ICES Journal of Marine Science, 53: 313–316.

1996

Acoustical investigation of phytoplankton

D. A. Selivanovsky, P. A. Stunzhas, and
I. N. Didenkulov

Selivanovsky, D. A., Stunzhas, P. A., and Didenkulov, I. N. 1996. Acoustical

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investigation of phytoplankton. – ICES Journal of Marine Science, 53: 313–316.

The existence of gas vacuoles and the dispersion of sound in water containing
phytoplankton is demonstrated under natural conditions by measurement of acoustic
reverberation and sound velocity, using phase, resonance, and other methods. The
volume of vacuoles ranges from 0.1–30% of the cell body volume. Two types of gas
cavities are observed: one is diminished elastically under compression, while the other
is decreased irreversibly. However, the volume is restored in a few hours with exposure
to light. The sound velocity has been measured over the frequency range 10–30 kHz, in
water containing phytoplankton cells at a volume concentration of 10 "4. The sound
velocity increased on average by 0.1% over this frequency range.

? 1996 International Council for the Exploration of the Sea

Key words: compression/decompression, gas cavities, phase and resonance methods,

reverberation, sound velocity dispersion, vacuoles phytoplankton.

D. A. Selivanovsky and I. N. Didenkulov: Institute of Applied Physics, RAS, 46 Ulyanov

St, Nizhny Novgorod 603600, Russia. P. A. Stunzhas: Shirshov Institute of Oceanology,
Krasikov St, 23, Moscow 117218, Russia. Correspondence to Selivanovsky [tel: +7 8312
384 584, fax: +7 8312 365 976].

Introduction acoustic reverberation properties; (2) static compressibil-

The acoustic significance of phytoplankton concen-
trations is well known. The possibility of remote study
of phytoplankton was proposed earlier when acoustic
instruments were first introduced into oceanography
(Cushing et al., 1956; Weston, 1958).
The existence of acoustically important elements such
as gas cavities in the cells was suggested by Bogorov
(1946) in discussing general concepts about the vital
activity of plankton. Clay and Medwin (1980) also
mentioned the possibility, based on the data of Watson
and Meister (1963). The fact that until now these
phenomena have not appeared as distinctly in blue-green
algae (Walsby, 1972) may be due to the earlier methods
of observation. For example, the size of gas vacuoles or
cavities in live cells may be too small to be resolved by
an optical microscope.
The ability of phytoplankton patches to cause sound
velocity to change with frequency, the phenomenon
known as dispersion, is also important, although this
effect has not been investigated so far.
Materials and methods
Attempts to identify gas cavities within phytoplankton
cells in water were undertaken by measuring: (1)
ity; (3) increase of the volume of the medium during
algal bloom; and (4) dispersion of sound velocity.
Experiments were carried out in the laboratory on
collections of cultures of marine algae and also in situ in
the tropical Atlantic.
1. Ultrasound reverberation measurements were done
at the same time as the compression measurements in
the expectation that the presence of gas cavities in
cells would markedly increase the level of reverber-
ation. The change in the size of cavities during
compression should also affect the level of the
reverberation.
Acoustical cross-sections for individual cells were
defined as the ratio of the volume scattering coeffi-
cient and the known concentration of cells (Sandler
et al., 1992).
2. Static compressibility was measured in an apparatus
which allowed the changes in volume of the medium
with and without algae to be compared during com-
pression. It is likely that a higher compressibility
of the medium containing algae could only occur
through the presence of gas cavities in the cells
(Sandler et al., 1992).
3. The increasing volume of the medium was measured
during the growth of the number of cells and related

1054–3139/96/020313+04 $18.00/0 ? 1996 International Council for the Exploration of the Sea
314 D. A. Selivanovsky et al.
to the increase of oxygen content. The aim of this
experiment was to compare the measured increase of
volume with the estimated volume increase which, as
a first approximation, may be determined from the
photosynthesis reaction whereby carbon-dioxide and
water are transformed into carbohydrate and oxygen.
The chemical formula for this reaction is
CO2 +H2O]CH2O+O2, from which the volume
increase may be calculated as described by Burlakova
et al. (1992).
4. The dispersion (variation with frequency) of the
sound velocity in media containing phytoplankton
must exist as a consequence of the presence of gas
cavities in the cells. It is known that the sound
velocity c decreases at frequencies below f0, the
resonance frequency of the gas bubbles, while for
frequencies above f0, the sound velocity increases.
We determined the velocities of sound at two
widely different frequencies (f1 =7.5 MHz and
f2 =0.75 MHz). One might expect that the ratio c
(f1)/c(f2) in a medium containing phytoplankton
would be more than in pure water.
The apparatus consisted of two sound velocity meters
with the same relative sensitivity, capable of detecting
proportional changes in velocity Äc/c>3#10 "6.
Changes in the velocity ratio were measured in water
containing monocultures of cells. In situ measurements
were made in the surface layer of water using the same
equipment, secured underwater to the side of the ship
travelling at a speed of 1–2 m s "1. In addition, as a
control check on the concentration of phytoplankton
and its biomass, the measurement procedure was sup-
plemented simultaneously by fluorimetry and by record-
ing dissolved oxygen from instruments positioned close
to the sound velocity meters.
These measurements showed that the sound velocity
variation in media containing plankton is more compli-
cated than previously thought on the assumption that it
was caused entirely by the presence of gas cavities in the
cells (Sandler et al., 1993). We therefore conducted a
detailed investigation of this phenomenon using the
phase and resonance methods over a wide frequency
range (200 Hz–11 MHz).
The phase method was performed in vessels having
the form of glass tubes with acoustic transducers at
the ends. The largest tube (for sound frequencies
7–250 kHz) had a length of L=760 mm and a diameter
of 210 mm. Smaller tubes were used for frequencies
75–2500 kHz (L=357 mm) and 1–11 MHz (L=28 mm).
The frequencies at which the emitted and the received
signals had the same phases were recorded. The
frequency detuning (Äf) is related to the sound velo-
city as c=Äf.L. Thus the velocity ratio is cmedia/
cwater =Äfmedia/Äfwater averaged over the frequency
range.
The resonance measurements were made in vertical
glass tubes with an elastic bottom, and a fixed height L
of the liquid volume which was kept constant within
&0.01 mm, using a tightly stretched polymer film to
displace the excess liquid. The sound velocity was esti-
mated as c=ëfres, where fres is the resonance frequency
of the liquid column and ë is the wavelength. The system
could be tuned to resonance with the sound pressure
node at the bottom and the particle velocity node at the
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polymer film. There were several resonant modes near
the frequencies corresponding to L~ë/4, 3ë/4, 5ë/4, etc.
There were additional resonances at the harmonics
(L~ë/2, ë) and the subharmonics (L~ë/8, ë/12) caused
by the finite impedances of the liquid column bound-
aries. The quality (Q-factors) of these resonances was
such that the accuracy of measuring the ratio cmedia/
cwater was about one part in 10 "5. Resonator tubes with
L=640 mm and 320 mm were used.
Results
1. Methods 1, 2, and 3 all indicated that gas cavities
exist in the cells of all the species we examined.
2. Measurements of the sound velocity dispersion show
that, together with the known acoustic effect of
increased absorption of sound, the presence of
phytoplankton causes complex changes in the sound
velocity.
(a) In the course of the reverberation measurements,
it was established that the acoustic cross-section
was one to two orders of magnitude more than
that expected from the cell tissue alone. Calcula-
tions show that, in general, the cell tissue is 10%
denser than water, and its compressibility is neg-
ligibly different from water (Urick, 1975). The
additional energy backscattered by the plankton
was attributed to the presence of gas cavities in
the cells. The sizes of the gas cavities were estab-
lished on the assumption that they are free gas
bubbles having a resonance with a quality factor
(Q) of about five.
When applying compression with an additional
pressure of 0.5 atm, it was found that the back-
scattering decreased irreversibly with some
species of cells. After compression, the measured
acoustic cross-sections of such cells became close
to the calculated values for the cell tissue. Over
some time (a few tens of minutes to hours) the
backscattering strength was restored, but it could
be reduced again by further compression.
For other types of cells, the scattering changed
elastically with the application of pressures up to
5 atm, and their measured acoustic cross-sections
returned to the original values as soon as the
pressure was removed.
 Acoustical investigation of phytoplankton
Table 1. Volumes (ìm3) of cells and gas cavities in the medium with some algae, measured by different
methods.
Volume
Species of alga of cells
Prorocentrum micans 1.4#105
Peridinium triquetrum 5.0#105
Olistodiscus luteus 600
Dunaliella salina 150
Platimous viridis 250
Phaeodactilum tricornutum 20
(b) By measuring the compressibility, we showed that
the cells which recovered elastically, as the
applied pressure was removed, obeyed Boyle’s
law (PV=constant). However, in the case of the
gas cavities in cells whose volume decreased irre-
versibly, these volumes were effectively missing
from the medium after removal of excess pres-
sure. Media containing these species of cells had a
compressibility differing very little from that of
water.
(c) Measurements in a medium containing an algal
bloom showed that the volume always increased
by more than the calculated quantity of new cell
tissue. This difference was ascribed to the forma-
tion of gas vacuoles connected with the growing
cells.
(d) It is suggested that measuring the sound velocity
at two frequencies provides another means of
determining the properties of gas cavities in the
cells. However, the anticipated effect (along
with its disappearance under compression) was
observed in only a few species of algae.
Field measurements in the sea showed that, in
phytoplankton concentrations, the sound velocity
ratio between frequencies f1 and f2 could be
smaller, larger, or equal to unity. However, in the
laboratory experiments an increase in sound
velocity was observed at both frequencies in
media containing phytoplankton. This effect was
proportional to the cube root of the cell concen-
tration. So far we have not ascertained the nature
of this phenomenon.
Phase and resonance measurement methods
showed a complex frequency dependence of the
sound velocity. The individual measurements are
fairly variable, but the average increase of sound
velocity is about 0.1% for frequencies of
10–30 kHz. Whenever the sound velocity for a
particular species of cells is below that in pure
water, the presence of gas cavities is indicated
since compression of the media increases the
315
Volume of gas cavities
Scattering Compression Volumetry
0.6–1.2 5000 100–600
0.2–0.9 5–10 7–100
0.2–0.4 30–70 10–60
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0.1–0.4 5–100 10–30
0.04–0.4 10–40 3–15
0.01–0.06 2–10 2–8
sound velocity which returns almost to that of
pure water.
By these methods, the mechanical and acoustical
properties of the media were measured for 32 species of
phytoplankton. We conclude that the results may be
explained by (1) the presence of gas cavities (vacuoles) in
(or near) the phytocells; and (2) the effect of the cells on
the sound velocity in the surrounding medium.
The volumes of gas in the same cells measured by the
various methods were very different (see Table 1).
Clearly, some or all of the models used for the analysis
were inadequate. However, our results show that while
the gas volume increases with the cell size, the relative
gas volume in small cells is greater than in large cells.
This is not consistent with the hypothesis that the basic
function of the gas in phytoplankton cells is to maintain
neutral buoyancy.
The existence of gas cavities in phytoplankton may
have many consequences. As an example, we note that
sound velocity dispersion can influence the propagation
of acoustic waves through regions occupied by phyto-
plankton. Furthermore, the presence of gas cavities
considerably increases the target strength of cells so that
phytoplankton concentrations in the ocean may be
observed acoustically.
References
Bogorov, V. G. 1946. The underwater world (in Russian).
Association of State Publishing House, Moscow, Leningrad.
48 pp.
Burlakova, Z. P., Krupatkina, D. K., Sandler, B. M.,
Selivanosky, D. A., and Stunzhas, P. A. 1992. Gas bubbles
and plankton. Volumetric measurements. Okeanologiya (in
Russian), 32: 481–485.
Clay, C. S. and Medwin, H. 1980. Acoustical oceanography:
principles and applications. A Wiley-Interscience Publica-
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Cushing, D. H., Lee, A. J., and Richardson, J. D. 1956.
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Sandler, B. M., Selivanovsky, D. A., Stunzhas, P. A., and
Krupatkins, D. K. 1992. Gas bubbles and sea phyto-
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plankton. Reverberation ultrasonic measurements. Oceanol-
ogiya (in Russian), 32: 92–100.
Sandler, B. M., Selivanovsky, D. A., and Stunzhas, P. A. 1993.
Sound velocity in media containing sea phytoplankton.
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