SCI. MAR., 61 (Supl.

1): 159-176 SCIENTIA MARINA 1997

LECTURES ON PLANKTON AND TURBULENCE, C. MARRASÉ, E. SAIZ and J.M. REDONDO (eds.)

Turbulence and ichthyoplankton: influence on vertical

distributions and encounter rates*
SVEIN SUNDBY
Institute of Marine Research. P.O. Box 1870 Nordnes, N-5024 Bergen, Norway.

SUMMARY: Two different aspects of the effects of turbulent mixing on eggs and fish larvae are considered here. In the
first topic the effects of physical processes on the vertical distribution of the eggs are considered. The physical processes
which determine the vertical distribution of fish eggs, and all other immobile plankton, are the buoyancy of the plankton and
the vertical mixing of the water column. Analytical models are presented to demonstrate the influence of the various terms.
A numerical model is also applied to show the effects of non-stationary solutions. The model results are compared with field
measurements. In the second topic the effects of turbulence on plankton encounter rates are considered. The processes are
illustrated by field data on first-feeding cod larvae feeding on Calanus nauplii. These field data show that wind-induced tur-
bulence strongly enhances the feeding rate of the larvae. For first-feeding cod larvae the feeding rate increases by a factor
of about 7 when the wind speed increases from 2 m s-1 to 10 m s-1. Model simulations show how the influence of turbulence-
enhanced encounter rate decreases as the larvae grow, increase their swimming speed and switch to larger prey of lower con-
centration. The simulations show that the turbulence-induced encounter rate decreases exponentially as cod grow, but sig-
nificantly influences feeding until the stage of 2 months old juveniles.

Key words: Fish eggs, buoyancy, vertical mixing, fish larvae, feeding rate, wind-induced turbulence, encounter rate.

INTRODUCTION understanding the effects of turbulence on plankton-

Physical processes influence planktonic life in a
large variety of ways, but only a handful of basic
physical parameters are involved in these processes.
The three most important parameters are tempera-
ture, light and turbulence. A large literature exists on
effects of temperature and light. These parameters
are easy to measure and relatively easy to simulate
and control in experimental designs. Turbulence,
however, is difficult to measure, particularly in field
experiments, and only during recent years accept-
able sensors for turbulence measurements have been
developed. More importantly because of the lack of
*Received December 5, 1995. Accepted April 17, 1996.
ic life many aspects of the theoretical framework are
still insufficiently explored. Mann and Lazier (1991)
gave an overview on turbulence from an ecological
perspective.
Traditionally, the detrimental effects of turbu-
lence have been given focus in ichtyoplankton
research. Much attention has been given especial-
ly to how turbulence by vertical mixing breaks
down patchy high concentrations of food, which
in turn is supposed to decrease feeding and sur-
vival of fish larvae (e.g. Lasker, 1975). Turbu-
lence, however, affects plankton dynamics in dif-
ferent ways, with both positive effects on plankton
production, as well as detrimental effects. Cury
and Roy (1989) studied pelagic fish recruitment in
upwelling areas. They found that there was an

TURBULENCE AND ICHTHYOPLANKTON 159

optimal level of wind velocity with respect to fish
recruitment, and suggested that this was caused by
the opposing effects of wind-induced upwelling,
which enhances plankton production, and turbu-
lent mixing, which has detrimental effects on lar-
val feeding. Studies on grazing copepods indicate
that turbulence may have multiple effects on the
feeding of zooplankton, particularly linked to
behavioral responses to turbulence (Alcaraz et al.
1988; Saiz and Alcaraz, 1992). Consequently, dur-
ing recent years we have become increasingly
aware of the adverse effects of turbulence. For this
reason, proxy variables for turbulence should be
used with care. Investigations of biological
responses and effects of turbulence should prefer-
ably be made by studying the basic biological and
physical processes.
Before exploring how biological processes are
influenced by turbulence, it is important to realise
that light and turbulent mixing are the most promi-
nent physical features of the upper layer of the sea,
the major site for all forms of planktonic produc-
tion. Obviously, light is the basic energy source
for synthesizing biomass. However, turbulence in
the upper ocean, induced by wind stress, transfers
energy from the atmosphere to the euphotic zone,
and turbulence is the most important means for
transferring momentum from the large scale to the
small scale. Therefore, from a general point of
view, it should not be surprising that turbulent
energy also contributes to planktonic biomass pro-
duction.
Principally, there are two quite different aspects
of how turbulence influences plankton. Firstly, tur-
bulence influences plankton distributions, as for
example might be described theoretically by Fick-
ian diffusion. This is a eulerian framework using
eddy exchange coefficients to describe the effects
of turbulence. Secondly, turbulence influences the
encounter rate between individual planktonic parti-
cles, i. e. predators and prey, which is most conve-
niently described in a lagrangeian framework with
direct measurements of the turbulent velocity. Of
the first aspect, I will focus on vertical distribu-
tions and use fish eggs as examples, since their
physical properties are simple in a hydrodynamical
sense and their biological properties are well
described. The second aspect, encounter rates
between planktonic particles, will be illustrated by
investigations on how cod larvae feeding on cope-
podite nauplii are influenced by turbulence-
induced particle interaction.
160 S. SUNDBY
VERTICAL DISTRIBUTIONS OF FISH EGGS
Basic equations
Let us assume that the horizontal variation in egg
distribution can be neglected compared to the verti-
cal variations. The vertical component of the diffu-
sion equation is then the basic equation in describ-
ing the vertical distribution of eggs:
∂ C(z, t) ∂ [ w(z, t)C(z, t)]
– =
∂t ∂z
∂ ⎡ ∂ C(z,t) ⎤
= K(z,t) + S(z,t) + M(z,t) (1)
∂ z ⎢⎣ ∂ z ⎥⎦
where
C(z,t) concentration of eggs in numbers per
unit volume
w(z,t) vertical velocity of the eggs
K(z,t) vertical eddy diffusivity coefficient
S(z,t) spawning (production) of eggs
M(z,t) egg mortality
If we neglect the influence of spawning and egg
mortality on the vertical distribution and consider
stationary conditions, equation (1) reduces to
∂ C(z)
–w(z) ⋅ C(z) = K(z) (2)
∂z
The equation expresses the balance between the
vertical velocity of the eggs (left hand side) and the
eddy diffusion flux of the eggs (right hand side).
Equation (2) can be solved to give the vertical
distribution of eggs. The solution depends on the
shape of the functions describing vertical velocity,
w(z), and eddy diffusivity, K(z). Let us, therefore,
first consider the factors which determine the values
of these two variables.
Buoyancy and vertical velocity of eggs
The terminal velocity of a particle moving in a
fluid is the constant velocity reached when there is
balance between gravity forces and frictional forces.
For a sphere like a fish egg it is a function of the
gravity acceleration, g, the egg diameter, d, the
buoyancy of the egg, ∆ρ= ρw - ρe, where ρe is the
specific gravity of the egg and ρw is the specific
gravity of the ambient water. When the Reynolds’
number, Re= wd/ν (where ν is the molecular vis-
cosity of the water), is less than about 0.5, the vis-
cous forces dominate the frictional force and the ter-
minal velocity is expressed by Stokes equation (See
Tim Pedley´s lecture, this volume):
w = 1/18 g d2 ∆ρ ν-1 (3)
When the Reynolds’ number exceeds 0.5 viscous
forces are starting to become less important due to
an increase of the turbulent forces. For a moderate-
ly large Reynolds’ number the frictional forces are
influenced by both viscous and turbulent friction
and an empirical expression for the terminal veloci-
ty was found by Dallavalle (1948):
w = KI (d -ζD) ∆ρ2/3 ν-1/3 (4)
where KI is a constant, ζ is a constant equal to 0.4
for spheres and D is the uppermost limit of egg
diameter to which the Stokes equation applies.
Hence, the terminal velocity becomes linearly pro-
portional to the egg diameter.
For completely turbulent conditions, when the
Reynolds’ number reaches the order of 103, the ter-
minal velocity is proportional to the square root of the
diameter. It is shown in Figure 1 that fish eggs are
FIG. 1. – Terminal velocity as a function of egg diameter for a range
of buoyancies, ∆ρ. Observed values of egg diameter and buoyan-
cies in their natural habitats for five different species of fish eggs
are plotted on the graph. 1. North Sea mackerel; 2. Arcto-Norwe-
gian cod; 3. North Sea plaice; 4. Namibian Cape hake; 5. Atlantic
halibut at the coast of northern Norway.
found in the Stokes’ regime or in the lower part of the
intermediate regime, where the Reynolds’ number is
generally lower than 5. The value of KI in equation
(4) is not a true constant in the whole range of the
intermediate region. However, for the actual range of
Reynolds numbers, i.e. < 5, we assume that KI is
approximately constant. A numerical expression for
KI can then be obtained by setting equations (3) and
(4) equal to the value of w at which Re=0.5. This
gives KI=19 (Sundby, 1983). In Figure 1 the terminal
velocity is plotted against egg diameter for a range of
buoyancy values, ∆ρ, using equations (3) and (4).
After Coombs (1981) introduced the density-gra-
dient column to measure the neutral buoyancy of
fish eggs, new possibilities appeared to understand
the processes behind the vertical distribution of
ichthyoplankton. By this laboratory technique it is
possible to measure the density of individual fish
eggs to an accuracy of about 0.04×10-3 g cm-3.
Many species of fish eggs have a specific gravi-
ty which makes them positively bouyant in the
upper mixed layer. They have a neutral buoyancy,
measured as salinity, which is lower than the salini-
ty of the upper layer. They are distributed as group
A in Figure 2 and are termed “pelagic” eggs (Sund-
by, 1991). Four of the egg species plotted in Figure
1 are pelagic eggs. Some species have eggs which
are heavier than the upper layer, but lighter than the
density of the deep layer. They are distributed as
group B in Figure 2 and are termed “bathypelagic
eggs”. Examples of such types are eggs of Pacific
FIG. 2. – Buoyancy distributions of three main groups of fish eggs in
relation to the salinity profile shown below, as defined by Sundby
(1991). A. Pelagic eggs, ∆ρ > 0; B. Bathypelagic eggs, ∆ρ = 0; C.
Bottom eggs, ∆ρ < 0. The salinity profile shows a typical profile for
coastal boreal regions with a upper mixed layer of lower salinity, a
halocline below and a deep homogeneous deep layer of high salinity.
TURBULENCE AND ICHTHYOPLANKTON 161
 FIG. 3. – Section through an Atlantic cod egg with developed embryo.
halibut (Thompson and van Cleve, 1936), Baltic cod
(Kändler, 1949) and Atlantic halibut (Haug et al.,
1984). Group C in Figure 2 is defined as bottom
eggs and are heavier than the density of the bottom
layer. Examples of such eggs are the saffron cod
eggs in the northeast Pacific Ocean (Dunn and
Matarese, 1986) and Barents Sea capelin eggs
(Bakke and Bjørke, 1973). It should, however, be
emphasized that eggs from various species may also
have buoyancy distributions which fall between the
three main groups defined above. Coombs et al.
(1985), for example, showed that eggs of sprat and
pilchard off the south coast of Great Britain had a
buoyancy distribution between pelagic and bathy-
pelagic eggs and their buoyancy changed through
development. Kendall and Kim (1986) demonstrat-
ed that bathypelagic eggs of walleye pollock may
substantially change their vertical distribution due to
changes in buoyancy during the egg development.
We now explore the properties which determine
the buoyancy of eggs. I will use investigations on
Atlantic cod eggs (Kjesbu et al., 1992) as an exam-
ple. Figure 3 shows a section through an Atlantic
cod egg with the embryo partly developed. The
diameter of this egg ranges from 1.1 to 1.6 mm.
The egg shell, the chorion, is the heavy fraction of
the egg. Kjesbu et al. (1992) found that the specific
gravity of the chorion was 1.20 g . cm-3. The thick-
ness is, however, only 5 - 9 microns. Holes in the
chorion allows seawater to penetrate it, and sea
water of the same density as the ambient seawater is
found in the perivitelline space, which is the space
between the chorion and the vitelline membrane.
The volume of the perivitelline space may vary. For
162 S. SUNDBY
Arcto-Norwegian cod eggs it is quite small, only 3 -
5% of the total egg volume.
The yolk and embryo constitute the light fraction
of the egg and is surrounded by the vitelline mem-
brane across which the osmoregulation occurs and
prevents the embryo and yolk from being dehydrat-
ed. The density of the embryo and yolk of Arcto-
Norwegian cod eggs is about 1.017 g cm-3. The
ambient salinity of Arcto-Norwegian cod eggs is 33
- 34 p.s.u. In contrast, Baltic cod eggs, which devel-
op in the halocline at low salinities, less than 10 -15
p.s.u., have developed eggs with a high water con-
tent which makes them much more buoyant
(Nissling and Westin, 1991). This is an ecological
adaptation in Baltic cod to prevent the eggs from
sinking down to the anoxic deep water of the Baltic
Sea. The specific gravity of the embryos of these
eggs is about 1.008 g cm-3, and the eggs are big, 1.8
- 1.9 mm in diameter, and have a thin chorion. It has
been shown that the spawning female cod invests a
relatively constant mass of chorion in the eggs inde-
pendent of egg size. This implies that bigger eggs
generally have thinner chorions. The specific gravi-
ty of the egg, ρe , can then be expressed:
ρe = ρi + (ρch - ρi) Vch/(4/3πr3) (5)
where ρi is the specific gravity of the embryo and
yolk, ρch is the chorion specific gravity, Vch is the
chorion volume and r is the radius of the egg.
Figure 4 shows graphs of equation (5) for three
different values of chorion volumes, the maximum,
the mean, and the minimum volumes. It can be seen
from the figure that egg size within the natural range

FIG. 4. – Egg specific gravity of Arcto-Norwegian cod as a function
of egg diameter for: the minimum chorion volume (26 x 106 mm3),
lower line; the mean chorion volume (33 x 106 mm3), middle line;
and the maximum chorion volume (40 x 106 mm3), upper line.
of variation (1.1 - 1.6 mm) contribute more to vari-
ations in specific gravity than the variation in chori-
on volume (26 x106 mm3 - 40 x106 mm3).
Vertical eddy diffusivity coefficient
The other variable which influences the vertical
distribution of eggs is the vertical eddy diffusivity
coefficient (K in equation (1) and (2)). Depending
on depth, wind velocity, stratification, surface
cooling and convection, tidal energy and bottom
stress, it varies over approximately five orders of
magnitude. It is normally largest in the mixed layer
and decreases to a minimum in the pycnocline due
to the strong buoyancy forces which act against
vertical mixing. In regions of strong bottom cur-
FIG. 5. – Qualitative shape of the eddy diffusivity coefficient, K in
cm2s-1, (right part of the figure) for a hydrographic profile (left part
of the figure) identical with the profile in Figure 2. The shaded area
indicates typical range of values for eddy diffusivities.
rrent the vertical eddy diffusivity coefficient might
increase in and above the bottom boundary layer
due to bottom friction. Figure 5 shows qualitative-
ly how the vertical eddy diffusivity might vary
through a 100 m deep water column with the pyc-
nocline at 30 - 50 m depth.
Estimating the mixed layer eddy viscosity coeffi-
cient is difficult, partly due to technical problems in
measuring the wave zone. Sverdrup et al. (1942)
derived estimates of the eddy viscosity coefficient
from Ekman theory. Sundby (1983) estimated over-
all eddy diffusivity coefficients for the mixed layer
from a model based on the vertical distribution of
pelagic eggs. Thorpe (1984) estimated the eddy dif-
fusivity coefficients in the surface layer based on a
model of the vertical distribution of air bubbles in
the sea. Although their results differ to some extent,
it may be concluded that the eddy diffusivity coeffi-
cient ranges from about 10 cm2 s-1 at wind speeds
near zero to about 103 cm2 s-1 during strong surface
mixing in stormy weather.
In the pycnocline the eddy diffusivity coeffi-
cient is inversely related to stratification and
directly dependent on energy input. Examples of
authors who have estimated the vertical eddy dif-
fusivity in pycnoclines are Gade (1970) for
Oslofjorden, and Kullenberg (1971) for shallow
coastal waters. Gargett (1984) reviewed the litera-
ture on vertical diffusivity coefficients in stratified
systems. Depending on the level of stratification,
the eddy diffusivity coefficient ranged from
0.5x10-2 cm2 s-1 to 4 cm2 s-1.
Bottom turbulence, which normally extends sev-
eral metres above the bottom, is mainly dependent
on the boundary layer velocity and bottom rough-
ness. Bowden (1962) reported values from several
authors. In areas of strong tidal mixing the eddy dif-
fusivity coefficient may exceed 100 cm2 s-1. Above
the seabed in deep oceanic areas 1 cm2 s-1 is more
common.
Analytical solution for pelagic eggs
Pelagic eggs which have a neutral buoyancy
distribution like A in Figure 2 will ascend towards
the surface. As we assume that there is no change
of density within the mixed layer, the vertical
velocity of eggs as a function of depth is constant
(w(z)=const.). If we also assume that the eddy dif-
fusivity coefficient through the mixed layer is con-
stant (K(z) = const.), the solution to equation (2)
becomes:
TURBULENCE AND ICHTHYOPLANKTON 163

FIG. 6. – Vertical distribution of pelagic eggs as shown in equa-
tion (6) for two values of w/K. Profile I: w/K is large. Profile II:
w/K is small.
C(z) = C(a) exp - w/K (z-a) (6)
where C(a) is the egg concentration at a given depth,
a. Equation (6) is a simple exponential solution
where the slope of the curve (Figure 6) is deter-
mined by the ratio w/K. When the ascending veloc-
ity of the eggs are large compared to the eddy diffu-
sivity coefficient (w/K is large) the egg concentra-
tion has the type I profile in Figure 6. When the
ascending velocity of the eggs is small compared to
the eddy diffusivity coefficient (w/K is small), the
egg concentration has the type II profile in Figure 6.
The solution in equation (6) is valid for one sin-
gle group of eggs with constant neutral buoyancy
FIG. 7. – Observed vertical distributions of Arcto-Norwegian cod
eggs based on measurements of Solemdal and Sundby (1981) (dot-
ted lines) and nonlinear regression of the observed values based on
the solution by Sundby (1983) (continuous lines).
164 S. SUNDBY
and diameter. However, as shown in Figure 2 one
population of eggs has a neutral buoyancy which is
most often Gaussian distributed around mean val-
ues. Sundby (1983) extended the solution to a
Gaussian distributed population of pelagic eggs.
This analytical solution is more complicated. How-
ever, for the buoyancy distributions in question
(Arcto-Norwegian cod eggs and North Sea macker-
el eggs), the results do not differ substantially from
the more simple solution of equation (6).
The extended solution by Sundby (1983) was
fitted by nonlinear regression to measured vertical
profiles of Arcto-Norwegian cod eggs off the coast
of Northern Norway for various wind situations,
and hence mixing conditions (Figure 7), and mea-
surements by Iversen (1973) of North Sea macker-
el eggs under wind conditions from Beaufort 0 to
6 (Figure 8).
The nonlinear regressions for the measurements
of Arcto-Norwegian cod eggs, North Sea mackerel
eggs and North Sea Plaice eggs made it possible to
estimate the eddy diffusivity coefficients for the
mixed layer, as a function of the wind speed (Figure
9) (Sundby 1983). The relation found was:
K (cm2 s-1)= 76.1 + 2.26 W2 (7)
where W is the wind speed in m s-1.
FIG. 8. – Observed concentrations (crosses) of North Sea mackerel
eggs by Iversen (1973) from Beaufort 0 to 6 wind force, and non-
linear regression of the observed values based on the solution by
Sundby (1983).
FIG. 9. – Computed eddy diffusivity coefficients, K, based on mea-
surements on vertical distributions of North Sea mackerel eggs,
North Sea plaice eggs, and Arcto-Norwegian cod eggs.
(After Sundby 1983).
The results are comparable with estimates by
other methods, and demonstrate that plankton, like
fish eggs, can be applied to estimate physical prop-
erties when precise measurements of buoyancy by
the method by Coombs (1981) are applied. In Figure
10, equation (7) is inserted in the model for pelagic
eggs (equation (6)) to demonstrate how wind mixing
influences the vertical distribution of Arcto-Norwe-
gian cod eggs and the lighter North Sea mackerel
eggs with higher ascending velocity. As seen in the
figure the lighter mackerel eggs are concentrated
more towards the surface during moderate wind
FIG. 10. – Theoretical computed vertical profiles of egg concen-
tration of mackerel eggs and cod eggs at K=80 cm2 s-1 (wind speed
of 2 m s-1), K=400 cm2 s-1 (wind speed of 12 m s-1), and
K=1150 cm2 s-1 (wind speed of 22 m s-1).
conditions. During strong wind conditions (e.g. 22
m s-1) vertical mixing dominates over the ascending
velocity of the eggs and the difference between the
vertical distribution of the two eggs species becomes
negligible. The figure also shows that the egg pro-
files undergo the largest changes in the range
between 0 and 12 m s-1 of wind speed.
The analytical solution presented above is a
steady state solution. Westgård (1989) developed a
numerical solution which allows for studying tem-
poral variations. Figure 11 presents some applica-

FIG. 11. – Development of the vertical profile of pelagic eggs (type A buoyancy distribution as
shown in Figure 2) from an initial distribution of spawning near the bottom at 120 m depth to steady
state distribution. Two events of mixed layer turbulence are shown. Upper part: wind velocity,
W = 0 m s-1. Lower part : wind velocity , W = 15 m s-1. (After Sundby 1991).

TURBULENCE AND ICHTHYOPLANKTON 165

tion of this numerical model as shown by Sundby
(1991). The figure presents two scenarios of cod
spawning at 120 m depth in a salinity profile as
shown in Figures 2 and 5. The profiles display how
the eggs ascend from the spawning depth and final-
ly reach the steady state solution when there is bal-
ance between the buoyancy and eddy diffusion flux-
es. The upper profiles show that the equilibrium pro-
file is reached after about 48 hrs. during calm con-
ditions, i.e. without wind and wind-generated turbu-
lence. During strong wind-induced mixing (wind
speeds of 15 m s-1) the equilibrium profile is reached
after only 30 hrs.

Analytical solution for bathypelagic eggs
Bahypelagic eggs which have a neutral buoy-
ancy distribution like B in Figure 3 will ascend
from the deep layer, but will descend from the
upper mixed layer. Their neutral buoyancy is
found in the pycnocline. We consider one single
buoyancy group of bathypelagic eggs. We assume
that the pycnocline is linear, and may therefore be
expressed:
ρ(z) = kz + b (8)
where ρ(z) is the density as a function of depth, z,
and k and b are constants.
The vertical velocity will then vary linearly with-
in the Stokes regime (equation 3), and may therefore
be written:
w(z) = m(z - zA) (9)
where m is a constant and zA is the depth level where
∆ρ(z) = 0, i.e. the level of neutral buoyancy of the
egg. We now assume that the eddy diffusivity coef-
ficient is constant in a linear pycnocline, and equa-
tion (2) can then be written:
- m(z - zA) . C(z) = K dC(z)/dz (10)
The solution to equation (10) is:
m eggs are large compared to other fish eggs, 3.0 - 3.5
C( z ) = C A exp ⎡⎢ − z − z A ) ⎤⎥
2
( (11)
⎣ 2K ⎦ (12) and (13), the standard deviation, σ, of the ver-
where CA is the concentration of eggs at the depth of
neutral buoyancy of the eggs, zA. It appears from
equation (11) that the bathypelagic eggs are verti-
cally distributed as a normal distribution with a stan-
dard deviation:
FIG. 12. – Vertical distribution of bathypelagic eggs in a strong pyc-
nocline where the eddy diffusivity coefficient is small and in a weak
pycnocline with a larger eddy diffusivity coefficient.
σ = (K/m)1/2 (12)
around the the mean depth level zA.
When the velocity of the eggs is confined within
the Stokes regime, the Stokes equation (equation 3)
for the terminal velocity is valid and the expression
for m in equation (12) becomes:
m = 1/18 d2 ν-1 ρw N2 (13)
where N is the Brunt-Väisälä frequency (with the
unit s-1).
Figure 12 shows the graphical form of the solu-
tion (equation (11)). The distribution is narrow
when the eddy diffusivity coefficient is small and m
(proportional to the the density gradient) is large.
To give an impression of the magnitude of the stan-
dard deviation of the vertical distribution of bathy-
pelagic eggs, let us take Atlantic halibut eggs in the
fjords of Northern Norway as an illustrative exam-
ple. Typically, as for the density profiles of Figure
13, the Brunt-Väisälä frequency ranges from
0.5x10-4 to 2.0x10-4 s-2. From the above-mentioned
literature on the influence of stratification on the
turbulence, the eddy diffusivity coefficients range
from 0.1 to 0.5 cm2 s-1. The diameter of the halibut
mm. When these values are inserted into equations
tical spreading of one bouyancy group of halibut
eggs will range from 0.4 to 1.6 m. However,
according to Haug et al. (1986) (Figure 13), the
older eggs (which have come to a steady state ver-
tical distribution) extend over a 150 - 250 m water

166 S. SUNDBY
 FIG. 13. – Density profiles and profiles of Atlantic halibut eggs in fjords of Northern Norway (after
Haug et al. 1986).

column. Consequently, the large vertical spreading
of halibut eggs observed in the water column must
be due to a spreading in the neutral buoyancy dis-
tribution of the eggs alone and is not caused by ver-
tical turbulence.
Atlantic halibut eggs are spawned in the deep
water below the pycnocline, most often near the bot-
tom at 400 - 800 m depth in Norwegian fjords. Con-
sequently, they have to ascend several hundred
meters before they reach the level of equilibrium at
the pycnocline. In Figure 14 one of the profiles from
Haug et al. (1986) is used to demonstrate the time it
takes to ascend from a spawning depth of 450 m.
The right part of the figure shows vertical position
as a function of time in days for three buoyuancy
groups of eggs. Group I is the lightest fraction and
has a specific gravity of 1.0270 g cm-3 which give
eggs an equilibrium level at 120 m depth. Group II,
the average fraction, has a specific gravity of
1.0272 g cm-3 with an equilibrium depth of 160 m.
Group III, the heavy fraction, has a specific gravity
of 1.0274 g cm-3 with the equilibrium level at 200 m
depth. Despite the small differences in specific
gravity the time to reach the respective equilibrium
depth levels are very different for the heavy group
compared to the two other groups. Groups I and II
reach the equilibrium level at about 4 - 5 days, while
the heaviest fraction which has the equilibrium level
at 200 m depth, only 250 m above the spawning
depth will use more than 9 days to reach that level.

TURBULENCE AND ICHTHYOPLANKTON 167


FIG. 14. – Left panel: Atlantic Halibut eggs profile (dashed line)
and water column density profile (continuous line) (After Haug et
al. 1986); Mid panel: Calculated vertical velocities for three buoy-
ancy groups of eggs, i.e. respectively 1.0270, 1.0272 and 1.0274 g
cm-3; Right panel: Calculated vertical positions as a function of time
for the three egg groups initially released at 450 m depth.
This is because the buoyancy of the heavy fraction
gives a very low ascending speed of those eggs in
the deep homogeneous layer.
LARVAL FISH FEEDING AND THE ROLE OF
TURBULENCE-INDUCED CONTACT RATE
Basic theory
The influence of turbulence on plankton
encounter rates is a biophysical process which sub-
stantially alters traditional considerations about
energy demands and behaviour of larval fish, and
consequently the recruitment processes. The basic
idea was presented by Rothschild and Osborn
(1988), and they developed the theoretical frame-
work necessary to investigate the influence of turbu-
lence on plankton contact rates in the laboratory and
in the field. Their theory was in turn built on the the-
ory of Gerritsen and Strickler (1977) on the
encounter rate between a cruising predator and a sto-
chastically moving prey.
To give an intuitive and qualitative understand-
ing of the processes which enhance contact between
predators and prey, let us look at Figure 15. The
three panels of the figure show a fish larva and its
prey, for example copepod nauplii. The tubes of the
three panels conceptually indicate the swept volume
by the fish larva between each prey encounter. The
upper panel indicates the traditional view of how a
larval cruise predator searches through the water for
prey. By this concept it is assumed that the relative
motion between the predator and prey is caused
168 S. SUNDBY
alone by the moving predator. Neither prey nor the
fluid environment has a velocity. The number of
encountered prey per time unit is then linearly pro-
portional to the search volume of the predator, and
can be expressed by the relation:
Z = N π R2. v (14)
where Z is the contact rate between larvae and their
prey, N is the number of prey particles per volume
unit, R is the perceptive distance for the fish larva,
and v is the swimming speed of the fish larva.
Hence, there is a linear relationship between the
contact rate, Z, and the swimming speed, v. Gerrit-
sen and Strickler (1977) pointed out that this is an
oversimplification. They showed that the relative
velocity between predator and prey is not a function
of the predator velocity alone, but also of the prey
velocity. They developed a model where the preda-
tor is cruising in a cloud of stochastically moving
prey. When predator speed is higher than the prey
speed the contact rate becomes:
Z = N π R2 (u2 + 3v2)/3v (15)
where u is the prey speed. The velocity component
of the contact rate, (u2 + 3v2)/3v, in the equation
above is larger than the velocity component, v, of
equation (14). The middle panel of Figure 15
shows the effect of the Gerritsen and Strickler
(1977) model, and it implies that the fish larvae
will need to swim a shorter distance to maintain the
same encounter rate as in the situation of the upper
panel. Hence, the tube of swept volume between
each encountered prey is shorter than in the upper
panel.
Let us use this model on the event of cod larvae
feeding on copepod nauplii. The mean swimming
speed of these nauplii are only 10% of the mean
cruising speed of the larvae. The enhancement of the
encounter rate using Gerritsen and Strickler’s model
is (u2 + 3v2)/3, and for cod larvae cruising at mean
speed it implies an enhancement in encounter rate of
only 0.3%. Only when the cod larva is cruising very
slowly, at speeds comparable to the speed of the
nauplii, is there a considerable improvement in the
contact rate, 33%, compared to the traditional model
(equation (14)). Even though the Gerritsen and
Strickler model did not give significant practical dif-
ferences in encounter rate for the case of cod larvae,
it was principally a very important theoretical devel-
opment.
 FIG. 15. – Three model concepts of larval encounter rate. Upper panel: The traditional concept of
encounter based on swept area by the larva. Only the larva is moving. Middle panel: Encounter model
according to the Gerritsen and Strickler (1977) model with stochastically moving prey. Lower panel:
Encounter rate model according to Rothschild and Osborn (1988) with stochastically moving prey
and ambient turbulence.

Rothschild and Osborn (1988) pointed out that in
nature turbulence contributes more to the stochastic
velocity component than the prey. They added the
turbulent velocity to the formulation in equation
(15) and arrived at a modified Gerritsen and Strick-
ler model:
Z = N π R2 (u2 + 3v2 + 4w2)/3(v2 + w2)1/2 (16)
where w is the root-mean-square turbulent velocity.
For typical turbulent velocities in nature the
velocity component in equation (16) becomes con-
siderably larger than in equation (15), and the larva,
in the lower panel, will have to swim a considerably
shorter distance to maintain the same encounter rate
as in the mid panel, again indicated by the length of
the tube. The other aspect of this model is that the
natural turbulence becomes so important that the
encounter rate is in fact quite insensitive to the
swimming speed of the cod larvae within its natural
range.
The nature of turbulence is that it occurs on all
scales at the same time and with a range of turbulent
velocities increasing with increasing scale. There-
fore, we need to sort out how we can apply in equa-
tion (16) the range of turbulence velocities. The
basic relation of scales in natural turbulence (ocean-
ic and atmospheric turbulence) as first postulated by
Kolmogorov (1941), is that turbulent energy cas-
cades from large scale to smaller and smaller scales
until it dissipates to heat by shear strain of molecu-
lar movement. A universal relation between the

TURBULENCE AND ICHTHYOPLANKTON 169

energy spectrum and wave number, k, of turbulence
of high Reynolds numbers in the inertial subrange
exists, and this is proportional to k-5/3. Therefore, tur-
bulence on all length scales are found simultaneous-
ly in a body of water in the ocean. Many later mea-
surements from the field have confirmed such a tur-
bulent energy spectrum (e.g. Grant et al. 1962). The
consequence of this physical nature is the well-
known fact that eddy diffusivity increases with
increasing length scale (Okubo, 1978). This means
that the spreading of a certain property in nature, for
example fish eggs, increases with time and with the
size of the distribution. This is simply because larg-
er and larger turbulent eddies contribute to the
spreading, or rearrangement of the patches of eggs.
Similarly, the relative motion between two individ-
ual particles due to turbulence will increase as the
distance between them increases. This basic process
contributes simultaneously to spreading of the parti-
cles and contact between them. So, both spreading
and contact rate is scale dependent in natural (aquat-
ic and atmospheric) turbulence.
Rothschild and Osborn (1988) developed the
relation between the turbulent scales and the associ-
ated turbulent velocities, the root-mean-square
velocity. This development was summarized in Fig-
ure 1 of their paper (Figure 16 here), and shows how
the turbulent velocity increases with increasing
length scale.
FIG. 16. – The relation between length scale, separation distance,
and the associated root-mean-square turbulent velocity at various
levels of turbulent energy dissipation rates (from Rothschild and
Osborn, 1988).
170 S. SUNDBY
Sundby and Fossum (1990) applied the theory of
Rothschild and Osborn (1988) to field observations
of first-feeding cod larvae feeding on Calanus fin-
marchicus nauplii. The turbulent scales which con-
tribute to increase the contact rate are at all scales
smaller than the typical separation distance between
predator and prey. The typical nauplii concentra-
tions, c, for first-feeding cod larvae are 1 - 50 liter-1,
which correspond to deterministic separation dis-
tances, r=c-1/3, from 10 to 3 cm. Against this back-
ground Sundby and Fossum (1990) used 5 cm as an
average separation distance. Depending on the level
of turbulent kinetic energy dissipation, ε, the root-
mean-square velocity, w, for such separation dis-
tances will typically range from the order of 0.01 -
1 cm s-1, as calculated according to Rothschild and
Osborn (1988). By applying equation (4) of Roth-
schild and Osborn (1988), valid for scales larger
than the Kolmogorov scale, the relevant turbulent
velocity, w, can be calculated:
w = 1.9 (ε d)1/3 (17)
where d is the turbulent length scale, and w is the
turbulent velocity associated with that length scale.
The turbulent length scales which increase the
contact rate between predator and prey are all length
scales smaller than the separation distance between
the predator and prey, because all such length scales
contribute to the relative velocity between them. For
fish larvae where the concentration is typically
much lower than the concentration of the prey items,
the separation distance is effectively equal to the
separation distance between the prey items, r. Tur-
bulent cells of a scale, d, larger than the separation
distance, r, will not contribute to increase the contact
rate, because those cells will move larger water
parcels without rearranging the relative distance
between the particles.
To get the right impression of the proper length
scales involved in plankton contact rates, and partic-
ularly the length scales involved in feeding cod lar-
vae, let us look at Figures 17 - 20. Figure 17 shows
the main prey organism of Arcto-Norwegian cod
larvae, a Calanus finmarchicus nauplius stage III.
The dots around the nauplius indicate the average
distance between phytoplankters in a typical spring
bloom of 106 cell l-1. In Figure 18 a first-feeding cod
larva is shown with a Calanus finmarchicus nau-
plius stage III, at the maximum reactive perceptive
distance which is about 0.45 cm. The vectors indi-
cate the average swimming speeds of the larva and
FIG. 17. – Calanus finmarchicus nauplius stage III, the main prey
organism for first-feeding Arcto-Norwegian cod larvae. The dots
indicate the food concentration of phytoplankters in a normal
spring bloom.
FIG. 18. – First-feeding cod larva and its main prey, Calanus fin-
marchicus nauplius stage III. The circle indicates the maximum
reactive perceptive distance, 0.5 cm, and the arrows indicate the
mean swimming speeds of the two organisms, 0.17 and 0.02 cm s-1,
respectively.
the nauplius, 0.17 and 0.02 cm s-1, respectively. Fig-
ure 19 shows the cod larva swimming in a situation
of very high food concentration of Calanus nauplii,
50 nauplii litre-1. The square-shaped clouds of parti-
cles around two of the nauplii indicate the phyto-
plankton particle densities in a spring bloom of 106
cells per litre, which is the same concentration as
shown in Figure 17, and of a particle density of 105
cell l-1. Figure 20 shows the first-feeding cod larva
in a low concentration of prey, 1 nauplii l-1. Even
this low concentration has been shown to give suffi-
cient feeding conditions for cod larvae under turbu-
lent conditions (Sundby and Fossum, 1990).
FIG. 19. – Cod larva (with the maximum perceptive distance indi-
cated by the half circle) in a very high concentration of nauplii, 50
nauplii l-1. The dimensional size of the nauplii are exaggerated; oth-
erwise they would have been invisible. The square shaped clouds of
dots around two of the nauplii indicate the concentration of a phy-
toplankton spring bloom of 106 plankters l-1 and a concentration of
105 plankters l-1. Also the dimensional size of the
phytoplankters are exaggerated to allow visibility.
FIG. 20. – Cod larva (with the maximum perceptive distance indi-
cated by the half circle) in a low concentration of nauplii, 1 nauplii
l-1, but still high enough concentration to survive in turbulent con-
ditions. The dimensional size of the nauplii are exaggerated;
otherwise they would have been invisible.
Application of theory on field data
Field evidence for the influence of turbulence on
the feeding rate of fish larvae has been provided by
Sundby and Fossum (1990) and Sundby et al.
(1994). They investigated the gut content, i.e. the
average number of nauplii in the gut, of cod larvae
sampled at the first-feeding areas in Lofoten, North-
ern Norway, and compared it to the nauplii concen-
tration in the sea under various wind situations.
TURBULENCE AND ICHTHYOPLANKTON 171
 The relation between the average number of prey
particles in the larval gut, A(c), (termed feeding
ratio) and the nauplii concentration in the sea, c, can
be expressed as follows:

A(c) = Amax [1 - e-(b · c)] (18)

where Amax is the maximum number of prey in the

larval gut, i.e. full gut, and b is the coefficient
which determines the slope of the function in equa-
tion (18). The function is shown in Figure 21. The
coefficient, b, contains all the environmental fac-
tors which influence larval feeding. Light condi-
tions and turbulence are the most important factors FIG. 22. – Feeding ratio of cod larvae versus naupliar concentration
influencing larval feeding in addition to prey con- in the sea for all data (259 points) presented by Sundby et al.
(1994). One point represents one sample depth. Graphs of equation
centration. (18) for b = 0.02 and 0.2 are plotted into the figure.
Sundby and Fossum (1990) and Sundby et al.
(1994) fitted their field data to equation (18) by
nonlinear regression analysis. Data sampled dur- feeding. The panels in Figure 23 show larval feed-
ing night time (10 p.m. to 2 a.m.) were omitted in ing under increasing wind conditions from wind
order to minimize the effect of reduced light on the speeds of 2.0 m s-1 to 10.5 m s-1, grouped in four
feeding conditions. During this period the feeding wind speed intervals: 2.0 m s-1, 3.2 - 4.0 m s-1, 5.4 -
ratio is lower than during day time. By omitting 6.9 m s-1, and 7.5 - 10.5 m s-1. In all there are 13
the data from the dark part of the day, it is expect- larval sampling stations distributed in the four
ed that the variations in the coefficient b are main- wind speed intervals. The different values of the
ly caused by variations in the turbulence-induced coefficient, b, for the 13 larval sampling stations
feeding rate. In Figure 22 all the data sampled by are plotted against wind speed in Figure 24. The
Sundby et al. (1994) is presented. Two graphs of figure indicates that turbulence-induced feeding
equation (18) for b = 0.02 and b = 0.2 are plotted rate is increased by a factor of 9 when the wind
onto the figure. Initially it seems that the data indi- speed increases from 2 to 10 m s-1.
cate no functional relationship. However, when
the data sampled under equal turbulent conditions
with respect to wind-induced turbulence were
grouped together and the coefficient b was deter-
mined for each event (Figure 23), it is evident that
wind-induced turbulence together with food con-
centration are the main causes of variable larval

FIG. 23. – Feeding ratio of cod larvae versus naupliar concentration

FIG. 21. – The functional relation between number of nauplii in the in the sea. The data are pooled into four wind groups. Panel 1: 2.0
larval gut (here termed feeding ratio) and nauplii concentration in m s-1; Panel 2: 3.2 - 4.0 m s-1; Panel 3: 5.4 - 6.9 m s-1; Panel 4:
the sea. 7.5 - 10.5 m s-1.

172 S. SUNDBY
FIG. 24. – The coefficient b in equation (18), derived from nonlinear
regression, as a function of wind speed.
At a wind speed of 4 m s-1 the turbulent veloci-
ty contributes more to contact rate than the larval
swimming speed. Figure 25 shows the contact rate
as a function of swimming speed for 4 different
events of turbulent conditions. The lower graph
shows the relation in the absense of turbulence, i.e
the Gerritsen and Strickler (1977) formulation
(equation (15)). The upper three graphs show the
Rothschild and Osborn (1988) formulation for tur-
bulent conditions corresponding to wind speeds of
2.5, 5.0 and 7.5 m s-1 respectively. The ranges of
the observed swimming speeds of first-feeding cod
larvae in the laboratory (Solberg and Tilset, 1984)
are indicated by the shading. It is clear from the
figure that at wind speeds above 5 m s-1 the benefit
for the larvae to increase the contact rate by its own
motion is negligible, and the larvae will, conse-
quently, waste its energy resources if cruising for
food, because turbulence will do the job anyway.
From this point of view the larvae should save its
energy resources for the short final attack on the
prey and for rapid escape reactions from predators.
Munk and Kiørboe (1985) indeed showed that this
is what occurs with herring larvae when the
encounter rate increases: The swimming activity is
reduced and the attact rate increases. Larval growth
itself is an important factor for survival, and it is
unlikely that the larvae are adaptated to a energy-
wasting behaviour like excessive cruising. Turbu-
lence (together with light) is the most characteris-
tic feature of the mixed layer, the site where most
of the plankton production occurs, and therefore it
is reasonable to believe that the cruising speed in
FIG. 25. – Theorically calculated contact rate (after the Roth-
schild and Osborn formulation) as a function of larval swimming
speed for four different turbulent conditions. Lower line is with-
out turbulence. The upper lines are for wind speeds of 2.5, 5.0,
and 7.5 m s-1. The dotted region shows the observed range of cod
larval swimming speeds.
the field is lower than in the laboratory where tur-
bulence intensities most often are low.
Above I have considered feeding by first-feeding
fish larvae which are small and slow-swimming
organisms compared to the turbulence velocities on
that scale. However, larger-scale turbulence may
also have the potential to enhance contact rates for
larger organisms, if the separation distance between
their prey is comparably larger. Sundby (1995)
analysed the influence of wind-induced turbulence
on cod larvae as they grow, increase their swimming
speed and change to larger prey items of lower con-
centrations.
Based on data of post-larval-cod growth rate
Sundby (1995) arrived at a relation for the average
swimming speed as a function of age:
v(t) = 0.152.exp{5.9.[1-exp(-0.0050 t)]} (19)
where v(t) is the age dependent swimming speed
and t is age in days.
As the Arcto-Norwegian cod larvae grow they
change mainly to feed on larger copepodite stages
of Calanus finmarchicus (Sysoeva and Degtereva,
1965). The main diet of early juvenile cod (70 d)
consists of copepodite stages IV and V (Helle,
1994).
Data on copepodite concentration from the Nor-
wegian Sea and the coast of Norway were assem-
bled from the literature. These data are synthesized
in equation (20):
TURBULENCE AND ICHTHYOPLANKTON 173
 N(t) = No e- (0.03 · t) (20)
where N(t) is the age-dependent (in days) cope-
podite concentration. No is the concentration of first-
feeding prey (nauplii stage III). These data were
used to calculate the prey separation distance as a
function of time.
Figure 26 shows the effect of wind-induced mix-
ing on the contact rate between the cod and the cope-
pod nauplii/copepodites, as the cod grows from larva
to juvenile. It shows the relative increase in contact
rate from first-feeding larvae to early juveniles for
three different wind speeds: 5 m s-1, 10 m s-1 and 15
m s-1, and for three different prey concentrations
(low, average, and high). The three prey concentra-
tions correspond to the low, average, and high con-
centration of equation (20). The lines converge
toward 1 when the turbulent velocity becomes much
less than the swimming speed of cod, and conse-
quently, no longer contributes to the contact rate
between the cod and its prey. The turbulence induced
contact rate is at its maximum for the slow-moving,
first-feeding cod larvae. The contact rate increases
by an order of eight for the situation of low prey con-
centration under a wind speed of 15 m s-1 compared
to non-turbulent conditions, while it increases by a
factor of five for high prey concentrations under the
same wind speed. Hence, the turbulence induced
contact rate contributes to reduce the effects of vari-
able prey concentrations, since the ratio between the
maximum and minimum prey concentration,
Nhigh/Nlow, is 63, while the change in concentration
experienced by the cod larvae (i.e. the change in the
number of encounters) is less: Zhigh/Zlow = 39. This is
due to the fact that at lower prey concentrations the
separation between prey particles becomes larger
and, consequently, larger turbulent cells with higher
kinetic energy contribute to increase the contact rate.
As the cod larvae grow and increase their swim-
ming speed, the influence of the turbulence-induced
contact rate decreases. However, the simulations
show that for strong mixing events the enhancement
of contact rate is still considerable for 2-mo-old
juveniles. At wind speeds of 15 m s-1 and at average
prey concentrations, the contact rate is higher by a
factor of 2.5 than it would be in non-turbulent con-
ditions. The prey concentration decreases as a func-
tion of time, and again, as mentioned in the para-
graph above, the increasing separation distance
between the prey particles contributes to maintain
the influence of turbulence as larger and larger
scales of turbulence come into effect.
174 S. SUNDBY
FIG. 26. – The relative increase (compared to non-turbulent condi-
tions) in the velocity component of the contact rate between Arcto-
Norwegian cod and its prey from first-feeding stage to 65 d after
hatching, for three wind speeds, 5 m s-1, 10 m s-1, and 15 m s-1, and
for three prey concentrations according to equation 5. Hatched
lines: Nlow = 0.8 e -0.03 t . Thick lines: Nav. = 6 e -0.03 t. Dotted lines:
Nhigh = 50 e -0.03 t .
CONCLUDING REMARKS
Previous opinions that turbulence generally has
detrimental effects on planktonic life is in opposi-
tion to theories and measurements developed over
the past two decades. The mixed layer is the larval
habitat for the major proportion of fish species. As
mentioned in the Introduction, light and turbulence
are the most predominant energy sources in this
part of the ocean. Organisms are likely to have
developed strategies to benefit from the effects of
these energy sources. MacKenzie et al. (1994)
modeled the optimal level of wind-induced turbu-
lence with respect to cod larvae encountering prey.
They found that the wind speed giving optimal tur-
bulence is probably at the level of about 15 m s-1.
The field results from Sundby et al. (1994) showed
that the optimal level must be higher than wind
speed of 10 m s-1, which is not inconsistent with
MacKenzie et al. (1994). This implies that the fish
larvae are able to benefit from quite strong turbu-
lent mixing. In the first-feeding areas of cod larvae
on the coast of northern Norway, the average
monthly wind speed (in May) is 5.6 m s-1 (Sundby,
1982), and wind speeds of 15 m s-1 or higher occur
less than 3% of the time. This leads to the conclu-
sion that detrimental effects of wind-induced tur-
bulence for pelagic larval cod encountering prey
rarely exist.
The models for vertical distribution of eggs show
that it is important to distinguish between stationary
distributions, where balance occurs between the
buoyancy forces and the vertical turbulent mixing,
and non-stationary distributions which are typical
for newly spawned eggs of less than 1 - 5 days old.
However, it is also important to realise that even a
stationary vertical distribution is not a static distrib-
ution on the individual particle level. The eggs are
continuously changing positions in relation to each
other at a rate depending on the level of turbulent
energy dissipation. For larval distributions, individ-
ual behaviour adds to the change of vertical posi-
tion. Sclafani et al. (1993) showed that the stochas-
tically vertical movement of larvae may result in a
sensed prey concentration for the individual larvae
which deviates considerably from the average lar-
vae. When there are large variations in the vertical
distribution of predator and prey, and particularly,
when peak concentrations do not overlap, the real
encounter rate between predator and prey will be
complicated to predict.
The topic of relevant turbulent scales for plank-
ton contact rates presently lacks a mathematical for-
mulation. In this presentation I have from physical
reasoning concluded that all turbulent “cells” of
length scales less than the separation distance
between predator and prey contribute to increase the
contact rate. This is because turbulent motion on
these length scales will continuously change the rel-
ative position between predator and prey. In the cal-
culations above based on larval cod from the field I
have, as an approximation, used an average concen-
tration of prey (i.e. Calanus finmarchicus nauplii) as
the basis for calculating the separation distance and,
hence, the relevant turbulent scale. Rothschild
(1988) stated that it is important to distinguish
between the deterministic separation distance, r, and
the probabilistic separation distance which is 0.55.r.
According to equation (17), the turbulence velocity
associated with the probabilistic separation distance
is 18% less than the turbulence velocity associated
with the deterministic separation distance. The level
of prey patchiness will influence whether the prob-
abilistic or deterministic distance is most relevant in
this context. However, the difference in the turbulent
velocities associated with these two scales is rather
small compared to the uncertainties for many of the
behavioural variables linked to the problem of
encounter rate between larvae and their prey.
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