Chapter 4

Mechanical Stimuli

4.1 Mechanical Stimuli

Mechanoreceptors respond to physical movement in the receptor. This can result
from water currents, sound, body movements, pressure or gravity. Each of these
types of stimulation is available in water and may be involved in migration and
orientation.
Fish mechanoreception is relatively difficult to investigate. The responses of
several different types of receptors seem to overlap. Mechanical stimuli are hard
to record without interfering with the stimulus itself, particularly stimuli from
touch or small turbulent water movements. Unlike vision and olfaction, it is rel-
atively difficult to neutralize the mechanical senses experimentally. Touch and the
lateral line sense are widespread on the body and would require nerve cutting,
which is more difficult than stuffing cotton into nostrils to eliminate olfaction, for
example. The senses of hearing and balance are virtually inextricably linked in
fish, so any attempt to deafen the fish will also interfere with its balance and orien-
tation to gravity. Although there are many experiments in which blind or anosmic
fish were tested for their ability to home or migrate, no studies that I know of test
the effect of deafness or lack of the lateral line on migration or homing.

4.2 Mechanoreceptors
4.2.1 Touch and Proprioception
These two senses have received little recent attention in fishes. The sense of touch
is well developed in fishes and seems to be mediated through the myriad of free
nerve endings in the skin (Marshall 1966). Touch probably plays a role in court-
ship and in responses to substrate. At least in some fish, the same receptors are
responsive to temperature and touch. It is not clear how the two types of informa-
tion are separated (Murray 1971). Proprioception, sensory information about the
movement and posture of the body, is present as well, and could provide informa-
tion about the action of water currents on the fins or other parts of the body. I
know of no studies on the degree to which proprioception is used by fish in orien-
tation or migration.

4.2.2 Pressure
Fish are able to respond to relatively small changes in water (hydrostatic) pressure
(Blaxter and Tytler 1978). In some species this ability is linked with the presence

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R. J. F. Smith, The Control of Fish Migration

© Springer-Verlag Berlin Heidelberg 1985
of a gas-filled swim bladder. Deflation of the swim bladder of the saithe, Polla-
ehius virens, for example, reduces the sensitivity to pressure changes by seven to
ten times, and reinflation of the bladder restores sensitivity (Tytler and Blaxter
1977).
This sensitivity is not dependent on connections with the inner ear. Qutob
(1960) found that blind European minnows still showed good bouyancy regula-
tion and responded to pressure changes of 4-7 cm of water at 70 cm min - 1 even
when the Weberian apparatus was removed. The Weberian apparatus links the
swim bladder mechanically to the ear in the Ostariophysi, minnows and related
fishes. Changes in pressure induced changes in the discharge rate of sensory
nerves from the swim bladder while cannulating the swim bladder and changing
its volume by adding or subtracting gas also induced changes in nerve discharge
(Qutob 1962).
It seems likely that, in fish with swim bladders, stretch receptors in the bladder
provide information on pressure changes. The sensitivity achieved is impressive.
Tsvetkov (1969) reported the thresholds of response of six teleosts as being be-
tween 4 and 20 mm of water. The weather loach, Misgurnus jossilis, responds to
changes in pressure of 4-14 mm of water (Tsvetkov 1972). McCutcheon (1966)
used a tilting tube to test 12 species offish. As the tube is tilted toward the vertical,
the height of the water column increases. He found thresholds as low as 2 mm of
water in some cases. The sea bass, Centropristis striata, showed a slight response
to 0.4 mm of water, a depth change of 0.004%. Blaxter and Tytler (1972) condi-
tioned haddock, Melanogrammus aeglefinus, cod and saithe to pressure changes
over a range of 1 to 20 atm. Responses followed Weber's Law with thresholds
about 0.5% of the adapted value in cod and saithe and 1.2% in hake.
Swim bladders are not essential for pressure sensitivity. Dabs, Limanda
limanda, which lack a swim bladder, could be conditioned to pressure changes of
1-2% of the adapted value (Blaxter and Tytler 1972). Larval plaice and larval
blennies, Centronotus gunnellus, which lack swim bladders, responded to changes
in water depth of25 em. Blennius pholis, which does have a swim bladder, showed
a lower threshold, 5-10 cm of water (Qasim et al. 1963). The mechanisms in-
volved in pressure detection without a gas space are not clear. The otoliths of
some fish are peizoelectric. They produce electric currents in response to mechan-
ical force. But it has not yet been determined whether this plays any role in sen-
sory reception of pressure changes (Morris and Kittleman 1967).

4.2.3 The Lateral Line System

The remaining mechanoreceptor systems: lateral line, hearing and the detection
of gravity and acceleration, are all based on the same basic type of receptor, the
sensory hair cell (Pumphrey 1950). This receptor cell has stereocilia arranged in
a steplike pattern leading up to a larger kinocilium (a true cilium). When the cilia
are bent toward the kinocilium by a water current or similar mechanical stimulus,
the cell becomes depolarized. Bending in the opposite direction polarizes the cell
(Flock 1971). The depolarization increases the spontaneous discharge rate of the
cell.

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