ISSN 1063-0740, Russian Journal of Marine Biology, 2007, Vol. 33, No. 6, pp. 417424. Pleiades Publishing, Ltd.

., 2007.
Original Russian Text B.E. Anninsky, G.A. Finenko, G.I. Abolmasova, Z.A. Romanova, 2007, published in Biologiya Morya.

ECOLOGICAL
PHYSIOLOGY

Somatic Organic Content of the Ctenophores Mnemiopsis leidyi

(Ctenophora: Lobata) and Beroe ovata (Ctenophora: Beroida)
in Early Ontogenetic Stages
B. E. Anninsky, G. A. Finenko, G. I. Abolmasova, and Z. A. Romanova
Institute of Biology of Southern Seas, National Academy of Sciences of the Ukraine,
Sevastopol, 99011 the Ukraine
e-mail: anninsky@ibss.iuf.net
Received May 17, 2007

AbstractThe organic matter content in the eggs and early larvae of the ctenophores Mnemiopsis leidyi and
Beroe ovata from the Black Sea was determined using the adapted microtechnique of dichromate oxidation.
The content of organic matter in the eggs of M. leidyi (0.25 g/indiv.) was 5 times less than in B. ovata (1.28
0.29 g/indiv.). The somatic content of organic matter was 0.25 0.09 g/indiv. (25.1 8.3 g/mg, wet wt) for
2-day-old larvae of M. leidyi (0.20.3 mm in body length) and 1.37 0.19 g/indiv. (67.1 5.7 g/mg wet wt)
for larvae of B. ovata (0.4 mm in body length). The specific organic content of larvae of both species steadily
decreased with an increase in ctenophore body size and weight, approaching 34 g/mg of wet weight for 2 mm
specimens of M. leidyi and 35 g/mg of wet weight for 6 mm B. ovata. The specific organic content of early
larvae was 2030 times higher than that of adult ctenophores. The results of this investigation could be useful
in the evaluation of the energy budget for somatic growth and generative production in these species. Calcula-
tions indicate that with specific wet weight growth rates of 0.43/day for M. leidyi larvae and 0.29/day for
B. ovata larvae, their true organic increases are respectively 30 and 38% less, i.e., no more than 0.31/day for the
former and 0.18/day for the latter species.

Key words: Organic matter content, growth, eggs, larvae, gelatinous plankton, Ctenophora, Mnemiopsis leidyi,
Beroe ovata.
DOI: 10.1134/S1063074007060090

Planktonic ctenophores are predatory gelatinous
invertebrates, which occur abundantly in the Black,
Azov, Caspian (from 1999), and other seas. Expansions
of ctenophores quite often give rise to changes in the
structure of biological communities. This is most viv-
idly exemplified by the naturalization of the cteno-
phores Mnemiopsis leidyi (from 1982) and Beroe ovata
(from 1999) in the Black Sea [4, 11, 13, 22, 32]. As a
result of high fecundity, individual differences in
growth rates, and the ability to reproduce as early as the
larval stage, the ctenophores size composition in natu-
ral populations is, as a rule, extremely wide. Without
doubt, such a population structure is important for the
existence of these animals, because it is beneficial for
their survival in conditions of food deficiency. How-
ever, many ecological studies assessing gelatinous
predators, in particular, in terms of their energy budget
do not take into account the ecological and physiologi-
cal features of the early larval stages of ctenophores.
It is common practice for the physiological data
obtained for the older age groups to be approximated
for the larvae. The ecological bioenergetics of larval
ctenophores have been determined in rare cases [24, 29,
33, 34].
This is largely due to methodological differences.
Because of the microscopic size and vulnerability of
the ctenophores eggs and larvae, it is hardly possible to
isolate them in the intact form, and when chemical
methods are used, the sea water and salts introduced
destroy the samples. As a consequence, repeated
attempts to determine the organic content of ctenophore
eggs and larvae smaller than 1 mm have proven to be
unsuccessful, despite modern technical means (e.g., the
CHN analyzer) [29]. Since such data were lacking, it
has been impossible to investigate the dynamics of the
organic matter content in the ontogenesis of cteno-
phores in detail and to estimate, in energy values, their
generative production and early somatic growth. At the
population level, underestimation of the specific
organic matter content variation in ontogenesis of
ctenophores can lead to the erroneous determination of
organic matter accumulated by the whole population
[29].
The purpose of the present research was to deter-
mine the organic matter content in the eggs and larvae
of ctenophores using the adapted microtechnique of
dichromate oxidation. In this regard, this work is also of
pure methodological interest because it analyzes

417
418
Body mass, mg
.
100 Y = 0.202 X 2.67
. Y = 0.111 X + 0.042
r2 = 0.99; n = 17
.
. . .
.
10
. 0.2
1
0.1
1 10
Body length, mm
Fig. 1. Relationship between body length and weight of
Beroe ovata larvae estimated by the weight method (white
circles) and by the volume method (black circles).
another technique for the determination of the organic
matter content (in wet weight) of gelatinous organisms.
It is well known that other approaches to solving this
problem (analysis of Corg, calorific value, and organic
composition) can yield, for a number of reasons, values
for organic matter content that are not always accurate
[18]. Using two species of ctenophores as an example,
we determine, for the first time, the contribution of the
so-called hydration component (caused by an increase
in the specific water content of the body) and the
organic component of the growth rate of early larvae
artificially grown in conditions of an initially abundant
food supply.
MATERIAL AND METHODS
Investigations on two species of ctenophores (Mne-
miopsis leidyi Agassiz A., 1865 and Beroe ovata Mayer
1912) were carried out near Sinop (Turkey) and at the
coast of the Crimea, at periods of the highest abundance
of these species in the plankton of the Black Sea (July
August 2003 and SeptemberOctober 2004). Eggs and
larvae were obtained from ctenophores that were main-
tained and reproduced in the laboratory. In addition,
large larvae were collected from net hauls.
Larvae and eggs were measured to the nearest
0.01 mm under a microscope. Their weight was deter-
mined by calculation, using the standard relationships
between the length and volume of regular geometric
bodies (spheres, cylinders, and ellipsoids). The density
of organisms in the calculations was taken to be 1 mg/l.
The volume of eggs was calculated as the volume of a
sphere (M. leidyi) or an ellipsoid (B. ovata); the volume
of larvae was determined as the volume of a cylinder
obtained by squeezing it between glass slides 0.2 mm
apart [20]. A special study of large larvae of B. ovata
RUSSIAN JOURNAL OF MARINE BIOLOGY
ANNINSKY et al.
Optical density, D
0.5
0.4 r 2 = 0.96; n = 19
0.3
0.1
0 1 2 3 4
Salt, mg
Fig. 2. Optical density of solution at dichromate oxidation
of the dry residue of sea water samples (25 l) with different
content of salts.
showed that weight of specimens determined by the
volume method or by weighing on a microanalytical
balance (with an accuracy of 0.05 mg) differed insignif-
icantly (p > 0.05) (Fig. 1). As well, the volume method
was the only appropriate method for calculating the
weight of eggs and early larvae.
Eggs and early larvae were concentrated in approx-
imately 1 ml of water by filtering through fine gauze
(100 m) and subsequently removing excess water with
microcapillary pipettes under a microscope at low mag-
nification. The final concentration of eggs and larvae in
the concentrated sample was 5001000 indiv./ml.
Twenty-five milliliters of the sample (sea water +
organisms) was taken with a dispenser and placed in
small glass vials. In the analysis of early larvae, one
sample contained 2035 individuals of M. leidyi 0.2
0.3 mm long or 1020 specimens of B. ovata 0.43 mm
long. In each sample, there were 35 larvae of M. leidyi
0.50.8 mm long. Larger larvae (M. leidyi, 12 mm;
B. ovata, 16 mm) were analyzed individually. The con-
trols were samples of sea water of equal volume (25 l)
in 35 replicates. All samples were dried in a drying
chamber at 100C for subsequent analysis. In all,
25 samples of M. leidyi larvae, 39 samples of B. ovata
larvae, and 20 samples of B. ovata eggs (545 in each
sample) were analyzed.
The organic matter content of ctenophores eggs and
larvae was determined using the microtechnique of
dichromate oxidation, as recommended for lipid analy-
sis [14, 18]. This method is very sensitive (~1 g OM);
however, in the case of highly mineralized samples, its
use is constrained because the salts distort the results
and lead to erroneous estimates (Fig. 2).
To avoid this, the salt effects are traditionally elimi-
nated, but we tried to create a uniform salinity back-
ground for all samples. Such a background was
Vol. 33 No. 6 2007
 SOMATIC ORGANIC CONTENT OF THE CTENOPHORES
achieved by strict standardization of the volume of liq-
uid samples (25 l) containing either sea water (con-
trol) or the investigated materialeggs and larvae
(or 50200 l of the tissue homogenate of large cteno-
phores)in sea water of the same salinity (in the range
of 17.618.2). The ctenophores are typical osmocon-
formers, and their eggs and larvae occupy a small vol-
ume of sample; consequently, negligible differences in
the degree of mineralization of the investigated samples
can be neglected if the salinity background is taken to
be the same as in an equal volume of sea water. One ml
of dichromate (8.5 mM K2Cr2O7 in 36 N H2SO4) was
added to vials containing the investigated samples and
controls and the vials were heated for 30 min in a dry-
ing chamber (150C). Samples were oxidized and
cooled to room temperature. Aliquots of the dichromate
solution (0.2 ml, 3 replicas) were diluted with 5 mls of
distilled water and the final change in the optical den-
sity of the solution was recorded on a spectrophotome-
ter (SF 46) at a wave length of 350 nm.
Protein in the ctenophore body is the dominant oxi-
dizable substrate, and makes up 8085% of the total
organic matter [15, 18]. Since it has an intermediate
degree of oxidability between that of carbohydrates and
lipids, the calibration curve was plotted from albumin
(human serum albumin, HSA, puriss.), which was also
dissolved in sea water of standard salinity (Fig. 3).
In calculations of the organic matter content from the
total chemical composition, only the protein content
was analyzed, using albumin as the standard [23]. The
total organic matter content of M. leidyi was calculated
from these data, the proportion of protein in the organic
matter of ctenophores was taken to be 85% [1].
For evaluation of the individual growth of M. leidyi,
larvae of this species were obtained in the laboratory
from eggs and kept in 20-l aquaria at a temperature of
2628C and a salinity of 18.2. The water in the
aquaria was renewed daily by 75%, simultaneously
supplying some microzooplankton (small orthoplutei
of Copepoda, rotifers, infusorians) that pass through a
gauze material with a mesh size of 100 m. The initial
concentration of ctenophore larvae in the aquaria was
about 50 indiv./l. In 1 to 2 days, they were examined
under a microscope, and the size of the largest speci-
mens (the fastest growing larvae) was recorded. After
that, all larvae were returned. The observation period
was 12 days.
To assess the growth rate in a group of M. leidyi
specimens, another series of experiments was con-
ducted on 2-day-old larvae. Preliminarily measured
specimens of the same size (0.20.3 mm) were placed
in three 6-liter jars with fresh sea water (10 larvae in
each) and maintained at the same temperature (26C)
and salinity (18.2). Simultaneously with the larvae,
microzooplankton filtered through a gauze of varying
mesh size was added to the jars. The mesh size was
100 m for the first jar, 50 m for the second, and
30 m for the third. The initial concentration of the
RUSSIAN JOURNAL OF MARINE BIOLOGY Vol. 33
419
Optical density, D
0.130
0.125 Y = 0.00195 X + 0.082
r2 = 0.94; n = 17
0.120
0.115
0.110
0.105
0.100
0.095
0.090
0.085
0.080
0 2 4 6 8 10 12 14 16 18 20
Albumin, g
Fig. 3. Optical density of the solution during dichromate
oxidation of the dry residue of sea water samples (25 l,
salinity 17.6) with different content of albumin (HSA).
largest component of microzooplanktonthe copepod
orthonauplii (L 0.1 mm)in the jars was 50, 45, and
5 indiv./liter, respectively. After 5 days of rest, all larvae
were measured, and subsequently measurements were
made every 2 days; the experiment continued for
9 days. No death of larvae was recorded over this period.
The individual growth of B. ovata was investigated
using 2-day-old larvae that developed from eggs
spawned by the ctenophores under laboratory condi-
tions. Larvae were kept in 5-l jars at a temperature of
1922C and a salinity of 18. The initial concentra-
tion of larvae was 1015 indiv./l. The water in the jars
was renewed by 3050% at 35 day intervals. Simulta-
neously, larvae were excessively fed on juveniles or
large fragments of adult M. leidyi. As a rule, this ration
was sufficient until the next portion was offered. However,
in some cases, the larvae were fasting for 1 to 2 days. Lar-
vae were examined and measured every 35 days.
Because of the rapid development of heterotrophic
microflora caused by the introduction of M. leidyi frag-
ments into the jars and because of damage to the larvae
through water collection, larval mortality was marked.
Over the 30 days of the experiment, larval numbers
decreased by 95%.
RESULTS
Correspondence of Data from Alternative Methods
The organic matter content in one specimen of Mne-
miopsis leidyi of medium size (L = 30 mm) was deter-
mined using two independent techniques: (1) dichro-
mate oxidation and (2) proximate chemical composi-
tion. The data from the two methods were comparable,
based on the assumption that not all organic com-
pounds would be fully oxidized by dichromate [6] and
a correction factor would be needed to convert from the
No. 6 2007
420 ANNINSKY et al.

Table 1. The organic matter content of the eggs and early larvae of the ctenophores Mnemiopsis leidyi and Beroe ovata from
the Black Sea

Body wet weight, Organic matter

Species L, mm n
mg g/indiv. g/mg of wet weight
Mnemiopsis leidyi
eggs 0.40 0.05 0.034 0.014 0.25* 7.4*
larvae 0.26 0.03 0.010 0.003 0.25 0.09 25.1 8.3 5**
Beroe ovata
eggs 0.85 0.07 0.322 0.063 1.28 0.29 4.0 0.8 20
larvae 0.43 0.01 0.020 0.001 1.37 0.19 67.1 5.7 16
Notes: is the standard deviation.
* Calculated from the organic matter content of early larvae.
** Each point is the mean of 510 independent determinations.

values obtained to the actual content of organic matter.
However, the results of concurrent analysis of the
somatic homogenate of one and the same ctenophore
did not differ significantly (n = 10; p > 0.05): the values
of the organic matter content of this ctenophore deter-
mined using the method of dichromate oxidation and
using the proximate chemical composition were 1.148
0.100 and 1.132 0.087 mg/g of wet weight, respec-
tively. Thus, either the two methods are equally appro-
priate for evaluating the organic matter content of
ctenophores or the errors of the methods have the same
vector of change. The latter is unlikely because results
from alternative methods for determining organic mat-
ter content in the ctenophores (analysis of Corg, calorific
value, and chemical composition) are usually very
comparable [16, 18, 31].
Specific OM, g per mg wet weight
Y = 28.8 X1.02
r2 = 0.94; n = 39
20
2 organic matter content of the larvae changes many-fold:
1 25.1 8.3 to 4.2 0.7 g/mg, wet weight; for B. ovata
Y = 6.97 X0.85; r 2 = 0.81; n = 25
2 4.2 0.3 g/mg, wet weight (Fig. 4).
0 1 2 3 4 5 6
Body length
Fig. 4. Variation of specific organic matter content (organic
matter, g/mg of wet weight) in relation to larvae length of
the ctenophores Mnemiopsis leidyi (1) and Beroe ovata (2).
Analysis of Eggs and Larvae
We failed to determine the organic matter content in
the eggs of M. leidyi in the strictly analytical sense.
Since the eggs spawned at night develop quickly (at 26
27C), many embryos in the egg masses examined dur-
ing the day had reached the stage of larva. Therefore, it
was virtually impossible to sample eggs from these
generations in the quantity required for analysis. Nev-
ertheless, as the differences in organic matter content
between eggs and early larvae are minimal (p > 0.05 for
Beroe ovata), then, with allowance made for negligible
error, the organic content of the eggs of M. leidyi can be
inferred from analysis of the larvae. By our calcula-
tions, the absolute organic matter content in the eggs of
M. leidyi approaches 0.25 g, which is 5 times less than
in the eggs of B. ovata. On the other hand, the relative
organic matter content was higher in the eggs of
M. leidyi (Table 1). This is due to the different hydra-
tion of the eggs: the developing larvae of M. leidyi
occupy about 30% of the egg volume, while in B. ovata,
this figure is less than 5%.
The larvae of B. ovata are almost two times larger
than the larvae of M. leidyi, and have a 2 times greater
wet weight, approximately 5 times higher absolute
organic matter content, and 2.5 times greater organic
matter content per milligram of wet weight (Table 1).
The last relationship remains the same until larvae of
B. ovata attain a size of 25 mm and above. The specific
for M. leidyi in the size range of 0.262.00 mm, from
in the size range of 0.436.02 mm, from 67.1 5.7 to
Growth
Larvae of M. leidyi obtained from the same egg
mass and maintained under the same conditions grew
and developed at different rates. As a consequence, the

RUSSIAN JOURNAL OF MARINE BIOLOGY Vol. 33 No. 6 2007

 SOMATIC ORGANIC CONTENT OF THE CTENOPHORES 421

Wet body mass, mg/indiv.

10 Y = 0.0032e 0.70X (a) Y = 0.095e 0.50X (b)
r2 = 0.95; n = 20 20 r2 = 0.95; n = 20
1

2
0.1

0.01 Y = 0.004e 0.44X;

0.2 Y = 0.12e 0.31X;
r2 = 0.64; n = 110 r2 = 0.62; n = 110
0.001
0 2 4 6 8 10 12 0 2 4 6 8 10 12

1000 Y = 0.02e 0.37X (c) Y = 1.39e 0.23X (d)

1000
100 r2 = 0.91; n = 24 r2 = 0.91; n = 24

10 100

1 10
0.1
1
0.01 Y = 0.006e 0.29X Y = 0.67e 0.18X
r2 = 0.57; n = 129 r2 = 0.57; n = 129
0.1
0 5 10 15 20 25 30 0 5 10 15 20 25 30
Days

Fig. 5. Dynamics of body wet weight (a, c) and organic matter content (b, d) in the course of larvae growth of ctenophores Mnemi-
opsis leidyi (a, b) and Beroe ovata (c, d) under experimental conditions. Black symbolsthe growth of the fastest growing larvae,
white symbolsthe growth of grouped specimens.

size of 7-day-old larvae differed by as much as two
times (0.450.90 mm) and body weight, by almost an
order of magnitude. We examined two characteristics
of growth: the growth rate of the fastest-growing larvae
and the average growth rate of larvae in a group (Fig. 5).
We assumed that the fastest-growing larvae are those
retaining the leading position in the process of develop-
ment and growth; in actuality, however, the possibility
of the alternation of these larvae cannot be ruled out.
In both cases, growth of larvae was described by an
exponential model, W = W0ekt, where W0 and W are the
initial and final body weight; k is the specific growth
rate; and t, the time (in days). Along with exponential
growth of body weight over time, such a relationship
assumes a proportional relationship between the
increase in weight and the existing weight, as well as
the constancy of the specific growth rate (k). These con-
ditions [7] are statistically appropriate for determining
the initial growth of gelatinous organisms, as supported
by our data in the case of the ctenophores (r2 = 0.62
0.95 for M. leidyi, r2 = 0.570.91 for B. ovata). In equa-
tions calculated for M. leidyi larvae, the values of the
index of the specific growth rate of body wet weight (k)
were 0.70 and 0.44, respectively, for the fastest-grow-
ing larvae and for the grouped specimens. The k values
in analogous relationships for somatic organic matter
content were 29 30% less (0.50 for the fastest-growing
larvae and 0.31 for grouped specimens). A detailed
analysis of the growth of M. leidyi larvae under differ-
ent feeding conditions was not a high- priority goal of
our study. We, however, note that during the first 7 days
larvae in the three experimental jars grew at a similar
rate, while between days 9 and 10, the growth rate of
larvae in the jar with the lowest concentration of cope-
pod orthonauplii was markedly lower (p < 0.05) than in
the jar with the greatest initial concentration of copepod
orthonauplii (Fig. 6). It is not clear which accounted for
this retardation in the growth of larvae, the qualitative
composition of food (predominance of infusorians and
rotifers) or an insufficient amount of food.
The growth rate of B. ovata larvae was 1.52 times
lower than in M. leidyi larvae. The indices of the spe-
cific daily increase in the wet weight of the fastest-
growing larvae (k = 0.37) and the grouped larvae (k =
0.29) exceeded the corresponding values (k = 0.23 and
k = 0.18) for the increase in organic matter content
by 38%.
DISCUSSION
Reliable values of somatic organic matter content
were previously determined for larvae of Mnemiopsis
leidyi at a minimum size of about 1.1 mm. According to
Reeve et al. [29], the Corg content of larvae is 13.1% of
body dry weight, and in oceanic conditions (34) it is
0.45% of wet weight. Our results show (Fig. 4) that the
organic matter content of larvae of this size can be up to
0.64%, and that Corg (organic matter/1.9) is 0.34% of the

RUSSIAN JOURNAL OF MARINE BIOLOGY Vol. 33 No. 6 2007

422 ANNINSKY et al.

Body length, mm It might be assumed that a high organic matter content

2.0 in larvae is necessary for the successful completion of
1 ontogenesis in other species of gelatinous organisms as
1.6 2 well. Initial storage of organic matter is required, not
only for the survival of larvae, but also during the sub-
3 sequent role of their competitive strategy, i.e., provid-
1.2 ing a maximal rate of growth at a minimal rate of
metabolism [16]. With increasing hydration of the body
0.8 during the development of these organisms, a decrease
in the capacity for movement and maneuvering is coun-
terbalanced by a large number of valuable qualities,
0.4
among them lower susceptibility to predators and ecto-
parasites, a larger size range for food organisms, a
0 greater range of capture of food organisms, rational uti-
7 9 11
Days lization of food energy as a result of a low active metab-
olism and the absence of expenditures on the formation
Fig. 6. Dynamics of length of Mnemiopsis leidyi larvae
of a skeleton [27]. Regarding the exponential character
growing under experimental conditions at different concentra- of ctenophores growth, its rate in M. leidyi (k = 0.77
tions of copepod orthonauplii (150 indiv./l, 245 indiv./l, and k = 0.43, respectively for the fastest-growing larvae
35 indiv./l). Vertical linesstandard deviation (). and for grouped larvae) allows the fastest-growing lar-
vae to double their body weight during a single day.
Analogous data were obtained experimentally for
wet weight. In Reeves work, the hypothetical content of M. leidyi from the Gulf of Biscay in the West Atlantic
Corg in the eggs of M. leidyi was taken to be 0.1 g/indiv. (k = 0.8, k = 0.69, and k = 0.4, respectively for the fast-
and in early larvae, was taken as approximately 20% of est-growing larvae, fast-growing juveniles, and nor-
dry weight (or 0.68% of wet wt). Our data show mally-growing juveniles) [30]. Under less favorable
(Table 1) that the Corg content in the eggs of M. leidyi is conditions, the growth rate of this and other ctenophore
0.13 g/indiv. and in early larvae, 1.32% of the wet species is lower and most often limited to k = 0.20.4
weight. It is well known that Corg in adult M. leidyi is [8, 10, 29]. In this regard, it is not clear to what extent
usually not above 0.065% of the wet weight [15, 16, the lower indices of the specific growth rate of B. ovata
29]. Thus, the results of the independent studies are larvae are species specific and to what extent they are
comparable and indicate more than 20-fold differences the consequence of conditions of maintenance that are
in the specific organic matter content between larvae not so favorable as for M. leidyi. Possibly because of
and adults of this ctenophore. This holds true for Beroe the lower temperature and irregularities in the food sup-
ovata, in which Corg in adult specimens was found to be ply, the specific growth rate of even fast-growing larvae
0.070.21% [25], 0.15% [4] or 0.13 0.03% [15]. of B. ovata did not reach maximum values. This may
also be suggested by the fact that the specific growth
The early larvae of ctenophores are very close to rate of B. ovata in Sevastopol Bay during the periods of
nongelatinous plankton in their Corg content, (Table 2). their seasonal expansion was 0.330.67/day [22].
Our results indicate that the large increase in the wet
Table 2. Content of Corg (% of wet weight) in ctenophores weight of larvae of the two ctenophore species was
and selected nongelatinous representatives of zooplankton largely (by 30% in M. leidyi and by 38% in B. ovata)
due to the intensive accumulation of water in the body.
Organism Corg, % Source This is probably a consequence of the predominant
Mnemiopsis leidyi 0.040.06 [15, 16, 29] development of the mesogloeal matrix, which is excep-
tionally hydrated but poorer in organic matter than
Beroe ovata 0.070.21 [4, 15, 25] other tissues (muscles and body coverings, etc.) [29].
Larvae of: If the so-called hydration constituent is subtracted, the
M. leidyi 1.3 Our data maximum values of k for larvae of M. leidyi and
B. ovata 3.5 Our data B. ovata will decrease to 0.50 and 0.23, respectively.
Under favorable feeding and temperature conditions,
fish 4.65.9 [17] such values are also characteristic of other representa-
Copepods 4.914.2 [16, 17, 28] tives of gelatinous plankton, for example jellyfish, cha-
Euphausiids 6.49.8 [19, 28] etognaths, and appendicularians [8]. During the initial
Amphipods 7.112.9 [28] period of growth, body hydration in the cteneophores is
likely to be more active, compared to other species of
Decapods 7.613.1 [16] gelatinous organisms.
Pteropods 2.98.9 [19, 28] It would be interesting to clarify whether or not the
Polychaetes 2.67.2 [17] high values of growth efficiency indices occasionally

RUSSIAN JOURNAL OF MARINE BIOLOGY Vol. 33 No. 6 2007

 SOMATIC ORGANIC CONTENT OF THE CTENOPHORES
observed in ctenophores (K2 = 0.60.8) [4, 26] are
related to the underestimation of changes in the body
hydration of growing individuals. Since data on the res-
piration of M. leidyi larvae were unavailable, such an
analysis was possible only for B. ovata. Calculations
showed that with a body weight of 1.37 g, an organic
matter and calorific value of 5.96 cal/mg, the daily
increase of B. ovata larvae would be 0.00188 and
0.00147 cal/indiv., respectively for the fastest growing
larvae and for grouped larvae [15]. Their daily meta-
bolic requirements at the same temperature (~20C)
can be estimated to be 0.00057 cal/indiv. (0.168 g
O2/indiv.) [34], i.e., the growth efficiency of B. ovata
larvae would probably be up to 77 and 72%. These val-
ues, as well as the above values of K2, are close to the
maximum values of growth efficiency for aquatic inver-
tebrates (K2 = 0.70.8) [3]. This could be connected, in
some measure, with the underestimation in ctenophores
of such components of metabolism as the specific
dynamic action of food [25] and anaerobic metabo-
lism [12]. At the same time, it is precisely the juvenile
stages of invertebrates, irrespective of their taxonomic
position, that are capable of growth with a K2 0.8 [3].
Body hydration accounts well for several other
physiological phenomena observed in ctenophore lar-
vae. In particular, respiration experiments with B. ovata
demonstrated that the rate of oxygen consumption by
newly hatched larvae was at least 20 times greater than
in adult specimens [34]. These authors believe that such
a difference in respiration rate could not be due to a
decrease in the specific organic matter content in grow-
ing larvae, because if this were the case, the Corg content
in early larvae would have an absolutely unrealistic
value of 3.2% wet weight. However, according to our
data, the content of Corg (Worg/Wwet/1.9 100%) in early
larvae of B. ovata can reach the even greater value of
3.5% of wet weight. Consequently, the differences in
respiration rate of different-size specimens of B. ovata
might largely be due to different content of metaboli-
cally active organic matter in their body. An analogous
conclusion is also true for the different specific ration
calculated from larvae respiration rates [34]. Values of
organic matter content in the ctenophores eggs are
undoubtedly important for the estimation of the energy
budget of generative production. With a good food sup-
ply, production of eggs in M. leidyi with a body Corg
content of about 15 mg (wet weight about 25 g, length
about 82 mm) can be 1600 [11] and even 3000 eggs per
day [29]. In this case, with an egg Corg content of
0.13 g/indiv., the share of generative production
respectively is 1.4 and 2.6% of the somatic Corg. With a
spawn of 8000 eggs [9], organic losses in these cteno-
phores can make up 6.9% of the somatic Corg. The daily
metabolic expenditures in ctenophores with a body
weight of 25 g at 26C are estimated to be 5.6% of the
somatic Corg [2]. Thus, despite the fact that the hypo-
thetical Corg content in eggs of M. leidyi (0.1 g/indiv.)
was earlier taken to be close to the real content [29], it
is difficult to agree with the statement that the contribu-
RUSSIAN JOURNAL OF MARINE BIOLOGY Vol. 33
423
tion of generative production to the total energy budget
of this species is negligible.
In sexually mature specimens of B. ovata, organic
matter losses via sex products are apparently even more
marked. At the period of active reproduction in the area
of Sevastopol Bay in 2003, the ctenophores spawned on
average 4500 250 eggs per day [11]. With an average
length of spawning specimens of about 53 mm (35
70 mm), their energy expenditures via sex products are
estimated to be 14.9% of their organic body weight
[15]. The daily metabolic requirements of B. ovata are
higher than in M. leidyi, and with the same size of spec-
imens and a weight of 15.4 g, they can reach 6 to 8% of
the organic matter at 1921C and 11 to 15% of the
organic matter at 2526C [5, 21, 32, 34]. Hence,
despite a higher level of metabolism than in M. leidyi,
losses via sex products in B. ovata are quite comparable
with respiratory expenses. This may be indicative of the
predominance of the generative strategy of metabolism
and the concurrent retardation of growth in B. ovata
during the reproductive period.
ACKNOWLEDGMENTS
We are deeply grateful to Dr. M.V. Propp (Institute
of Marine Biology, Far East Division, Russian Acad-
emy of Sciences) and an anonymous reviewer for con-
structive suggestions that have improved the manu-
script of this paper.
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