Salmonidae, Clariidae, Esocidae) and fish having the
body canal shifted in the dorsal (Trachiidae, Uranoscopidae) or ventral direction (Exocoetidae, Trichiuridae) (from Nelson, 1994).

JOURNAL OF ICHTHYOLOGY Vol. 43 Suppl. 2 2003

S182 KASUMYAN

Carangidae
Centrarchidae

Chaetodontidae Pentoceratidae

Pleuronectidae Balistidae
Fig. 8. Examples of fish having curved (Centrarchidae, Carangidae, Chaetodontidae, Pentoceratidae, Pleuronectidae) or zigzag
(Balistidae) body canal of the lateral line (from Nelson, 1994).

in Sand, 1984). Pores are especially pronounced visu-
ally at the head because of their larger size. For exam-
ple, such relatively small fish (TL 8–10 cm) as Baikal
Batrachocottus baikalensis and stone sculpin Paracot-
tus knerii, the diameter of pores in the mandibular canal
reaches, respectively, 0.6–0.8 and 0.4–0.5 mm (Janssen
et al., 1999). In gobies, large pores have special name—
genipores (genal pores).
In fish covered with scales, e.g., in salmons and
many others, pores of body canals perforate the scales
in several points, which provide for the links between
neuromasts and the environment (Fig. 13). In macker-
els, such scales are much larger and thicker and are
called scutes. In pikes, the scales of the lateral line have
a characteristic notch in the free posterior part and a
depression on the dorsal side for the body neuromasts.
In some fish (cartilaginous fish, certain teleosts, e.g.,
Dorosoma) the basic canal of the lateral line is linked
with the environment via a relatively long canal, which,
in turn, could branch several times and open to the body
surface with several pores (up to ten in Gymnura
micrura) (Maruska, 2001) (Fig. 14). Usually, canal
branching and the formation of accessory canals is
observed in large individuals (Disler, 1977). It is
accepted that this provides for a good connection
between canal neuromasts, whose number remains
constant with the environment regardless of the contin-
ued growth of the fish and the respective increase of the
body canal. In the European sole Solea vulgaris, pores
in the rostral part of the body canal are multiplied soon

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 THE LATERAL LINE IN FISH: STRUCTURE, FUNCTION, AND ROLE IN BEHAVIOR S183

Scaridae Cichlidae

Rhodeus sericeus Leucaspius delineatus

Pomacentridae Hexagrammidae

Clupeidae
Fig. 9. Examples of fish having broken body canal of the lateral line (Scaridae, Cichlidae), incomplete body canal (bitterling Rho-
deus sericeus, verkhovka Leucaspius delineatus, Pomacentridae), having several parallel body canals (Hexagrammidae), and the
fish having no body canals (Clupeidae) (from Nelson, 1994).

after the metamorphosis with the formation of a con-
glomerate of short and narrow tubes (Fig. 15). It is
thought that the sizes of the pores are reduced due to the
increase in their number, which protects the canals of
these fish from pollution by large particles (Appelbaum
and Schemmel, 1983).
In Notopteridae, Mormyridae, and certain other
fish, the pores of the canals are closed with an elastic
membrane isolating them from the environment. Many
cartilaginous fish, especially skates (e.g., G. micrura,
see Fig. 14), have lateral line canals or regions of
canals, in which pores are completely absent, but typi-
cal neuromasts are retained. The link of neuromasts
with the environment is maintained because such
canals or their regions are a part of a system of related
canals most of which have normally developed pores
(Maruska, 2001).
In flatfishes, the systems of canals on different sides
of the body are non-symmetric with respect to each
other and developed to different degrees, more on the
eye side of the body and less on the blind side (Fig. 16).
In skates, canals on each side of the body form similar
systems connecting with each other by special canals
perforating the fish body in the dorso-ventral direction
(Fig. 14). However, regardless of the symmetry, there
are certain differences between the systems of ventral
and dorsal canals of the lateral line. For example, in the
stingray Dasyatis sabina, additional canaliculi on the
dorsal surface of the body are more numerous and more
branching, and the number of pores in the lateral line
here is almost three times more than at the ventral sur-
face of the body (Fig. 17). Some of the canals on the
ventral side of the body (infraorbital) or their long areas
(hyomandibular) lack pores. However, ventral canals

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S184
(‡)
(b)
(Ò)
Fig. 10. Lateral line canals at the head of the common chop
Zingel zingel (a), ruffe Gymnocephalus schraetser (b), and
Hoplostethus mediterraneus (c) (from Jakubowski, 1967,
1974).
have many more neuromasts, and the diameter of these
canals is wider than in the dorsal (Fig. 17). Systems of
the lateral line on various sides of the body differ as
much in other skates: Raja eglanteria, Narcine brasil-
iensis, and G. micrura. Such differences are less pro-
nounced at the head of the hammerhead shark Sphyrna
tiburo (Maruska, 2001). It was proposed, that the struc-
tural characteristics of lateral line canals at the ventral
and dorsal sides of the body are associated with their
involvement in feeding, defensive and reproductive
behavior of the fish.
KASUMYAN
1
2
4
5
Fig. 11. Lateral line canals at the head of the common ruffe
Gymnocephalus cernuus: (1) upper temporal or frontal
commissure; (2) temporal; (3) supraorbital; (4) suborbital;
(5) preoperculo–mandibular. White ovals degignate the
positions of canal neuromasts (from Jakubowski, 1967).
Fig. 12. Network of canals at the head of Sardina pilchardus
(from Wohlfahrt, 1937).
The cavity of the lateral line canals is filled with vis-
cous liquid. Its chemical composition is still inade-
quately studied. Neuromasts are located at the bottom
of the canal at approximately equal distances from the
adjacent pores. Such a location of neuromasts in canals
is considered evolutionary advanced, apomorphic. It is
characteristic of Actinopterigii and such more ancient
fish as coelacanths (Latimeria) and some dipnoans
(Neoceratodus) (Fig. 18). The canal usually becomes
narrower at the neuromast locality (Figs. 19 and 20)
(Jansen et al., 1987). In cephalic canals, perforating the
epidermis, the limited region of the canal with neuro-
masts could be surrounded by a bony cover. Anoplo-
gaster (Anoplogastridae) and Notopterus chitala
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 THE LATERAL LINE IN FISH: STRUCTURE, FUNCTION, AND ROLE IN BEHAVIOR S185

(‡) 3
2 (a)
1 1

4
2
3
6 5 4
(b)

4
1

(b)

1
5
4
(c) 2

5 cm

Fig. 14. Scheme of lateral line canals at the dorsal (a) and
ventral (b) side of the body in Gymnura micrura ray:
(1) supraopercular canal; (2) hyomandibular canal; (3) pos-
terior canals; (4) infraopercular canal; (5) hyomandibular
canal (based on Maruska, 2001).

Fig. 13. Morphological characteristics of the body lateral

line canals in fish with scales. (a) Sagittal section of the
canal: (1) neuromasts; (2) canals perforating the scales;
(3) pores of canals; (4) scale; (5) neural fibers enervating
neuromasts; (6) basic lateral line canal (based on Bond,
1979). (b) Lateral line scales of the American pike Esox
americanus: (1) neuromasts; (4) scales with grooves (based
on Merrilees and Crossman, 1973). (c) Scales of the
mediolateral body canal of Hexagrammos decagrammus,
viewed from above and from aside (based on Wonsettler and
Webb, 1997).

(Notopteridae) have internal epithelial folds or septae

in wide head canals dividing the canal into several com-
partments. Neuromasts in such canals are situated in a
small hole present in each septa (Fig. 19) (Denton and
Gray, 1988; Coombs et al., 1988).
Cartilaginous fish, chimaeras, and bipnoans (Lepi-
dosiren, Protopterus, and many fossil bipnoans) may
have several separate neuromasts in the canal between
adjacent pores. It is the more primitive plesiomorphic
type of neuromast location (Fig. 18). In many sharks
(Squalus, Mustelus, etc.) and chimaeras (Chimaera
monstrosa and Hydrolagus collei), neuromasts are
located close to each other creating a practically unin-
Fig. 15. Multiplied pore in the anterior part of the body
canal in the European sole Solea vulgaris (based on Appel-
baum and Schemmel, 1983).
terruptible sensory field overlapped by a common cup-
ula (Johnson, 1917; Tester and Kendall, 1969; Hama
and Yamada, 1977; Ekström von Lubitz, 1981; Field
et al., 1993). In the ontogeny of these fish, multiple
canal neuromasts between adjacent pores are formed

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S186 KASUMYAN

4 (‡) (b)
2 2

5
1
3 1

7
6
7

Fig. 16. Lateral line canal system on the eye (a) and blind (b) sides of the body in the plaice Pleuronectes platessa: (1) body canal;
(2) supratemporal canal; (3) temporal canal; (4) supratemporal commissure; (5) supraorbital canal; (6) infraorbital canal; (7) preo-
perculo–mandibular canal (based on Voronina, 2002).

from an unitary epidermal anlage, placode) (Johnson,
1917). In greenlings of the genera Hexagrammos and
Pleurogrammus, neuromasts were found only in the
medio–lateral body canal, which is connected to the
cephalic canals. In other body canals of greenlings neu-
romasts are absent. The function of such canals still
remains unclear (Wonsettler and Webb, 1997).
(a)
300
Number of
Canal neuromasts are significantly larger than
superficial neuromasts (Fig. 21). Their length may
reach several mm (Table 2), and the number of hair
cells, several thousands. For example, various authors
counted from 1000 to 5000 hair cells in neuromasts of
the ruffe Gymnocephalus cernuus (van Netten and
Kroese, 1987; Gray and Best, 1989). However, kinoci-
lium and stereocilia of receptor cells of canal neuro-
masts are shorter than in free neuromasts (Münz, 1989;
Song and Northcutt, 1991). Canal neuromasts differ
from those on the surface not only in size, but also other

properties. This is revealed by selective disturbance of

pores

only the canal neuromast’s hair cells when the fish are
150 maintained in the gentamicin solution (0.0002%), an
aminoglycoside antibiotic. Free neuromasts after such
treatment remain intact (Song et al., 1995; Montgom-
0 ery et al., 1997), which is probably associated with the
(b) characteristics of their cupula composition.
1200
Hair cells in canal neuromasts, as well as in free
Number of
neuromasts

neuromasts, are oriented in two mutually opposite

600 directions, but always parallel to the longitudinal axis
of the canal. The maximum deviation of the orientation
of individual cells from the axis of the canal is not more
0 than 10°–15° (Blaxter, 1987). Regions with similarly
oriented cells in neuromasts are absent. Unlike hair
Diameter of canals, µm

1500 cells, the base of canal neuromasts may be oriented

(Ò)
1000
500
0 D V Structural characteristics of the lateral line (the
Fig. 17. Morphological parameters of lateral line canals at
the dorsal (D) and ventral (V) sides of the body in the stin-
gray Dasyatis sabina. (a) Number of pores at the body sur-
face; (b) number of neuromasts in canals; (c) average size of
the hyomandibular canal (based on Maruska, 2001).
along the length of the canal and have an elliptical or
stripe shape, as they are in relatively narrow canals. In
wide canals, the base of neuromasts are situated per-
pendicularly the to longitudinal axis of the canal (e.g.,
ruffe G. cernuus, Aphredoderus sayanus, G. schraetser,
Zingel zingel, Percarina demidoffi) (Fig. 22).
number and pattern of lateral line canals, curvature and
the length of canals, the number of pores etc.) are stable
morphological characters and are often used as key
traits for the species identification of the fish, determi-
nation of intra- and interspecific relationships and char-

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 THE LATERAL LINE IN FISH: STRUCTURE, FUNCTION, AND ROLE IN BEHAVIOR S187

I II

Chondrichthyes
Lepidosiren
Protopterus Polypterus
I Neoceratodus Chondrostei
I Tetrapoda
II Lepisosteus
Latimeria
I? Amia
Teleostei
II

II
Sacropterigii Actinopterygii

Gnatostomata

Fig. 18. Plesiomorphic (I) and apomorphic (II) types of neuromast distribution in lateral line canals in different groups of fish (based
on Webb and Northcutt, 1997).

(a) (b)

(c) (d)

Fig. 19. Scheme of the internal structure of lateral line canal. (a) and (b) respectively, narrow and wide canals at the fish head:
(1) sagittal section; (2) frontal section; (3) transverse section (based on Coombs et al., 1988); (c) wide canal with internal partitions
and thin cover membrane at the head of Anoplogaster cornuta (based on Denton and Gray, 1988); (d) canal with branching canal-
liculi, terminated by pore (based on Coombs et al., 1988).

acteristics of large taxa: genera, families, and suborders
(Andriyashev and Jakubowski, 1971; Nelson, 1994;
Balushkin, 1996; Evseenko, 2000).
Vesicles of Savi and the Spiracular Organ
Skates (Torpedo, Narcine, and Dasyatis) have char-
acteristic structures, the vesicles of Savi, which also
belong to the lateral line system. The origin and func-
tions of Savi vesicles still remain unclear in spite of the
fact that these structures have been described since the
middle of the nineteenth century (Savi, 1844, cit. in
Maruska, 2001). The vesicles of Savi represent closed
cavities, located at the depth 0.5–2.0 mm under the
layer of epidermis, derma, and connective tissue. They

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S188 KASUMYAN

1 row, 45° in the rostro-caudal axis of the fish body. The

2 orientation of the vesicles of adjacent rows is orthogo-
0 1 2 3 4 5 6 romast, and the vesicles are found only on the ventral
cm
Fig. 20. Cast of the inner cavity of the hyomandibular canal
of the lateral line of Dissostichus mawsoni (TL about 1 m):
(1) canal pores; (2) narrowing of the canal cavity near the
neuromast zone (based on Montgomery et al., 1994).
are surrounded by a cartilaginous cover and isolated
from the external environment. The vesicles of Savi are
located in symmetrical rows at the head of skates, but
are most numerous at the ventral surface. Vesicles have
an oval shape and are oriented similarly in the same
nal to each other. Each vesicle usually has one large
central neuromast at the bottom and two smaller neuro-
masts, each having individual cupula and innervation.
(Fig. 23) (Nickel and Fuchs, 1974; Barry and Bennett,
1989; Marushka, 2001). However, in representatives of
the genus Dasyatis, vesicles of Savi have only one neu-
side of the head, and are connected between themselves
and with one canal of the lateral line (Maruska and
Tricas, 1998).
The function of the vesicles of Savi is still unclear.
Electrophysiological data point out and that the maxi-
mum sensitivity of vesicles of Savi is within the range
150–200 Hz (Szabo, 1978), i.e., in a relatively high-fre-
quency part of the spectrum. It is hypothesized (Barry
and Bennett, 1989) that such spectral sensitivity may be
explained by the isolation of the vesicles from the envi-
ronment and low-frequency oscillations. It is proposed

(b)

(a)

(c)

Fig. 21. Relative sizes of free (a) and canal (b, c) neuromasts in Cottus bairdi (based on Coombs, 1997).

1 mm

1 2 3 4 5 6 7 8

Fig. 22. Orientation of canal neuromasts and their relative size in perches: (1) Percarina demidoffi (TL 9.5 cm), (2) Gymnoceph-
alus schraetser (18.5 cm), (3) Gymnocephalus acerinus (15 cm), (4) Gymnocephalus cernuus (18 cm), (5) Zingel zingel (22 cm),
(6) Z. streber (16 cm), (7) Perca fluviatilis (18 cm), (8) Stizostedion lucioperca (20 cm). The sensory zone of the neuromast is
shadowed (based on Jakubowski, 1967).

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 THE LATERAL LINE IN FISH: STRUCTURE, FUNCTION, AND ROLE IN BEHAVIOR S189

Table 2. Maximum size of canal neuromasts and the relative area of their sensory zone in some perciid fishes (based on
Jakubowski, 1967)
Body length, Relative area
Species Maximum neuromast size, mm
cm of the sensory zone, %
River perch, Perca fluviatilis 18 1.3 × 0.5 47
Zander, Stizostedion lucioperca 20 1.4 × 0.6 51
Ruffe, Gumnocephalus cernuus 15 2.0 × 1.0 78
Don ruffe, G. acerinus 18 2.5 × 1.5 45
Ruffe, G. schraetser 18.5 2.8 × 1.7 70
Chop, Zingel zingel 22 2.4 × 0.9 64
Small chop, Z. streber 16 1.0 × 0.8 71
Percarine, Percarina demidoffi 9.5 1.5 × 1.0 37

that fish can use the vesicles of Savi to discover the
source of vibrations within the bottom substrate, to
obtain information about the character of the substrate
during touching, or touching the partner of group or
touching of prey (Nickel and Fuchs, 1974; Barry and
Bennett, 1989; Maruska and Tricas, 1998; Maruska,
2001). These hypotheses are supported by the fact that
the vesicles of Savi are more common in bottom-dwell-
ing fish (skates) and are absent in pelagic fish
(Maruska, 2001).
Cartilaginous and the majority of teleost fishes have
the spiracular organ. It represents an isolated cavity in
the hyoid arch zone. The morphology and the functions
of the spiracular organ are still poorly understood. It
was hypothesized that it could have proprioreceptor
functions and control the movements of the hyoman-
dibular apparatus in fish (Barry and Bennett, 1989).
on the body, and certain other nerves. Ancient fossil
animals had, probably, not less than four nerves in the
lateral line, and some of them had seven nerves (North-
cutt, 1989).
Central nuclei of the lateral line are situated in the
acoustic–lateral area of the medulla above the auditory
centers and close to them, as well as in eminentia gran-
ularis of the cerebellum. Ascending pathways link
these centers with regions of the mesencephalon (torus
(a) 5
6
1
2

INNERVATION OF THE LATERAL

LINE ORGANS
3
It has been considered for a long time that the inner-
vation of the lateral line organ involves the nervous 4
trigeminus (V), n. fascialis (VII), n. glossopharyngeus 7
(IX), and n. vagus (X). It was thought that these nerves 8
unite into the so-called nervous lateralis anterior, inner- 5
vating neuromasts on the head of the fish. Neuromasts (b) 1 mm
of the body and the caudal peduncle was innervated by
n. lateralis posterior, formed by branches of n. glos-
sopharyngeus and n. vulgaris. Some branches of the
n. lateralis posterior may also innervate neuromasts,
located in posterior regions of the head. For example,
one branch of the n. pharyngeus innervates the middle 2
part of the supratemporal canal and the adjacent free 5 cm
neuromasts.
Recently, this scheme of innervation was rejected
(Northcutt, 1989; Coombs et al., 1992). It is thought Fig. 23. Lateral line organs at the dorsal (a) and ventral (b) body
that teleost fishes have at least three separate nerves in sides in a weakly electric ray Narcina brasiliensis; vesicle
the lateral line (n. lateralis) with their own ganglia: dor- of Savi is given within the border. (1) Supraopercular canal;
(2) vesicles of Savi; (3) infraorbital canal;
sal and ventral anterior nerves of the lateral line, inner- (4) hyomandibular canal; (5) electric organ; (6) posterior
vating free and canal neuromasts at the fish head, pos- canal; (7) peripheral neuromasts; (8) central neuromast
terior nerve of the lateral line, enervating neuromasts (based on Maruska, 2001).

JOURNAL OF ICHTHYOLOGY Vol. 43 Suppl. 2 2003

S190 KASUMYAN
semicircularis), and then with the thalamus and telen-
cephalon (McCormick, 1989). The important role of
the cerebellum in the lateral line is illustrated by the
fact that its extirpation causes the loss of previously
acquired and blocks the development of newly condi-
tioned reflexes to lateral line signals (Malyukina,
1958).
FUNCTIONAL CHARACTERISTICS
The function of the lateral line remained unclear for
a long time, even though its structure has been well
known to anatomists and morphologists since the 16th
century. The earliest hypothesis about the role of the
lateral line was that it secretes the skin mucus covering
the body of the fish. Only in the 19th century, Jakobson
(1813, cit. in Bleckmann, 1993) and then Leydig (1850,
1851, cit. in Sand, 1984) determined on the basis of
detailed morphological analysis (in particular, innerva-
(a)
+ its cell membrane is hyperpolarized and the frequency
– of electric impulses significantly increases in the affer-
Hyperpolarization
Depolarization
(b)
1 2 Impulse activity
Fig. 24. Alteration of the sensory potential of hair cell (a)
and impulse activity in the afferent nerve (b) under the
effect of stimuli with opposite direction. 1 and 2, respec-
tively, afferent and efferent nerve fibers; the direction of
stimulation is shown by arrows; “+” and “–” point to the
vector of the receptor potential alteration (based on Blaxter,
1987).
90°
0° 180°
270°
Fig. 25. The intensity of the response in afferent nervous
fibers during the stimulation of the cupula at different
angles to the polarization axis of the neuromast hair cells
(the polarization axis of hair cells 0°–180°) (based on
Bleckmann, 1993).
JOURNAL OF ICHTHYOLOGY
tion of neuromasts) that these structures have a sensory
function. At the beginning of the 20th century, the
American physiologist Parker (1904) found that neuro-
masts are a kind of mechanoreceptors. Several years
later, direct experiments revealed that the lateral line is
used to perceive water currents and stimuli from under-
water moving objects (Hofer, 1908). Later, the validity
of this conclusion was corroborated by many authors,
including the studies by Sven Dijkgraaf. Dijkgraaf pro-
posed to call the sense perceived by the lateral line the
distant touch sense, because the presence of a movable
object is discovered indirectly due to the perception of
water disturbances it causes (Dijkgraaf, 1963). To
honor Sven Dijkgraaf, who did very much in the study
of lateral line functional and structural characteristics,
another name for this sense was proposed “svenning”
(by analogy with hearing) (Platt et al., 1989).
The use of electrophysiological and conditioned
reflex-based methods, and direct observations of fish
behavior, revealed that the lateral line receptors are
detectors of shift waves caused by closely located
sources of underwater oscillations. Reaching the neuro-
mast, the shift wave deflects the cupula and, therefore,
kinocilium and stereocilia of hair cells. If stereocilia are
deflected towards the kinocilium, the cell gets exited,
ent nerve. When the cupula is deflected in the other
direction, the electrical activity of the cell is sup-
pressed, the cell membrane depolarizes and the fre-
quency of electric impulses is reduced (Fig. 24). During
the intermediate effects on the neuromast, the response
is proportional to the cosine of the angle between the
direction of the stimulus and the cell orientation. If the
stimulation occurs parallel to the cell polarization axis,
the response is not evoked. The intensity of the electric
response of hair cells directly depends on the intensity
of the stimulus, i.e., on the amplitude of the oscilla-
tions, which, in the case of a movable object is deter-
mined by the distance to this object, its shape and the
movement speed. The movement direction of the object
is also important when its direction changes to the
opposite, the electrical response of the polarized cell
also switches to the opposite (Mogdans and Bleck-
mann, 1998). It should, however, be noted that the
polarization axis does not coincide with the polariza-
tion direction of the neuromast in all hair cells. This is
why a relatively weak electric response may be caused
in the afferent nerve by stimulation of the source
located at 90° (Fig. 25).
The oscillations perceived by the lateral line recep-
tors are in the low-frequency spectrum range: from 1–5
to 100–200 Hz. Within this range, the areas of the max-
imum sensitivity of canal and free neuromasts differ. In
canal neuromasts, the maximum sensitivity zone is
shifted to a higher frequency zone of the spectrum and
in most fish includes the range from 20–30 to 100 Hz
(Sand, 1981). However, free neuromasts are most sen-
sitive to oscillations with the frequency from 2–5 to 10–
Vol. 43 Suppl. 2 2003
 THE LATERAL LINE IN FISH: STRUCTURE, FUNCTION, AND ROLE IN BEHAVIOR
15 Hz (Fig. 26). In the sculpin Cottus bairdi, for exam-
ple, the maximum sensitivity of canal neuromasts is
observed at 50 Hz, and free neuromasts, 10 Hz
(Coombs and Janssen, 1990). Due to these functional
characteristics, the role of free neuromasts is usually
linked with obtaining information about the presence
and the direction of the water current and with the per-
ception of oscillations caused by large movable objects.
In the current, when free neuromasts are significantly
affected by it, they are not sensitive to weak biologi-
cally significant stimuli. Canal neuromasts, on the other
hand, allow the fish to determine alterations in the
hydrodynamic situation, even in a constant current
(Bleckmann, 1993; Montgomery et al., 1995; Engel-
mann et al., 2000).
Functional specialization of canal neuromasts is
reached, primarily due to their isolation from the envi-
ronment. Low-frequency components of the shift wave
penetrating via the pores to the canal are damped much
easier than higher frequency waves. The damping of
low-frequency oscillations is increased due to such
anatomical characteristics of canals as narrowing in
neuromast zones (Fig. 20), which further increases the
relative efficiency of the higher frequency components
(Montgomery et al., 1995).
It should be noted that the zones of spectral sensitiv-
ity of the lateral line and auditory organs in part coin-
cide. However, the zones of maximum sensitivity are
located in very different parts of the spectrum. The
basic difference between the lateral line and auditory
systems is that the receptors of the lateral line system
respond to shift waves, whereas receptors of the audi-
tory system, to pressure waves. Sound oscillations
(pressure waves) cannot be perceived with the lateral
line because the fish body is acoustically transparent:
the oscillations of the fish body, cupules of neuromasts
and the surrounding water are the same. The absence of
a relative shift (deflection) of the cupula, and therefore
the respective shift of the kinocilium and stereocilia,
makes the evocation of the electric signal impossible.
The stimulation of receptors by pressure waves in the
auditory organ (labyrinth) is reached by the inertia of
the otolith, almost three times more dense than the fish
body.
The lateral line is characterized by high sensitivity
in most fish. Free neuromasts are able to respond to
very weak water currents, with speed from 0.03 mm/s
and higher, canal neuromasts, to alterations of the cur-
rent speed from 0.3 to 20 mm/s2 (Bleckmann, 1993).
The threshold amplitude of oscillations in canal neuro-
masts is 0.1–0.5 µm (Sand, 1981), their sensitivity is
highest to oscillations parallel to the fish body surface
with respect to the orientation of the canal rather than
to oscillations at straight angle to the body. Differential
sensitivity of the lateral line organ is, as determined by
the conditioned reflex method, 3 Hz (Carassius aura-
tus) (Malyukina, 1956). Aplocheilus lineatus, feeding
on fallen insects at the surface, can distinguish oscilla-
JOURNAL OF ICHTHYOLOGY Vol. 43 Suppl. 2 2003
S191
Response intensity,
arb. units
1.0
0.8
0.6
0.4
0.2
1
0.1 2
10 20 40 80
Frequency, Hz
Fig. 26. Spectral sensitivity of free (1) and canal (2) neuro-
masts in Dissostichus mawsoni (based on Montgomery
et al., 1994).
tions differing by 4–15% from the baseline frequency
(Bleckmann et al., 1981). To reduce the hydrodynamic
noises and efficient extraction of the useful signal, fish
developed several adaptations: separation of neuro-
masts from the turbulence zones evoked by the fin func-
tioning (characteristic curves of the body lateral line
canal near the pectoral fins or its shift to the dorsal and
ventral direction), alteration of active swimming with
short-time stops or inertial slipping, temporal termina-
tion of breathing movements, etc.
The discrimination ability of neuromasts is deter-
mined by the sensitivity of the cupula to adequate stim-
uli and its ability to respond to even very weak stimula-
tions. The more the cupula deviates, the stronger the
signal evoked in hair cells. The high sensitivity is
reached first by morphological and anatomical charac-
teristics of the neuromast and its cupula.
It is obvious that the higher the cupula, the weaker
are the effects required to bend it. The depth of the cup-
ula is especially important for free neuromasts. When
the current speed is low, when the current is almost
absent, or during passive transmission of the fish body
within a laminar current, the borderline immobile layer
of water may be quite thick and isolates cupula from the
external signals. Therefore in fish living in such condi-
tions, e.g., Gobius, anglers Neoceratias, Phrynichthys,
and other representatives of Ceratoidea, neuromasts are
situated at the tips of high papillae shifting cupula out-
side of the borderline layer (Fig. 4). In Phrynichthys
wedli with a body length of only 4 cm, each body side
bears 400 free neuromasts, located at papillae with the
height of 5 mm, which is 12.5% of the body length of
these fish! The cupula in these fish is often not cylindri-
cal but compressed, and hair cells are oriented perpen-
dicular to the transverse axis of the neuromast, making
them most sensitive to water shifts with respect to the
body surface. To reduce hydrodynamic disturbances
S192 KASUMYAN
created by the fish, anglers can for a long time stay
immobile except for only rare breathing movements:
they have neutral buoyancy and a low metabolism rate.
Canal neuromasts in anglers are completely reduced
(Denton and Marshall, 1958; Marshall, 1996; Mont-
gomery and Pankhurst, 1997). Neomorphic, the sec-
ondary free neuromasts, which are located far from the
lateral line canals and differing in their orientation, are
especially important for the detection of differently
directed oscillations.
The shift wave is distributed along the lateral line
canal. Therefore, the maximum response and, there-
fore, the maximum sensitivity to the stimulus is pro-
vided by the orientation of the hair cells along the canal
axis. This also depends on such structural characteris-
tics of the canal as its narrowing in places of neuromast
location, internal partitions with small gaps for neuro-
mast cupules (Fig. 19), flattening of the cupula perpen-
dicular with respect to the longitudinal axis of the
canal, overlapping of the entire canal by the cupula, etc.
It has been noted earlier that the neuromasts hidden
in canals are significantly shielded from the outer low-
frequency hydrodynamic noise caused by active swim-
ming of the fish, or due to turbulent or laminar water
current. Filtration of such noises makes possible the
reception of only the useful higher frequency signal by
the neuromasts. Therefore, rapidly swimming fish or
fish living in water currents usually have better devel-
oped canals in the lateral line. Isolation of canal neuro-
masts from the sources of hydrodynamic noises well
explains the characteristic curving of the body canal in
proximity of pectoral fins where the turbulent currents
appear during fish swimming. This also accounts for
the shifting of the lateral line canal to the ventral sur-
face of the body in fish swimming due to the undulation
of the caudal fin (Figs. 7, 8) (Bone, 1971).
Membranes closing the pores of the canals are also
involved in it, which significantly improves the sensi-
tivity of canal neuromasts to more high-frequency stim-
uli, generated by plankton invertebrates. According to
other hypotheses, membranes are involved in resonance
amplification of 5–10 Hz oscillations, significantly
increasing the sensitivity of the fish. One example is
bathypelagic fish Anaplogaster (Anaplogastridae), in
which the wide and shallow canals of the lateral line are
separated by bony partitions and covered by a thin layer
of skin from above (Fig. 19). They have no pores and
cover almost the whole surface of the head. In other
bathypelagic fish, Poromitra (Melamphaidae), the very
wide canals (up to 7 mm in individuals with the length
5 cm) also cover almost the whole surface of the head
(Marshall, 1996). Even though the skin membrane has
pores with diameter of about 0.4 mm, resonance is also
possible in such canals. Computations indicated that the
resonance increases the sensitivity of the lateral line in
Poromitra by a factor of eight (Denton and Gray, 1988).
Another way to increase the sensitivity of the lateral
line to adequate stimuli is associated with the increase
JOURNAL OF ICHTHYOLOGY
of the number of receptor cells in the neuromast.
Because the density of receptor cells differs insignifi-
cantly, this requires the increase of the linear size of the
neuromast and usually is accompanied by the respec-
tive increase of the canal width. For example, the pres-
ence of not only the very large neuromasts and
extremely wide canals in Poromitra provides for the
additional growth of sensitivity of the lateral line by the
factor of 100 (Denton and Gray, 1988). The wide and
well-developed canals of the lateral line are usually
found in inactive fish or fish living in weak current or
no current (Jakubowski, 1967; Marshall, 1996). Func-
tionally, it brings about an increase in the sensitivity to
relatively low-frequency oscillations, as in the fish with
numerous free neuromasts inhabiting stalled or slowly
moving waters (Dijkgraaf, 1963; Marshall, 1971; Mer-
rilees and Crossman, 1973; Coombs et al., 1992).
A unique way to increase the sensitivity of the lat-
eral line is characteristic in herring (Clupeidae) and
Mormyridae, unlike most other fish having no body
canals (Fig. 9). However, the lateral line is well devel-
oped on the head of these fish. A specific characteristic
of these fish is the direct link (recessus lateralis)
between these canals and the labyrinth, the intra-canal
liquid and the perilymph are separated by only a thin
elastic membrane. In turn, the labyrinth in herrings has
a direct contact with paired gas-filled bladders, the out-
growth of the swimbladder connected with it by a thin
duct (precoelomic duct) (Fig. 27) (Denton and Blaxter,
1976). Due to such structural characteristics, the lateral
line in these fish is highly sensitive not only to shift waves,
but to alterations of the volume of the swimbladder caused
by pressure waves from the source of underwater oscilla-
tions, caused, for instance, by small plankton organisms. It
is proposed that the feeding efficiency of these fish signif-
icantly increases in this way.
The presence of complex branchy canal system on
the head is linked with the control of the swimming
speed in fish, i.e. the speed of the incident flow. It is
obvious that in relatively slowly moving fish (e.g., Tra-
churus mediterraneus ponticus and Scomber scom-
brus), the ramification of the head canals of the lateral
line is distributed to the dorsal surface of a part of the
body, i.e., to the turbulence zone. In significantly faster
swimming fishes, e.g., sailfish Istiophorus platypterus,
the formation of turbulence occurs at the rostrum, which in
this species is covered by a dense network of branching
canals of the lateral line (Chubukov et al., 1987).
The multiplicity of body canals characteristic of
several fish species (Bleckmann and Münz, 1990) was
considered as an adaptation directed to increase the dis-
criminative capacity of the lateral line system. How-
ever, detailed studies, conducted on Hexagrammos
decagrammus and H. stelleri, did not support this
hypothesis (Wonsettler and Webb, 1997). It turned out
that out of the five canals on the lateral side of the body
isolated from each other, neuromasts were found only
in one canal and only this canal is connected with the
Vol. 43 Suppl. 2 2003
 THE LATERAL LINE IN FISH: STRUCTURE, FUNCTION, AND ROLE IN BEHAVIOR S193

(a)

(c)
7 8 9

5
1 3
6
4
(b)

10
2
11

Fig. 27. Links between the lateral line, labyrinth and the swimbladder in clupeid fishes. (a) Lateral line canal system at the head of
juvenile Alosa pseudoharengus; (b) labyrinth; (c) links between lateral line canals at the head, labyrinth and the swimbladder.
(1) Membrane, separating the lateral line canal from the perilymphatic cavity of the labyrinth and the location of the membrane on
the head of the fish; (2) gas-filled bladder; (3) endolymph of the utriculus; (4) macula of the etriculus; (5) perilymph of the labyrinth;
(6) links between the cavities of gas-filled bladders and the swimbladder; (7) canal neuromast; (8) cavity of the lateral line canal;
(9) canal pore; (10) rostral area of the swimbladder; (11) membrane, separating perilymph from the cavity of the gas-filled bladder
(based on Denton and Blaxter, 1976 and Janssen et al., 1995).

head canals of the lateral line. The function of the addi-
tional canals is still unclear.
The role of the central regulation in the functioning
of neuromasts and the lateral line system as a whole is
still poorly understood. It was found that the activity of
neuromasts is significantly reduced at the moment of
sharp drops. Such an inhibitory effect may be consid-
ered as an important adaptation protecting the system
from functional overload (Russell, 1974).
The shift wave is a vector value, i.e., the direction of
the water particles shifts is important. The shifts per-
ceived by the lateral line receptors allow the fish to
determine the direction of the source of the oscillations.
This is also enabled by the location of lateral line recep-
tors, which usually form elongated rows or canals: the
rapidly extinguishing shift waves, reaching different
parts of these rows, have different amplitudes, which
make the orientation to the source of the stimulus eas-
ier.
The ability of the fish to determine the location of
the source of the shift waves is demonstrated by the
experiments conducted by G.A. Malyukina on the gold
fish. A conditioned reflex was developed in the fish
involving the location of a generator of low-frequency
oscillations hidden by a shield. Later, a second genera-
tor was placed in the second corner of the aquarium, to
which the positive response of the fish was not rein-
forced by the food. After a prolonged generalization
stage, when the fish positively responded to both posi-
tive and negative stimuli, the fish could differentiate
between them without error (after 70–90 trials), i.e. it
could determine the direction of the source of oscilla-
tions (Malyukina, 1956).
The shift waves have a limited distribution zone
because of the rapid reduction of oscillation amplitude
at a distance from the source. Therefore, the lateral line
as a sensory system is efficient only at small distances.
It was determined that fish respond to immobile objects
at a distance of several millimeters to several centime-
ters or dozens of centimeters depending on the size of
the objects and the amplitude and frequency of the
oscillations. The higher the amplitude and frequency of
the oscillations emitted by the object (of course, within
the adequate ranges), the longer is the distance at which
the fish could detect its presence. It is accepted that the
maximum distance from the source of oscillations at
which the fish could perceive it does not exceed one
body length (Popper and Fay, 1993). It has been shown
experimentally that minnows Phoxinus phoxinus can
easily detect a water current (2.5 cm3/min) released
from a tube with a diameter of 0.7 mm at the distance
of 2–3 cm and an opaque glass string with the diameter
of 0.25 mm at the distance of 10 mm (Dijkgraaf, 1934,
1952, 1963). Baikal gobies Batrachocottus baicalensis
and Paracottus knerii with the length of 80–95 mm
respond to the swimming of nearby amphipods Gme-
lenoides fasciatus with the length of 6.6 mm at the dis-
tance of 7–9 and 3–4 mm (Janssen et al., 1999) (Table 3).
Cottus bairdi in 50% of cases can find live daphnia with

JOURNAL OF ICHTHYOLOGY Vol. 43 Suppl. 2 2003

S194 KASUMYAN

Table 3. The maximum response distances to immobile A blinded muskellunge Esox masquinongy makes a
prey or its acoustic model served by the lateral line directed strike to the prey from 3.2 cm (New et al.,
2001). Most probably, the maximum distance of the lat-
Maximum
Species
distance, cm
Data source eral line is 50–60 cm. It was the distance at which gold
fish could respond to oscillating plates hidden by
Stone sculpin, 0.3–0.4 Janssen et al., 1999 shields (Malyukina, 1955).
Paracottus kneri
The maximum sensitivity and the maximum
Bream, Abramis brama 0.5 Townsend and
Risebrow, 1982 response distance is provided for by the largest neuro-
masts; the spectral characteristics of neuromasts, dif-
Sculpin, Cottus bairdi 0.7 Coombs and Jans- ferent in size and localization, are usually the same
sen, 1989 (Hoekstra and Janssen, 1986; Coombs and Janssen,
Wide-head sculpin, 0.7–0.9 Janssen et al., 1999 1990). Each canal of the lateral line has a sensitivity
Batrachocottus zone, which depends on the length and location of the
baicalensis canal and the size of neuromasts (Fig. 28). To deter-
Chologaster, 2–3 Poulson, 1963 mine the direction of the source of the stimulus and the
Chologaster agassizi distance to it, the fish require information from neuro-
Muskellunge, 3.2 New et al., 2001 masts located in different canals. Blinded Cottus bairdi
Esox masquinongy with an intact lateral line show a strong correlation
Blind fish, 2–4 Poulson, 1963 between the distance to the source of oscillations and
Ambloysis splelaca the length of the drop to the source of the stimulus. If
Cave fish, Typhlichtys 4–5 Poulson, 1963 the body canal is blocked and the head canals remain
subterraneus intact, this correlation is not observed, the distance of
the drop is unrelated to the distance to the target
Pike, Esox lucius 5–10 Wunder, 1927
(Fig. 29) (Janssen and Corcoran, 1998). As hypothe-
Scorpionfish, 30–35 Andriyashev, 1944a sized by some researchers, the determination of the
Scorpaena porcus direction of the source of oscillations by the fish is
Dragonfish, 30–40 Andriyashev, 1944b facilitated by the fact that the lateral line is stretched
Trachinus draco along the longitudinal, rostro–caudal axis of the body
(Sand, 1984).
the length of 2 mm, approaching to 7 mm (Coombs and Fish can determine not only the direction of the
Janssen, 1989). Cave fish of the genus Amblyopsis can source of oscillations and the distance to it, but also
find daphnia at a much greater distance of 2–4 cm respond to alterations in the direction and speed of
(Poulson, 1963). Lepomis macrochirus can detect large movement of the object and discriminate its size and
active prey (juvenile fish) at the distance of 3 cm (Enger shape. For example, in blinded goldfish Carassius
auratus, the frequency of breathing gill movements
et al., 1989). Esox lucius, devoid of eyes, can catch altered if the movement speed of a vertically presented
active prey fish from the distance of 5–10 cm, but can- Plexiglas stick differed by 1 cm/s (5 and 6 cm/s), or if
not hunt if the lateral line is destroyed (Wunder, 1927). sticks with different cross-section areas were presented
(1 × 1 cm2 and 1.4 × 1.4 cm2), or with different shapes
(square, round, or triangular section) (Vogel and Bleck-
55° mann, 2001).
3
160°
NATURE AND SOURCES OF ADEQUATE
2 STIMULI FOR THE LATERAL LINE.
HYDRODYNAMIC CAMOUFLAGE IN FISH
Any movable object in water is a source of acoustic
oscillations, generating both shift and pressure waves.
1 Pressure waves spread in water faster, are extinguished
slower, and create a so-called far acoustic field. Shift
waves have a slower speed, are extinguished much
130° faster, and therefore have a much smaller distribution
zone near the acoustic field. As has been noted earlier,
lateral line receptors are sensitive to relatively low-fre-
Fig. 28. Sensitive zones in various groups of free neuro-
masts in Aplocheilus lineatus: (1) postorbital; (2) supraor- quency shift waves and any other movements of water
bital; (3) nasal groups. Numbers designate the angular size or the fish itself, causing the shifts of water particles
of the sensitive zones (based on Schwartz, 1967). with respect to the fish body surface.

JOURNAL OF ICHTHYOLOGY Vol. 43 Suppl. 2 2003

 THE LATERAL LINE IN FISH: STRUCTURE, FUNCTION, AND ROLE IN BEHAVIOR
2
Drop distance, mm
50 (b)
40
30
20
10
0 10 20 30 40
Fig. 29. Relationship between the drop distance (1) in blind Cottus bairdi and the distance to vibrating target (2): (a) scheme of the
experiment and the measures recorder; (b) responses of the fish with intact body canal of the lateral line; (c) responses of the fish
with blocked body canal. Dashed line shows the initial position occupied by the fish before the drop (based on Janssen and Corcoran,
1998).
The sources of such hydrodynamic stimuli are the
fish themselves, other aquatic animals actively moving
in the water, the bottom substrate or at the surface of
water, water currents or water layers with different
hydrodynamic properties (speed, turbulence). Immo-
bile objects reflecting underwater oscillations or dis-
turbing the hydrodynamic field are secondary sources
of hydrodynamic stimuli. As it has been found, this
allows a fish to find active prey even if it is temporarily
ceases to move and sinks to the bottom (Montgomery
and Milton, 1993).
Small local currents, induced by many aquatic ani-
mals during their movements, e.g., various plankton
organisms or immobile filterers like bivalve mollusks
(Fig. 30), are another class of stimuli for the lateral line.
According to different data, the current speed at the
outflow from the syphon of a mollusk can range from 6
to 14 cm/s (Montgomery et al., 1995). That such cur-
rents can be used by fish to detect the prey follows from
the data about the significant proportion of molluskian
syphons in feeding of many nearshore marine fishes
(Peterson and Quammen, 1982).
The intensity and spectral characteristics of hydro-
dynamic disturbances in water depend on many param-
eters, the speed, trajectory, acceleration of the moving
object, its size and shape, resistance by the water envi-
ronment, the type of the propeller, etc. It was found that
JOURNAL OF ICHTHYOLOGY Vol. 43 Suppl. 2
S195
(a)
1
(c)
50 0 10 20 30 40 50
Distance to the stimulus source, mm
a calmly swimming fish generates oscillations with the
frequency from several Hz to 10 Hz, but sharp drops
and sudden alterations of the movement direction
induce oscillations with the frequency up to 100 Hz
(Bleckmann et al., 1991). Plankton invertebrates gener-
ate oscillations with the frequency from several Hz to
30–40 Hz, which lies in the range of the maximum sen-
sitivity of the fish feeding on them (Fig. 31) (Kirk,
1985; Montgomery and Macdonald, 1987; Montgom-
ery et al., 1988; Montgomery, 1989).
The swimming fish not only is a source of oscilla-
tions of certain frequency, but also releases a hydrody-
namic trace representing a system of turbulent micro
currents. Visualization of these turbulent currents, gen-
erated by a swimming fish or a fish standing in a cur-
rent, allows us to see their real pattern and scale
(Fig. 32). This microturbulence can exist for a rela-
tively long time, from several dozens of seconds to sev-
eral minutes, and spread to a relatively large distance,
up to several meters (Bleckmann, 1993) depending on
the size of the fish and the speed and the pattern of its
swimming. Experiments involving the release of micro
particles with neutral buoyancy into the water and special
laser recording technology revealed that goldfish Caras-
sius auratus (TL 6–10 cm, swimming speed 7–14 cm/s)
freely swimming along straight line trajectories induce
microturbulence, the spatial structure of which is
2003
S196 KASUMYAN

Response intensity, %
(a) 100 (a)

80

60

40

20

(b) 0 20 40 60 80 100
Response intensity, arb. units
(b)

(c)

Fig. 30. Examples of the sources of low-frequency under-
water oscillations: (a) water currents evoked by a filterer
bivalve mollusk; (b) hydrodynamic field, created by fish
moving by inertia; (c) whirl path, evoked by an actively
swimming fish (based on Montgomery et al., 1995).
retained for 30 s. The particles moving with the speed
exceeding the threshold for being detected by the lat-
eral line receptors (0.2 mm/s) can be found in water for
3–5 min (Hanke et al., 2000). Finding oneself in such a
trace, a fish can determine the time passing from the
moment of the whirl formation, the size of the fish that
released the turbulence, the speed and the direction of
its movement, and perhaps even more specific informa-
tion such as the species. It was hypothesized that pred-
atory fish can track down active prey, following it by its
hydrodynamic trace (Hanke et al., 2000).
The hydrodynamic trace makes the fish easily
detectable (Montgomery et al., 1995). Certainly, its
intensity and the duration of existence depend on the
size of the fish and the swimming type: fish with a tor-
pedolike and arrowlike body release more pronounced
hydrodynamic traces. An eellike swimming pattern is
usually accompanied by less intense hydrodynamic
effects. This is why most deep-sea fish and many noc-
turnal fish (e.g., Anguillidae, Siluridae, Muraenidae,
etc.) have elongated bodies and use the eellike swim-
ming pattern allowing them to be successful predators
without being too conspicuous to the prey. This swim-
ming type may be considered as a kind of hydrody-
2
3
0 20 40 60 80 100
Frequency, Hz
Fig. 31. Spectral sensitivity of the lateral line of Pagothenia
borchgrevinki (a) and the frequency–amplitude characteris-
tics of the hydrodynamic disturbances (b) created by zoop-
lankton organisms; (1) Euchaeta antarctica (Calanoida),
(2) Euphausia crystallorophias (Euphausiacea),
(3) Orchomene plebs (Amphipoda). Dashed lines on the
spectrograms show the peak, evoked by the laboratory
equipment (based on Montgomery and Macdonald, 1987).
namic camouflage (Montgomery et al., 1995; Mont-
gomery and Pankhurst, 1997).
There exist other types of hydrodynamic camou-
flage and ways to reduce the hydrodynamic noise
caused by the fish. For example, many species use the
saltation type of search behavior, when random (non-
vectored) movements are altered with short-term stops
allowing them to reduce the level of hydrodynamic dis-
turbances and approach the prey undiscovered. This
tactic is used by the notothenioid fish Trematomus ber-
nacchii (Janssen et al., 1991), Cottus bairdi (Coombs,
1993, cit. in Montgomery et al., 1995), and fish larvae
(O’Brian et al., 1990). Short-term immobility not only
hides the predator, but also allows it to detect signals
with low intensity produced by small food objects. Sev-
eral fish (Hyporhamphus ihi, Aphanopus carbo) use the

JOURNAL OF ICHTHYOLOGY Vol. 43 Suppl. 2 2003

 THE LATERAL LINE IN FISH: STRUCTURE, FUNCTION, AND ROLE IN BEHAVIOR
Fig. 32. Hydrodynamic disturbances evoked by swimming
danio Brachydanio rerio in a media with stratified tempera-
tures. The visualization was based on light diffraction in a
shallow aquarium, vertical temperature gradient 2°C/cm
(based on McCutchen, 1976).
tactic of alteration between short-term rapid move-
ments and gliding by inertia (Montgomery et al., 1995).
Prolonged immobility, characteristic of ambush preda-
tors, can also be considered as a type of hydrodynamic
camouflage. Deep-sea fish even suspend breathing
movements with this aim, which significantly reduces
the level of hydrodynamic noise making it easier to
detect the useful signal (Marshall, 1996).
THE ROLE OF THE LATERAL
LINE IN FISH BEHAVIOR
The lateral line is involved in the feeding, defensive,
schooling, reproductive, and migration behavior in
fishes, allows them to orient in darkness and perform
rheoreaction. When the illumination level is high, other
sense organs are involved along with the lateral line, in
the realization that these behavioral forms may be more
important. In nocturnal fish with or cave or deep-sea
fish, the hierarchy of the sensory systems is different
and the information coming from lateral line receptors
becomes most important.
Feeding Behavior
The ability of the fish to respond to small objects
allows them to determine the presence of the prey,
detect its location with high precision and make a
directed strike. The role played by the lateral line is
clear from the location of the lateral line canals, their
sizes, and the number of neuromasts. For example, the
largest and most conspicuous canals of the lateral line
JOURNAL OF ICHTHYOLOGY Vol. 43 Suppl. 2
S197
Time, min
3
5
Fig. 33. The time course of microwhirls generated by swim-
ming goldfish Carassius auratus (TL 6 cm). Hydrodynamic
disturbances were created by the fish swimming with the
average speed 9.0 cm/s and crossing the recording zone
(upper part of the scheme) for 2.8 s; the width of the record-
ing zone is 27 cm; dark contour line notes the regions, in
which the speed of the water particle movement exceeded
the sensitivity threshold for the lateral line (0.002 cm/s).
The duration of the recording was five min (based on Hanke
et al., 2000).
in many fishes are located on the head, allowing it to
obtain information about the location and movements
of food objects in proximity to the mouth and thereby
catch the prey (Montgomery and Pankhurst, 1997). In
fish feeding on fallen insects, large neuromasts are
located at the dorsal surface of the head (Schwartz,
1967).
The feeding behavior of many predators having a
well-developed lateral line is to a large extent based on
the information provided by the lateral line receptors.
Using the example of the American muskellunge
E. masquinongy, it was shown that visual reception
plays the major role in the detection of prey and the
slow approach to it, whereas the final strike and catch-
ing of the prey is mostly based on the lateral line
(Fig. 34). Intact muskellunges (TL 15–25 cm) struck
the prey (Pimepales promelas, TL 5 cm) at an average
distance of 13.5 cm. After the lateral line receptors were
turned off (the anterior and posterior nerves of the lat-
eral line were cut), the directed strike was observed
from a significantly shorter distance, 10.1 cm. Blind
pikes (visual nerve cut) took the prey only if it acciden-
tally approached closer than 3.2 cm. Fish devoid of
both visual and seismosensory systems did not even try
to catch prey. The loss of any of these two sensory sys-
tems required more precise orientation to the prey
2003
S198 KASUMYAN

(a)

1
ε, α

(b)

2 3

Fig. 34. Measures used in a study of the responses of the muskellunge Esox masquinongy to active prey: coordinates of the prey (a)
and the locomotor response of the predator (b). ε, elevation, α, azimuth, (1) distance to the prey, (2) ambush distance, (3) drop dis-
tance (based on New et al., 2001).

before the final strike: in fish with the visual nerve or
lateral line nerves cut, the angle between the direction
to the prey and body position was mush larger than in
intact individuals (New et al., 2001).
The ability of blind fish to hunt successfully is char-
acteristic of the pike E. lucius (Hofer, 1908; Disler,
1967). Such fish as the burbot Lota lota, catfish Silurus
glanis, eel Anguilla anguilla, Corvina nigra, Gaidrop-
sarus mediterraneus, and many other nocturnal or twi-
light fishes feed much more efficiently in darkness than
in good illumination (Pavlov, 1959, 1962, 1963; Girsa,
1962, 1981). The bluegill sunfish Lepomis macrochirus
actively feeds in darkness, however, after exclusion of
Swimming speed, mm/s
1 more successful are the fish in their behavioral
2 responses. Comparisons of night feeding in the ruffe
25
15
5
1 2 3 4 5 6
Response distance, mm
Fig. 35. Relationships between the response distance in
darkness in the common ruffe Gymnocephalus cernuus
(TL 5.6–6.2 cm) (1) and the yellow perch Perca flavescens
(TL 5.3–5.8 cm) (2) to live daphnias (length 3.0–3.5 mm) and
the swimming speed of the fish (based on Janssen, 1997).
the lateral line, it catches the prey goldfish only after
accidental immediate contact with it. The deprived
predator cannot find the prey even if it is only 0.2 cm
away (Enger et al., 1989). Juvenile ruffe G. cernuus can
feed on large plankton organisms even in darkness
(Smirnov, 1969). The Chinese perch Siniperca chuatsi
(TL 10–12 cm) can equally feed on Pseudorasbora
parva (TL 3.5–4.0 cm) when both its vision and lateral
line are intact and if any of these sense organs are
excluded (Liang et al., 1998). Lepomis cyanellus and
Micropterus salmoides trained to take food from a pipe
actually catch the water current rather than individual
visual stimulus of the prey, which is also an example of
the lateral line dominance over the visual system (Jan-
ssen and Corcoran, 1993).
The better the development of the lateral line, the
better developed are neuromasts and canals and the
G. cernuus and perch P. flavescens indicated that the
former species is more successful in feeding on live
daphnia. It not only can detect the prey from a greater
distance than perch, but also during the movement by
inertia, when the fish does not move fins; in contrast,
perch could catch daphnias only when immobile
(Fig. 35). The more efficient lateral line in the ruffe
depends on structural characteristics of the lateral line:
larger canal neuromasts, and elastic membranes closing
the canals (Jakubowski, 1967; Janssen, 1997). With
respect to the development of lateral line canals, the
ruffe significantly differs from many other representa-
tives of perches and is more similar to inhabitants of the
bathypelagial zone. It was hypothesized that mem-
branes sort out the low frequency hydrodynamic noise
evoked when the fish moves forward by inertia: they
remain concave, but the intra-canal liquid does not

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 THE LATERAL LINE IN FISH: STRUCTURE, FUNCTION, AND ROLE IN BEHAVIOR
(a)
20° 0°
40°
60°
5 mm
Fig. 36. Response zones, served by different regions of the lateral line (a) and repositioning (b) in the sculpin Cottus bairdi: the
dashed line limits the area, where the prey is captured without prior preorientation (Hoekstra and Janssen, 1986).
move. It is this time when the fish can successfully
detect the oscillations created by daphnias. According
to another hypothesis, the membrane may be used as a
resonator amplifying the signal from the prey (Denton
and Gray, 1988, 1989).
Due to the lateral line, many fish, most importantly,
pelagic planktivores such as clupeids, certain cyprinids,
and juveniles of many species, can retain the ability to
feed in complete darkness or if their visual reception is
excluded, and catch individual plankton animal (Jans-
sen et al., 1995). Blocking of lateral line receptors with
streptomycin causes the blind fish to loose this ability
(Blaxter and Hoss, 1981). Experiments on the sculpin
Cottus bairdi feeding at night indicated that the main
role in it is played by canal neuromasts. Blinded fish
(body length 7.5 cm) responded by alteration of their
orientation and directed strikes at live daphnia 2 mm in
size or bead of 3 mm, generating oscillations with the
same frequency as the swimming daphnia 3 Hz (Kirk,
1985). The response distance reaches 5 mm when the
largest neuromasts are stimulated at the lowest side sur-
face of the head (mandibular canal) and at the gill cov-
ers (opercular canal). When the less sensitive sector of
the lateral line was stimulated, the fish changed its posi-
tion in such a way that the source of the stimulus was in
a more sensitive zone and struck only after this
(Fig. 36) (Hoekstra and Janssen, 1986).
Repositioning, i.e., such alteration of the body posi-
tion that the source of oscillations was in the most sen-
sitive zone, is very important for precise orientation and
capture of the prey. In most cases (83%), unsuccessful
final drops of the sculpin were observed if the source of
oscillations was located immediately in front of the fish
(sector ±20°). When the sculpin moved in short darts to
the source of oscillations, its location in the most sensi-
tive place at the side was retained. Whenever the fish
turned one side to the source of oscillations all the time,
the movement trajectory represents an arch. Fish can
JOURNAL OF ICHTHYOLOGY Vol. 43 Suppl. 2
S199
(b)
Prey position
original after repositioning
5 mm
approach the source perceiving it with left or right side
receptored in an alternating manner (Coombs and Con-
ley, 1997).
To discover prey in darkness, pelagic planktivores
often use the so-called saltation search when periods of
active swimming are interchanged with short-term
stops. The prey is detected in the moment when the fish
is immobile and the ratio between the hydrodynamic
noise and the useful signal is the most optimal. The dis-
tance of the prey detection and directed drop usually
does not exceed one body length of the predator
(Norton, 1991; Norton and Brainerd, 1993).
Fish living in stagnant or slowly moving water envi-
ronments are relatively inactive. Many deep-sea and
bathypelagic fish use free neuromasts to detect active
prey or approaching predators. This is possible due to
several structural and physiological adaptations reduc-
ing the level of low-frequency hydrodynamic noises
affecting the receptors. These adaptations include high
laterally compressed cupules of neuromasts, the large
size of neuromasts, dermal tubercles and papillae pos-
iting neuromasts outside the immobile borderline layer,
the inactivity of the fish and low level of their metabo-
lism. Many deep-sea fish have very wide and shallow
canals of the lateral line on the head, which in the func-
tional respect (spectral sensitivity) are not very differ-
ent from free neuromasts. Many deep sea fish have an
eellike body and swimming pattern causing a much
lower level of oscillations in the environment. This
reduces the unfavorable hydrodynamic noises and
makes the fish less conspicuous to predators and prey.
It was recently found that the lateral line, along with
the auditory reception, is involved in the search of
active objects not only in water, but also in the bottom
substrate. Sculpins C. bairdi can easily find a 10 Hz
vibration, imitating the moving ground benthos organ-
isms (Nereis and Lumbriculus). The response to the
vibration is observed only if the fish is placed on the
2003
S200 KASUMYAN

(a) (b)
Target object

β α

Vibrator 2
4

8 5 3
6
9 7
10 cm

Fig. 37. Discovery of a source of underwater oscillations (10 Hz) imitating an active food object and hidden in the bottom substrate
by the sculpin Cottus bairdi. (a) Target linked with a vibrator and hidden in sand ground at the depth 4 mm from the surface;
(b) search movements of the sculpin. Silhouettes (1–9) point the subsequent alterations of the position and orientation of the fish,
the angles between the direction of the initial orientation of the fish (before the stimulation), and the direction to the source of oscil-
lations (α) and the direction of the fish after pre-orientation after the first drop (β) (Janssen, 1990).

bottom, the fish touches the surface of the substrate
with its lower jaw, and holds its breathing movements
for more than 1 s. If it receives the signal in this
moment, it rapidly turns to the source of the stimulus.
A series of short darts towards the vibration occurs after
this. It is very important that the fish touches the sub-
strate with its lower jaw after each dart. The response
terminates by a precise directed strike at the ground in
the place of the oscillation source. The fish can respond
at the distance up to 30 cm, i.e., four–five body lengths.
If the vibration is not contacting the ground, the
response distance is lower, 1–2 cm. The treatment of
mandibular neuromasts at the lower surface of the
lower jaw in large fish with streptomycin or cobalt salt
brings about partial or complete disturbance of the
response: the fish touched the substrate with the lower
jaw but did not dart toward the oscillation source
(Fig. 37) (Janssen, 1990). Most probably, many bottom
fish feeding on infauna can use the lateral line together
with the auditory reception to catch hidden prey objects
from the bottom substrate.
In fish specializing on feeding on fallen insects, the
lateral line provides for not only the information about
the prey, but also allows the fish to determine the dis-
tance to it. The mechanism of the precise detection is
associated with the distribution of the surface waves
evoked by the insect and the ability of the fish to per-
ceive and analyze these oscillations. There are several
important differences between the surface waves and
the waves distributed in water. In particular, the low-
frequency components of surface waves distribute at
higher speeds than the high-frequency, whereas the
reverse is characteristic of shift waves. On the whole,
the speed of surface waves is more than 1000 times
lower than waves distributing in the water column.
They hardly penetrate into the water column, therefore
the fish specializing feeding on fallen insects usually
stay at the surface.
The detection of the surface waves is based on canal
and free neuromasts located on the head and is well
developed in such fish as Pantodon buchholzi (Pant-
odontidae), Galaxias fasciatus (Galaxiidae, Osmeri-
formes), Fundulus notatus (Cyprinodontidae) or
Aplocheilus lineatus (Cyprinodontidae) (Schwartz,
1965; Hoin-Radkovsky et al., 1984). The direction of
the prey is determined due to the difference in the time
and the amplitude of the waves coming to symmetrical
neuromasts on both sides of the fish head (Schwartz,
1965; Schwartz and Hasler, 1966). The distance to the
source is determined due to the well-developed ability
of these fish to analyze the spectral composition of sur-
face waves coming from the prey and its dynamics. An
insect fluttering in water generates oscillations in a
wide frequency range, much wider than the baseline
oscillations of the water surface evoked by the wind or
the fish itself (Fig. 38). But after a short time after the
signal dispersed a small distance, its spectrum changed
because of the much faster attenuation of high-fre-
quency components and their slower distribution. The
oscillation amplitude significantly reduces with the dis-
tance from the source (Fig. 39).
Analyzing the frequency, time, and amplitude
parameters of the signal reaching the receptors on both
sides of the body, fish can easily determine the distance
to the prey. The deviation error in the drop is usually
about ±5° (Bleckmann et al., 1989). The response dis-
tance in fish reaches 50–60 cm in Fundulus and up to
1 m in Aplocheilus. The response latency is usually
only 0.038 s (Schwartz, 1967). It is quite enough if the
A. lineatus detects only the first eight–ten wave oscilla-
tions to localize the source (Bleckmann and Schwartz,
1981). If the fish are experimentally presented with a
signal, similar in its spectral and frequency parameters

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 THE LATERAL LINE IN FISH: STRUCTURE, FUNCTION, AND ROLE IN BEHAVIOR S201

Amplitude, dB
(a) 0

–30

–60

(b) 0

–30

–60

(c) 0

–30

–60
500 µs 0 30 60
Frequency, Hz

Fig. 38. Amplitude, dynamic, and frequency parameters of surface waves, generated by fallen at water surface flying insects (a),
fish (b), and wind (c) (based on Beckmann et al., 1989).

to the signal from a larger distance, the fish make an Schooling Behavior
error and perform a more distant drop (Bleckmann,
1993). Fish retain the capability to orient and catch There is not much data pointing to the possible
fallen insects even if they lack almost all neuromasts involvement of the lateral line organs in schooling
(Bleckmann and Schwartz, 1982), but loose this ability
if the artificial source generates a monofrequency rather (a)
4 cm 8 cm 16 cm
200 mV

than a polyfrequency signal (Bleckmann, 1980). 100 mV

1V

Defensive Behavior
20 µs
Large active objects evoke relatively strong, high-
amplitude, and low-frequency oscillations, which can
distribute far in water and can be easily perceived by
lateral line receptors, primarily, by free neuromasts.
Large fish are usually the sources of such signals. The
behavioral response to such stimulus expressed in
active escape or hiding, allowing to avoid predators, Frequency, Hz
especially at night and in twilight hours, when the dan- (b)
ger cannot be so easily detected with other sense 160
organs. The defensive response to high-amplitude, low- 120
frequency oscillations is innate and appears early in the
fish ontogeny, after the first appearance of free neuro- 80
masts in larvae before the beginning of exogenous feed- 40
ing or at the beginning of the larval period (Disler,
0
~
~
~
~

~
~

1960). The involvement of the lateral line in the realiza- 3 5 7 9 16 19

tion of the active escape response is supported by the Distance, cm
links between the Mauthner cells (involved in the active Fig. 39. Amplitude, dynamic (a) and frequency (b) parameters
escape response) and lateral line nerves (Zottoni and of the surface wave, recorded at different distance from the
Danielson, 1989). source of oscillations (based on Montgomery et al., 1995).

JOURNAL OF ICHTHYOLOGY Vol. 43 Suppl. 2 2003

S202 KASUMYAN
behavior in fish. For example, in observations of a blind
Anchoviella individual, it always joined the school and
repeated all schooling maneuvers (Moulton, 1960). In
experiments with the tuna Euthynnus affinis, the tested
fish separated by a transparent partition were located
with respect to each other differently than when they
were in a single school, pointing to the possible role of
the lateral line in the within-school pattern (Cahn,
1972). Blinded individuals of pollack Pollachius virens
retained the within-school contacts but lost them after
the lateral line nerve was cut (Pitcher et al., 1976; Par-
tridge and Pitcher, 1980). Unfortunately, many of these
experiments did not control well the completeness of
the deprivation during the whole experiment. The
attempts to associate the transition to schooling with
massive formation of neuromasts in juvenile fish (Blax-
ter, 1987) are not convincing.
The examples indicating that the lateral line is not
important in schooling behavior of fish are much more
abundant (Pavlov and Kasumyan, 2000), when the
retention of the lateral line but exclusion of vision
caused the complete destruction of the school and com-
plete loss of schooling behavior. It is most likely that
the lateral line plays only a secondary role in the real-
ization of schooling behavior and provides for precise
coordination during rapid maneuvers, along with
vision, when the distance between schooling partners is
very short (Gray and Denton, 1991). However, as is
supported by many authors, even in compact schools
the involvement of the lateral line seems not completely
convincing because, for example, herrings (Clupeidae)
with very well-developed schooling behavior, have no
lateral line canals on the sides of the body (Sand, 1984;
Popper and Platt, 1993). It was hypothesized (Chu-
bukov et al., 1987) that in Clupeidae, Carangidae,
Scombridae, and Istiophoridae, the complex network of
canals on the head is primarily used for mutual control
of the swimming speed in the school.
Spawning and Parental Behavior
The complex organization of spawning behavior, char-
acteristic of many fishes, is often accompanied by dis-
plays, shifts, dances, vibrations, and other specific behav-
ioral patterns. Obviously, the low-frequency oscillations
may bear very important information and can help in the
synchronization of spawning behavior and spawning. The
efficiency of such signalization depends on a small dis-
tance between the partners, usually less than one body
length, i.e., in the near acoustic field.
The involvement of the lateral line in spawning
behavior has long bee only hypothesized (Tinbergen,
1951; Keenleyside, 1979). Experimental data support-
ing this have been obtained only recently in sockeye
salmon Oncorchynchus nerka (Satou et al., 1994a,
1994b). The spawning behavior of males and females
of these fish represents an array of displays: sharp
bends of the body of the female in the nest make the
male somewhat downstream move rapidly towards the
JOURNAL OF ICHTHYOLOGY
female and a characteristic quivering (vibration) causing,
in turn, a quivering response of the female, signal its readi-
ness for spawning. In response, the male approaches the
female, quivers, opens its mouth, and releases the sperm.
This releases the female’s response: quivering and release
of eggs. During the body bending and quivering, the fish
cause oscillations with the frequency from 2 to 37 Hz, and
the myograms in males and females at various stages of
spawning behavior are different.
To analyze the role of the low-frequency oscillations
in spawning behavior of these fish, the responses of
dwarf sockeye salmon males to plastic models of a ripe
female (the length of the model 24 cm, depth 6 cm, and
width 4 cm) were observed. The model, vibrating
in the vertical hyperplane with the frequency 2–32 Hz
and the amplitude 1.7–2.5 mm brought about all the
spawning repertoire including the release of the sperm
in males located in a basin with a gravel substrate
(142 × 42 × 45 cm, current speed 5–6 cm/s). Depending
on the frequency of quivering, males performed from
2–4 to 7–10 spawning cycles during two minutes of the
experiment. Characteristics of the model coloration
(black or white or models with black upper or lower
parts) did not affect the behavior of the male signifi-
cantly. The efficiency of an opaque quivering model
was significantly weaker. The spawning behavior was
not observed in the absence of quivering, when the
vibrating model was shielded by a glass partition, or if
visual and hydrodynamic models were separated from
each other. Interestingly, the dart and quivering of the
male were best stimulated by the model oscillations
with the frequency 2 Hz, whereas opening the mouth
and sperm release, with the frequency of 5 and 21 Hz,
which agreed with the frequency of the quivering per-
formed by female sat the respective stages of the
spawning behavior (Fig. 40).
Signals perceived with the lateral line receptors may
be very important in parental behavior of certain fishes.
Males of the fighting fish Betta splendens use such sig-
nals to rapidly attract the juvenile offspring swimming
near the surface at a distance from the nest in case of
danger. In such cases, the male takes an oblique pos-
ture, touches the water surface, and makes fluttering
movements with its pectoral fins. This evokes a surface
wave, which can be perceived by the young at a dis-
tance up to 40 cm, and which rapidly causes directed
movement to the source of the oscillations. The
response is observed both in light and in darkness. It
can be easily released in response to an artificial signal
with the optimal frequency 8–10 Hz and amplitude,
13 µm. The efficiency of such signalization is quite
high, about 70% of the brood approaches the nest after
1–2 min (Bleckmann, 1993).
Orientation in Space
and the Response to the Current
Many fish species experience water currents. The
ability to determine its presence, strength, direction,
Vol. 43 Suppl. 2 2003
 THE LATERAL LINE IN FISH: STRUCTURE, FUNCTION, AND ROLE IN BEHAVIOR S203

(‡) Number of fish expressing rheoreaction

4.0 (a)
1
2
Number of spawning attempt by male 3.5
* 3
10 (b)
* 3.0
6
2.5

2 ***
*** *** 2.0
0 2 4 6 8 10 12 14 16
1 2 3 4 5 6 7 (b)
Type of the artificial female model 4.0 4
Fig. 40. Responses of male sockeye salmon Oncorhynchus 3.5
nerka to artificial hydrodynamic models of female:
(a) spawning nest and female model, the arrows point to the
direction of the model vibrations; (b) the number of spawn- 3.0
ing attempts of the male (vibration, opening the mouth, and
releasing a portion of sperm), caused by different female 2.5
models. 1, 2, 3, and 4, black and white models with the
vibration frequency 2, 5, 21, and 37 Hz; 5, immobile black
and white model; 6, transparent model with the vibration 0.5 1.0 1.5 2.0 2.5 3.0
frequency 21 Hz; 7, black and white model with the vibra- Current speed, cm/s
tion frequency 21 Hz, located within a shield. * and ** points
to the significance of the differences with respect to the Fig. 41. Sensitivity of the blind fish Astyanax fasciatus to
male response to the model 1, respectively, p < 0.05 and p < the water current. (a) The number of fish exhibiting rheore-
0.001 (based on Satou et al., 1994b). action: intact (1), with canal neuromasts blocked by gen-
tamicin (2), with the surface neuromasts removed (3);
(b) the number of intact fish, exhibiting rheoreaction in
and the structure is extremely important for the survival presence of the food extract (4) (based on Baker and Mont-
and most behavioral forms: defensive, territorial, gomery, 1999b).
spawning, schooling, migration, etc. (Pavlov, 1979,
1980, 1986). This ability is especially important for the (a) (b)
species living under conditions of turbulence or in dark-
ness, or who are active at night. Orientation against the
current is necessary to the fish feeding on drift because
it allows them to obtain olfactory information about the
approaching prey, for energetically more efficient resis-
tance to the current, for migrations or redistribution
within the same habitat, as in nearshore marine fishes of
the tidal zone, and to retain their position within the
(c)
current and so on.
The perception of the current by the fish is possible
due to free neuromasts, the cupules of which are
directly affected by the pressure of the water passing
along the fish body (Engelmann et al., 2000). Removal
of free neuromasts causes the same increase of the fish
response threshold to the current as the simultaneous
removal of all types of neuromasts (Cheimarrichthys
fosteri, Pagothaenia borchgrevinki, blind form of Asty-
anax fasciatus). Blocking of only intra-canal neuro-
masts does not cause any significant deviations in the
fish sensitivity to the current. When food scent appears
in water, the sensitivity of the fish to the water current Fig. 42. Scheme of the hydrodynamic field, generated by
significantly increases (Montgomery et al., 1997; the fish swimming by inertia (a), disturbance of the field by
Baker and Montgomery, 1999a, 1999b) (Fig. 41). an immobile object (b), and the scheme of the hydrody-
namic gradient at the surface of the fish body when it locates
Removal of free neuromasts significantly interferes in proximity to an immobile object (c) (based on Montgom-
with the rheoreaction in marine cartilaginous fishes, as ery et al., 2001).

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S204 KASUMYAN

(a) (b)

(c) (d)

Fig. 43. The development of the peripheral network of the anterior nerve of the lateral line in the ontogeny of Eigenmmania sp.:
(a) juveniles at the age three days with the length 5.5 mm, neuromasts are absent on the head; (b) juveniles at the age four days with
the length 6.25 mm, there are 17 neuromasts on the head; (c) juveniles at the age 5 days with the length 6.5 mm, there are 23 neu-
romasts on the head; (d) juveniles at the age seven days with the length 7.0 mm, there are 35 neuromasts on the head. The shadowed
zone indicates the location of the fluorescent stain injection, which spread along nervous fibers. Arrows point to the branches of the
anterior nerve of the lateral line, simultaneously enervating neuromasts and electroreceptor organs (based on Vischer et al., 1989).

has been shown in the shark Heterodontus portusjack-
soni (Peach, 2001).
Due to the lateral line, fish can orient in the environ-
ment in complete darkness and detect obstacles. It has
been hypothesized first by Dijkgraaf (1963), and exper-
imentally corroborated by Campenhausen et al. (1981
etc.) in cave fish. Cave fish constantly perform hydro-
dynamic monitoring of their environment, specific
hydrodynamic location, comparable with the echoloca-
tion of bats (Chiroptera) and Noctuidae butterflies
(Lepidoptera) or the active electrolocation of weakly
electric fishes (Lapshin et al., 1993; Lapshin, 1995;
Moller, 1995). As a result of this location, the fish cre-
ate a three-dimensional “hydrodynamic representation”
or “hydrodynamic image” of the environment (Hassan,
1989). And this makes their orientation in darkness
possible. Moving forward by inertia, i.e., without their
own active movements, the fish induces wave oscilla-
tions and creates a hydrodynamic field, which is dis-
turbed by any objects in proximity reflecting these
waves (Fig. 42). The pattern of the field disturbance
depends on the size and shape of the object, the speed
of its movement, the pattern of swimming, etc.
Observations of blind Mexican fish Astyanax fascia-
tus (Characidae) indicated that when the fish locate the
space and find a novel object, it uses characteristic tac-
tics: approaching the object, they swim in proximity to
it by inertia. Such behavior could be repeated several
times, the fish changes the direction approaching the
object from a different side. If the fish passes above the
object, it shifts from the vertical axis so that the side
surface of the body was directed to the novel object.
The cave fish could easily discover cylindrical objects
with a diameter up to 2 mm at the distance of several
mm in water, which was determined using the condi-
tioned reflex methods. The fish could discriminate grids
if the distance between the vertical partitions differs by
12.5% towards its increase or decrease, e.g., grids with
the distance between the partitions 10 mm from grids
with the step 8.75 or 11.25 mm (Campenhausen et al.,
1981; Weissert and Campenhausen, 1981).
If the task is more complicated and the fish are
transferred into a completely new environment rather
than presented with novel objects, exploratory behavior
looks different. In this case it involves more rapid
swimming during the first days (Teyke, 1985). Smaller
individuals swim with relatively higher speed than
large, which is probably associated with the need to
generate more high-amplitude shift waves, i.e., for the
formation of a larger near acoustic field (Teyke, 1988).
Blocking the lateral line receptors with salts of heavy
metals brings about the compensatory increase of the
fish swimming speed. That is, the loss of sensitivity
causes the tendency to generate a stronger signal. The
hydrodynamic image of the environment (cognitive
map) is retained in the fish memory at least for several

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 THE LATERAL LINE IN FISH: STRUCTURE, FUNCTION, AND ROLE IN BEHAVIOR S205

(a)
1

(b)

(c)

Fig. 44. The appearance of free neuromasts in early ontogeny of fish: (a) Paralichthys olivaceus, (1) newly hatched embryos,
(2), (3), (4), (5) after 9, 21, 30 h and 9 days after hatching (Kawamura and Ishida, 1985); (b) one-day old juveniles of Lateolabrax
japonicus; (c) one-day old juveniles of the goldfish Carassius auratus (based on Iwai, 1980).

days, because individuals returning after two days to a
familiar environment spent much less time on explor-
atory behavior than if placed in a novel environment
(Teyke, 1989). Hydrodynamic location and the orienta-
tion of cave fish involves primarily the canal neuro-
masts (Abdel-Latif et al., 1990).
The use of the lateral line to avoid encounters with
obstacles in darkness was experimentally shown in
other fish species: intact dace Leuciscus leuciscus with
the length of 4–5 cm could successfully avoid encoun-
ters with a plate placed into the water current with the
speed gradient from 0.3 to 2.0 cm/s/cm at the distance
not more than 20 cm (Pavlov and Tyuryukov, 1993,
1995). The reception of oscillations, reflected by large
underwater objects, allows fish to use these objects as
stationary landmarks, which is very important for terri-
torial fishes, such as juvenile salmons, retaining their
position not only in the light but also in the dark part of
the day, and successfully resist the current. After the
nerves enervating neuromasts on one side of the body,
the fish can retain orientation with respect to such
objects due to receptors on the intact side (Sutterlin and
Waddy, 1975).
ONTOGENY OF THE LATERAL LINE
The first neuromasts appear in embryos under the
shell, as for example, in carp Cyprinus carpio and many
other fish species (Disler, 1960; Iwai, 1967; Appelbaum
and Riehl, 1997). The formation of neuromasts is

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S206 KASUMYAN

Neuromast length, µm larvae of herring and cod, its growth is 1.4–4.1 µm/h
140 (a) (Blaxter, 1984, 1987). Due to the proliferation ability,
Canal neuromasts free neuromasts appear during the whole life of the fish
120 or at least during a significant period rather than in only
100 a limited period of early ontogeny (Disler, 1960;
O’Connel, 1981). The rate of new neuromast formation
80 differs in fish with a different mode of life, especially
60 Free neuromasts the hydrodynamic conditions of further development.
For example, in fish laying eggs on water vegetation
40 (roach Rutilus rutilus, bream Abramis brama,
20 verkhovka Leucaspius delineatus) free neuromasts
develop faster and the recently emerged embryo have
0 almost two times more neuromasts per unit area of the
34 36 38 40 42 44 46 48
Body length, mm body than in embryos of fish spawning in mountain riv-
Number of neuromasts
250 ers with a stony bottom (Diptychus maculatus,
(b) D. dybowskii, Schizothorax pseudoksaiensis issikkuli,
Leuciscus schmidtii (Disler, 1960). The sizes of free
200 At the body neuromasts in juveniles of older age groups differ less
(Fig. 45) (Wonsettler and Webb, 1997).
150
The formation of lateral line canals occurs signifi-
cantly later, but the vector is the same: head canals are
100 the first to appear, and only later, the body canal of the
At the head
lateral line. The canals development is induced by neuro-
50 masts: separate free neuromasts first appear in places of
the canal formation. Then, their number increases, they
0 form even rows and continuously go down to the epider-
5 10 15 20 25 30 35 mal groove. The roof of the canal is developed later. It
Body length, mm appears in the body canal in its anterior part and then dis-
tributes in the caudal direction. The relative sizes of the
Fig. 45. Alterations in the size of free and canal neuromasts pores, leading to the canal are reduced. In larger individ-
in Hexagrammos stelleri (a) (based on Wonsettler and uals, they could rise above the epithelium surface as
Webb, 1997) and the number of neuromasts on the head and short tubes.
the body (b) in juveniles of Carassius carassius as it grows
(based on Disler, 1960). The canal neuromasts become larger with the
induced by lateral line nerves, which develop from gan- growth of the fish (Fig. 45). The number of receptor
glion cells originating from the respective placods. The cells in them increases, but the number of canal neuro-
growth of nerve fibers and their branching rapidly masts after canal formation does not change. In juveniles
makes the peripheral nervous network of the lateral line of Paralichthys olivaceus (Kawamura and Ishida, 1985),
more and more complicated (Fig. 43). The first neuro- the first to develop is the mandibular canal, on the 25th
masts appear where the branching nervous fiber grows day after hatching (at the temperature 16.2–20.7°C) All
down to the epithelium. canals are formed at the head to the 53th day. The for-
mation of the body canal occurs by the 67th day. In flat-
Free neuromasts are the first to appear. In many fish fishes, the development of the lateral line canals on both
species (Lateolabrax japonicus, Acanthopagrus sides of the body is asynchronous—rapidly on the eye
schlegeli, Kareius bicoloratus, Tridentifer obscurus, side and with a delay on the blind side (Harvey et al.,
etc.) they are conspicuous on the head even in just 1992). In Latimeria chalumnae, lateral line canals are
emerged embryos, regardless of their small size and formed extremely early, in embryos with a yolk sac, the
rarity (Fig. 44). In other fish (Hypomesus transpacifi- number of pores on the head approaches 300, in early
cus, Tribolodon hakonensis, Zacco platypus, Oryzias juveniles, 500, and in mature individuals, almost 3000
latipes, etc.) neuromasts become easily visible due to (Hensel and Balon, 2001).
the cupula about one day after emergence (Iwai, 1972).
The size and number of free neuromasts increases rap- Almost immediately after the appearance of the first
idly and already several days or even hours after emer- neuromasts, the lateral line becomes active and func-
gence they can be found on the body, first in its anterior tional. Early juveniles of fish and even prolarvae (e.g.,
part, and then in the caudal part, and finally homoge- keta O. keta) can respond to stimuli adequate for the lat-
neously cover the body of the larvae (Figs. 44 and 45) eral line, such as currents and large underwater objects.
(Disler, 1960; Iwai, 1967; Vischer et al., 1989). Very soon, later larvae and early fry exhibit responses
The depth of the cupula at the moment of neuromast to weak hydrodynamic signals evoked by smaller active
formation rapidly increases. As it was determined in objects like food organisms (Disler, 1960).

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 THE LATERAL LINE IN FISH: STRUCTURE, FUNCTION, AND ROLE IN BEHAVIOR
EFFECTS OF CONTAMINATION
As in most other sensory systems, lateral line recep-
tors are very susceptible to the effects of contaminating
agents. Heavy metal ions exert the strongest effects on
the function of the lateral line. For example, 24 h of
exposure in a solution of a Cd+2 with the concentration
of 450 µg/l causes complete elimination of the electric
response to adequate stimulation of nervous fibers in
Pimephales promelas (Yan, 1995; cit. in Bake and
Montgomery, 2001). Heavy metals disturb the ability of
the fish to orient in the water current: the reduction of
the rheoreaction is observed in Galaxias fasciatus
already after one day of exposure in water with a con-
centration of cobalt ions 2 µg/l. After returning to clear
water, the normal rheoreaction can be retained, but this
requires about 14 days (Baker and Montgomery, 2001).
Heavy metals are often used to exclude lateral line
receptors temporarily. Certain other preparations, like
streptomycin sulfate antibiotic can also be used for this
aim (it blocks both free and canal neuromasts) (Kaus,
1987, cit. in Baker and Montgomery, 2001).
CONCLUSIONS
This review includes only some materials from the
numerous data on the morphology, function, and the
role of the lateral line in fish behavior. The studies in
this area are very intense. They are often conducted
using the most recent methods and approaches, includ-
ing different electrophysiological apparatuses, elec-
tronic microscopy, immunohistochemistry, etc. They
allow us to obtain a large amount of data on the charac-
teristics and structural organization of the peripheral
lateral line and the main functional characteristics of
this sensory system. The involvement of the lateral line
in several behavior forms was revealed.
However, many questions are still poorly studied.
This is especially true for such structures as the vesicles
of Savi and the spiracular organ. Their functions are
still unknown. Very little is known about the internal
structure of the cupula, its mechanical properties,
development and the regeneration speed, its chemical
composition, and about how the cupula is attached to
the neuromast base. The ultrastructure of hair cells and
the innervation of neuromasts in various groups of fish
are weakly studied. Certain functional characteristics
of the lateral line, the structure of its central regions,
and especially characteristics of information process-
ing and the efferent organization of neuromasts are
inadequately known. The role of the lateral line in
schooling and certain other kinds of behavior, interac-
tion between the lateral line and other sensory modali-
ties, their complimentarity are also poorly understood.
The pattern of lateral line development during the
ontogeny is not well known. The evolution of the lateral
line organs and especially the pattern of the evolution-
ary transformations, that caused the development of
such related systems as the auditory and vestibular is
JOURNAL OF ICHTHYOLOGY Vol. 43 Suppl. 2 2003
S207
far from being completely understood. Further studies
are necessary on the vulnerability of lateral line organs
to various types of pollutants and strong artificial seis-
mic factors, such as underwater explosions, used in oil
and other geological prospecting on the marine shelf.
ACKNOWLEDGMENTS
I am very grateful to S.A. Smirnov and
E.A. Marusov for many valuable comments on the
manuscript. I am also grateful to J. Janssen, S. Coombs,
M. Jakubowski, J. Montgomery, N.J. Marschal,
R.G. Northcutt, H. Bleckmann, and S.M. Van Netten
for copies of their papers, which were very valuable
during the preparation of this paper. This study was
supported by the Program “Leading Scientific Schools”
(Project 00-15-99769) and the program “Universities
of Russia” (topic UR.07.03.011).
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Translated by S.V. Budaev
2003