JOURNAL OF

EXPERIMENTAL
MARINE BIOLOQY
Journal of Experimental Marine Biology and Ecology AND ECOLOGY
ELSEVIER 183 (1994) 179-192

Biochemical composition of Aurelia aurita in relation to age

and sexual maturity

C.H. Lucas
Department of Oceanography, The University, Highfield, Southampton, SO9 5NH, UK

Received 17 February 1994; revision received 17 May 1994; accepted 27 May 1994

Abstract

Dry weight (DW), ash free dry weight (AFDW), and proximate biochemical composition were
examined for a full range of sizes of whole medusae of the scyphomedusa Aurelia aurita. Dry
weight ranged from 3.07 to 3.91 y0 of wet weight (WW), and was fairly uniform throughout the
size groups. Ash free dry weight ranged from 25.6 to 46.0% of DW, with values significantly
greater in ephyrae and medusae < 20 mm. The sum of carbohydrate, protein and lipid fractions,
termed “total organics”, is often taken to be synonymous with AFDW, but values were consis-
tently lower than AFDW, ranging from 5.56 to 22.76% of DW. Even after AFDW had also been
calculated assuming water of hydration to be 11.7%) there were still differences between the two
values. Whole Aureliu aurita medusae displayed the typical characteristics of gelatinous preda-
tors, with low carbohydrate (0.10-l. 12% of DW), intermediate lipid (3.50-l 1.50% of DW), and
high protein (2.07-28.56x of DW) content. The biochemical composition of isolated umbrellar
and gonadal tissues of medusae > 50 mm were examined, and in every size group, “total
organics” were greater in gonadal than umbrellar tissue. The greater organic content of gonadal
tissue was the result of an increase in all three biochemical fractions, with carbohydrates and
proteins displaying the greatest differences. Mean values ranged from 0.33 to 0.85% carbohy-
drate, 2.34-8.31% protein, and 0.87-2.91x lipid in umbrellar tissue, compared with 1.08-
2.10%,4.38-22.98x, and 2.59-6.08%, respectively, in gonadal tissue. The biochemical content
of ripe oral arms, i.e. those containing dividing eggs and planula larvae, was similar to that of
the maturing gonads. Protein content was greatest in gonads of medusae 60-110 mm, before
decreasing significantly with increasing bell diameter. Rapid growth and maturation in the 60-
110 mm size group was coincident with the spring peak of mesozooplankton abundance in
Southampton Water, while the decline in protein content may be explained by loss in condition
of “spent” medusae.

Keywords: Aurelia aurita; Biochemistry; Sexual maturity; water of hydration

0022-0981/94/$7.00 0 1994 Elsevier Science B.V. All rights reserved

SSDI 0022-098 1(94)00089-V
 180 C.H. Lwos ‘J. t.;‘\-p.Mar. Bid. Ecd. 1X3 (lYY4, 17%IY?

1. itttroductioa

Knowledge of the biochemical composition of organisms is a major element to the

understanding of the transfer of energy and matter within an ecological community
(Percy & Fife, 1981). Until very recently, the elemental and biochemical composition
of gelatinous zooplankton had not been examined, but increasing awareness of the
importance of these predators within the pelagic community has attracted attention to
this area (Clarke et al., 1992).
Of the comparatively few gelatinous biochemical studies, most have been carried out
on ctenophores (Hoeger, 1983; Morris et al., 1983; Clarke et al., 1987, 1992; Schneider.
1989). By contrast, little is known about the biochemical composition of medusae, with
studies limited to those of Percy & Fife (1981). Schneider (1988a), Arai et al. (1989)
and Clarke et al (1992) two of which are from polar regions. Gelatinous zooplankton
are characterised by high water and mineral ash contents (Ikeda, 1972; Percy & Fife,
1981; Larson, 1986b), and in relative terms, a high protein, intermediate lipid, and very
low carbohydrate content is typical (Mayzaud & Martin, 1975; Hoeger, 1983; Schneider.
1988a). Data in the literature for ctenophores give values, expressed as a percentage
of dry weight (DW), ranging from 3.06 to 23.8”,, protein, 1.7?, to 14.49, lipid, and
0.5”” to 1.1s; carbohydrate (Percy & Fife, 1981; Hoeger, 1983; Schneider, 1989) with
similar values for medusae (Percy & Fife, 1981).
According to Giese (1967), in order to study the relation of the biochemistry of an
animal to its nutritional or reproductive state, analysis of isolated body components are
more informative than analysis of the whole body. The gonads of medusae. for example,
often represent a substantial fraction of the total organic content, and are directly af-
fected by these short-term changes in food availability (Larson, 1986b). The vast ma-
jority of research into the biochemical content of gelatinous zooplankton has been
carried out on whole animals, and this includes studies of changes in biochemical
composition in response to food supply (Reeve et al., 1970; Hoeger, 1983; Morris et al.,
1983; Arai et al., 1989). The elemental, i.e. carbon and nitrogen, content of isolated
body components of medusae have been examined by Schneider & Weisse (1985),
Shenker (1985) and Larson (1986a), but the distribution of organic substances within
a gelatinous predator has only previously been described for a relatively small sample
of the scyphomedusan Aurelia aurita from the Baltic Sea (Schneider, 1988a).
On a seasonal basis, Aurelia aurita is a significant member of the gelatmous preda-
tor community of Southampton Water, forming up to 96”: of the total annual carbon
biomass (Lucas & Williams, 1994). As part of a wider programme to examine the
bioenergetic flux within the zooplankton community of Southampton Water (Lucas,
1993) the temporal relationship between size and biochemical composition of whole
and isolated tissues of A. aurita ephyrae and medusae will be described.

2. Methods

2. I Collection

Aurelia aurita medusae were collected from March through to the end of June as part
of a twice-monthly zooplankton sampling programme in Southampton Water, either by
 C.H. Lucas /J. Exp. Mar. Biol. Ecol. 183 (1994) 179-192 181

towing 212 pm and 340 ,um cod-end plankton nets, or by dip-netting them from the
surface. The medusae were maintained in seawater and transferred to the laboratory
within 3 h of capture. Care was taken not to damage the animals, particularly the
tentacle areas. During capture and maintenance it was noted that there was no mucous
production, an indication that the medusae were not “stressed”. Bell diameter was
measured, and excess salt rinsed off using distilled water.

2.2. Water and organic content

Wet weight (WW) was obtained after carefully “blotting” the medusae to remove
superficial water, but without causing damage. After wet-weighing, each individual was
placed in weighed pre-ashed crucibles and dried at 70” for 24 h, or until a constant
dry weight (DW) was obtained. The crucibles were cooled in a desiccator prior to
weighing. The dried samples were ashed in a muffle furnace set at 550” for 24 h to burn
off organic material. The resulting ash weight (AW) was measured after cooling in a
desiccator, and ash free dry weight (AFDW) calculated as the difference between DW
and AW.

2.3. Biochemical composition

Proximate biochemical analysis was carried out on x 100 whole individuals, but in
order to determine the biochemical composition of reproductive organs, up to 50 larger
(> 50 mm) Aureliu medusae were dissected carefully to remove the four horseshoe-
shaped gonads from the gastric cavity in the umbrellar tissue. Where planula larvae
were present in the brood pouches on the oral arms of mature adults, usually > 90 mm
bell diameter, the oral arms were also separated from the umbrellar tissue. Specimens
were either freeze-dried immediately or frozen at -20 o in individual sealed bags for up
to 9 months prior to freeze-drying. Apart from ephyrae < 10 mm, which were pooled
together in groups of 30, biochemical assays were made on individuals.
The resulting freeze-dried material was pulverised into a powder using a glass-rod
homogeniser. The ground, dried material was weighed into three approximately equal
aliquots of 5-50 mg each, one for each biochemical assay, although l-5 mg samples
were used where only small quantities of tissue were available. Each aliquot was pre-
pared for the appropriate assay by adding either 4ml O.lN NaOH for total protein
analysis, 4ml distilled water for total carbohydrate analysis, and 4ml 2: 1 chloroform/
methanol for total lipid analysis. The samples were thoroughly mixed using a Janke &
Kunkel IKA T25 homogeniser for ~2 min at 10000 rpm.
Protein was determined calorimetrically according to a modification of the Lowry
et al. (1951) Folin Ciocalteau method, with BSA (V), diluted using 0.9% w/v KC1 as
a standard. Carbohydrate was determined calorimetrically directly from freeze-dried
material using the phenol-sulphuric acid method of Dubois et al. (1956), with D-Glucose
as a standard. This method is particularly suitable for gelatinous material because
of its sensitivity to micro-quantities of sugar. Total lipid was first extracted using
2: 1 chloroform/methanol (Bligh & Dyer, 1959) and determined gravimetrically accord-
ing to Folch et al. (1957), following washing of the crude extract with 0.05N KCl.
In all cases, each sample was assayed in triplicate, and the results expressed as
mg g. DW-‘.

3. Results

Apart from Fig. 2 and Table 2 where data are presented for individuals, the medu-
sae have been sorted into size groups, and the results are presented as the mean of
individual analyses within the size group.
Mean dry mass for whole medusae ranged from 3.07 to 3.91’~ of wet weight (WW)
(Table l), and apart from slight increases in the smallest and largest medusae, was fairly
uniform throughout the size ranges. Ash free dry weight as a percentage of DW was
more variable, with values ranging from 25.6 to 46.0%. AFDW of ephyrae and 1 l-20
mm medusae was signifkantly greater than in all but 91-100 mm medusae (p < 0.05).
AFDW also increased significantly to > 30 0/i in medusae larger than 80 mm, probably
reflecting gonadal ripening which occurred at these sizes (Lucas, 1993).
The sum of carbohydrate, protein, and lipid fractions, termed “total organics” was
calculated where each substrate could be determined from the same individual. In whole
medusae, values generally decreased with increasing size, and ranged from 5.56 to
22.76’?, of DW (Table 1). Total organic content was consistently lower than AFDW
in A. aurita medusae, comprising 22 to 5 1 g,; of AFDW. This discrepancy is usually
assumed to be the result of “water of hydration” that is still retained in tissues after
drying at 60”, but lost during ignition at 550” (Madin et al., 1981; Larson, 1986b).
AFDW and total organics were calculated assuming water of hydration to be 11.7?,

Table I
Mean ( + SD) percentage water and organice content of whole A. uurira size groups

Size DW (‘,“WW) AFDW (“,,DW) Total organics (“,DW)

(mm)
I, measured II measured revised II measured revised

-c l(l 22 3.91 (1.83) 14 45.96 (16.71) 35.20 _

I I-20 28 3.44 (0.65) ‘5 44.21 (17.30) 34.25 9 12.76 (7.72) 25.57
71-30 7 3.31 (0.27) 6 29.00 (16.71) 21.68 7 ) 2.29 (4.47) 13.92
31-30 4 3.12 (0.15) 4 28.42 (4.04) 17.70 6 9.67 (4.32) 10.95
‘tl-SO 5 3.32 (0.37) 5 25.57 (1.76) 14.71 5 6.44 (2.29) 7.29
5 l-60 7 3.10 (0.39) 7 26.04 (2.89) 22.00 10 5.79 (2.00) 6.56
6 I-70 11 3.07 (0.65) 11 26.43 (2.96) 22.78 6 10.64 (3.88) 12.05
71-80 15 3.32 (0.67) 16 27.23 (3.81) 16.11 1 5.56 (0) 6.30
8 l-90 II 3.69 (0.49) II 31.59 (6.91) 22.25 I x.23 (0) 9.32
91-100 9 3.43 (0.77) 9 36.04 (8.40) 22.58
> 100 IO 3.79 (0.84) 10 32.69 (9.17) 22.79 11 (I.77 (1.93) 7.66

Mean 128 3.42 (0.76) 120 32.46 (14.321 23.7u 56 9.79 (7.11) 11.07
--_--
Ash free dry weight (AFDW) and total organic (sum of protein, lipid and carbohydrates) content, are given
as the actual “measured” values and “revised” values, assuming water of hydration is 11.7”, of DW (Lar-
son. 1986b).
 C.H. Lucas/J. Exp. Mar. Biol. Ecol. 183 (1994) 179-192 183

of DW (Larson, 1986b), and although the discrepancy was reduced, total organics were
still only 30-75% of AFDW (Table 1).
The typical gelatinous zooplankton pattern of very low carbohydrate, intermediate
lipid, and high protein content of different size groups of whole medusae is illustrated
in Fig. 1. Mean carbohydrate, typically 0.35 to 1.12% of DW, showed little significant
variation with size at the 95% confidence level, although the value of 0.096% for the
91-100 mm size group is extremely low. Mean protein content ranged from 2.07 to
28.56% of DW, with the greatest values of > 9% of DW in ephyrae, small medusae
< 20 mm, and medusae 6 l-80 mm in size. Mean lipid content showed a similar trend
to that of proteins, with greatest values of between 3.50% and 11.50% of DW in small
medusae < 40 mm, before declining rapidly to 1.17% to 3.39% of DW in larger me-
dusae.
The relationships between the weights of carbohydrate, protein, and lipid as a func-
tion of total DW of individual medusae were determined on log-transformed data using
least squares regressions (Fig. 2). Carbohydrate (C), protein (P) and lipid (L) content
increased with increasing DW according to the equations, log C (mg) = 0.9020 +
1.3042 log DW (g), I = 0.9392, log P (mg) = 1.7483 + 1.1253 log DW (g), I = 0.9501,
log L (mg) = 1.1855 + 0.7712 log DW (g), Y = 0.8303.
The biochemical compositions of isolated umbrellar and gonadal tissues from me-
dusae > 50 mm bell diameter were examined, and in every size group total organics
was greater in gonadal tissue than in umbrellar tissue. Total organic content (as a
percentage of DW) of umbrellar tissue showed little variation with size, ranging from
3.68 to 11.16%, whereas in contrast, total organics of gonadal tissue decreased sig-
nificantly with increasing size from 36.13 y0 in the 5 l-60 mm size group to 7.14% in
the 141-150 mm size group (Fig. 3). The resulting U:G ratio of total organics increased
with increasing size from 0.17 to 0.56 (Fig. 3).
The greater organic content of gonadal tissue was the result of an increase in all three
major biochemical fractions (Fig. 4). Carbohydrates displayed the greatest relative
difference, with mean values of 0.33% to 0.85% in umbrellar tissue significantly (p <
0.01) less than the 1.08% to 2.10% in gonads. Similarly, proteins formed 2.34% to
8.31% of umbrellar DW compared with 4.38% to 22.98% of gonadal DW, with the
most significant differences in the 61-100 mm and 11 l-120 mm size groups (p < 0.01).
The lipid contents of umbrellar and gonadal tissue were more comparable, with val-
ues ranging from 0.87 to 2.91% of DW in umbrellar tissue compared with 2.59% to
6.08% of DW in gonadal tissue. Statistically significant differences (p < 0.05) in lipid
content of umbrellar and gonadal tissue were only apparent in the 91- 100 mm, 11 l-
130 mm, and 141-150 mm size groups. Overall, there were very few significant size-
related differences in biochemical composition of umbrellar tissue, and similarly, there
was little variation in carbohydrate and lipid content of gonadal tissue. By contrast,
protein content of gonadal tissue decreased significantly with increasing bell diameter
(r = 0.885, F = 36.23, sig = O.OOOl), which accounts for the total organics pattern
of Fig. 3.
The carbohydrate, protein, and lipid content of isolated ripe oral arms, i.e. those
containing dividing eggs and planula larvae, were determined from seven medusae
between 71 mm and 160 mm in size (Table 2). The results showed no apparent size-
1x1 C.H. h~?rs .f. t.yp bdar. &ot. htt. t8.i !iYY4/ t 7Y-iv_’

related trends, but in general, the biochemical composition of planula larvae were
similar to that of the gonads. with mean values of 1.08”;, carbohydrate, 11.38”, pro-
tein, and 3.37:$ lipid.

0
50

40

30

20

10

0
25 _______.___
_...._
‘1 .--- I.,1I-“‘!S ’

20 -1 i,
15

70

Fig. 1, Carbohydrates, proteins, and lipids content of different six groups of whole ,4 cauitamcdusac. Figure
shows mean, standard deviation. and sample number.
 C.H. Lucas /J. Exp. Mar. Bid. Ecol. 183 (1994) 179-l 92
185

103

lo2

10'

loo

10-l

1o-2

1O-3

1o-4

1o-5

lO+j I I I I I I I

lo-5 1o-4 1o-3 10-z 10-l 100 10' 102

lo4
log P (mg) = 1.7483 + 1.1253 log DW (g)
lo3 r = 0.950. n = 103

lo2

10'

loo

10-l

1o-z

1o-3

1o-4

1o-s I I I I I I I

1 o-5 1o-4 1o-3 1o-2 10-l

104
G L (mg) = 1.1855 + 0.7712 IagDW (g)
P = 0.830, n = 57
lo3

IO2

10'

loo

10-l

1o-2
I I I I

1o-3 10-z 10-l loo 10' lo2

log,, g Dry Weight

Fig. 2. Relationship between the weight of carbohydrates, proteins and lipids as a function of dry weight of
whole A. au&a medusae, with the resulting equations solved by leastsquares regressions.
1% C.H. Lucus J. E.xp. Mar. Bid. lhd. 183 1lYY41 17%IY.’

Fig. 3. Total organics content of whole, gonadal and umbrellar tissue from different sire groups of A. uuriru
medusae. Figure shows mean and standard deviation.

4. Discussion

Almost without exception, published data on the water and organic contents of
medusae have been presented as single values from whole animals of an unspecified
size. Shenker (1985) and Matsakis & Nival (1989) are alone in presenting DW and
carbon contents for both mature and immature Chrysaom and Phi~lidium, respectively,
with the mature medusae typically having the greater DW (as”., of WW). Clarke et al.
(1992) examined the DW, AW, and elemental composition of a range of sizes of

Table 2
Biochemical composition of isolated oral arm tissue

Size “,, of DW
--_l------.-
(mm)
Carbohydrates Proteins Lipids

71 (1.88 IS.27 I.56

9x 1.25 IO.61 1.70
100 0.9’ 15.21 3.96
11x 1.17 13.08 3.62
I’X 1.12 IO.71 2.07
13X 0.63 4.07 1.33
160 1.21 IO.70 7.5X
 C.H. Lucas / .i. Exp. Mar. Biol. Ecal. 183 /I9941 179-192 187

Southern Ocean gelatinous organisms, but in contrast to the present study where
AFDW was greatest in ephyrae and small medusae, no significant variation with size

5 t
Carbohydrates

4-

20
Lipids

IO -

5-

Size group (mm)

Fig. 4. Carbohydrates, proteins and lipids content of isolated gonadal and umbrellar tissue from different
size graups of A. auritu medusae. Figure shows mean and standard deviatian.
 was reported. Dry weight and AFDW values for a range of sizes of‘4. auritu medu-
sae in the present study, are in broad agreement with those found in the literature
obtained from A. auriru of unspecified sizes (Lowndes, 1942; Larson, 1986b).
Ash free dry weight and organic weight have often been taken to be synonymous,
but a discrepancy between the sum of carbohydrate, protein, and lipid content, termed
“total organics” (Madin et al., 1981) and AFDW has been highlighted. In the present
study. the total organic content of A. mrita comprised 22-5 1Ouof AFDW, with similar
observations made for salps where total organics were 19-51.8”, of AFDW, and the
sum of carbon, nitrogen, and hydrogen was 21-67”,, of AFDW (,Madin et al., 1981).
This discrepancy between total organics and AFDW is usually assumed to be the re-
sult of “water of hydration” that is lost during ignition at 550 (Madin et al.. 1981:
Larson, 1986b). However, even when the effect of 11.7”,, water of hydration (Larson.
1986b) was taken into consideration for A. m&o, total organics were still 25-70”; less
than AFDW. Clarke et al. (1992) have suggested that gelatinous zooplankton contain
5- IO”, (of DW) amino-carbohydrates and/or glycoproteins that are missed or under-
estimated by conventional assay techniques. However, because this still would not
account for all of the discrepancy between total organics and .4FDW in the present
study, it remains unclear what the exact reasons for the differences are. As a result,
Hoeger (1983) has suggested that AFDW used as a basis for organic matter will
therefore lead to great errors. with carbon and nitrogen content providing a more
meaningful value.
0ne obstacle encountered in the present study was the very small quantities of ge-
latinous tissue available after drying, often S-10 mg, representing 3-40,, of the origi-
nal WW. Many authors have overcome this by pooling up to 50 individuals for analysis
together as one “batch” (Percy & Fife, 198 1: Hoeger. 1983; Morris et al., 1983; Arai
ct al.. 1989). In the present study this common practice was avoided where possible,
so that individual and seasonal variation in biochemical composition could be deter-
mined. The carbohydrate, protein, and lipid contents of whole A. mm7u medusae ob-
taincd in the present study fall within the relatively wide range of literature values
obtained primarily for polar medusae (Table 3), and are also comparable with the oni)
other data published for .4. auritL/ from the Baltic Sea (Schneider. 1988a). In terms of
size-related trends, protein and lipid showed the greatest range, with higher values
typically found in ephyrae and small medusac.
.4lmost without exception, lipids have been found to be an intermediate biochemi-
cal component of gelatinous predators. comprising 1.9-22.1°0 of DW of medusae
(Table 3). Because of the absence of typical storage lipids such as wax esters (Lee, 1974;
Clarke, 1984), proteins rather than lipids are thought to be the main storage product
in gelatinous zooplankton (Morris et al., 1983; Clarke, 1984). Indeed. lipids in gelati-
nous predators are comprised chiefly of phospholipids (67”“) which are viewed as
having a structural role, and this would explain the greater proportion of lipid, relative
to proteins, found in umbrellar tissue when compared with gonadal tissue of A. auritL~
in the present study.
In order to relate the biochemistry of an animal to its nutritional or reproductive state,
analyses of isolated body components are more informative than the whole body (Giese,
1967). The gonads of medusae, for example, often represent a substantial fraction of
 C.H. Lucas / J. Exp. Mar. Biol. Ecol. 183 (1994) 179-192 189

Table 3
Review of the carbohydrate, protein, and lipid composition of medusae, as a percentage of dry body weight

Species Area Carbohydrate Protein Lipid Ref.

Anthomedusae
Bougainviliia superciliaris Arctic 0.7-1.0 7.7-14.9 6.8-10.0 1
Sarsia princeps Arctic 0.4-0.8 14.5-14.7 7.8-9.1 1
Hybodocon prolger Arctic 0.8 23.0-31.0 13.1-22.1 1
Halitholus cirratus Arctic 0.7-0.8 10.4-18.2 4.6-7.6 1
Calycopsis borchgrevinki * Antarctic 1.12 11.01 2.15 2

Leptomedusae
Aequorea victoria British Columbia 0.7 6.6 1.9 3

Scyphomedusae‘
A. aurira Baltic Sea 2.9 5.9 2.17 4
A. aurita Southampton 0.10-1.12 2.07-28.56 1.17-11.50 P.S.
A. wyvelii* Antarctic 1.71 16.95 4.17 2
Rhizostoma pulmo Mediterranean - 8.6 3.5 5

Trachymedusae
Aglantha dig’rale Arctic 0.4-0.9 21.6-22.1 6.0-6.9 1
Aglantha digitale Bering Sea 0.8 56.5 3.0 6

References: 1. Percy & Fife (1981), 2. Clarke et al. (1992), * Value obtained using WW to DW conversion
from paper, 3. Arai et al. (1989), 4. Scheider (1988), 5. Ceccaldi et al. (1978), 6. Ikeda (1972), P.S. Present
study.

total carbon content, and are directly affected by short-term changes in food availability
(Larson, 1986b). AFDW has been found to comprise 29% of oral arm tissue compared
with 23% of umbrellar tissue in the scyphomedusan Cyanea capillata, and 92% of
gonadal tissue compared with 25 yO of umbrellar tissue in the hydromedusan Stomotaca
atra (Larson, 1986b). The elemental content (C,H,N) of isolated tissues of various
medusae have been examined by a number of authors, such as Schneider & Weisse
(1985), Larson (1986a; 1986b), and Schneider (1988b). Values of 4-15x C and l-4%
N are found in whole scyphomedusae (reviewed by Larson, 1986b), with the resulting
low C:N ratios of 3.4 to 4.3 typical of a high protein content (Morris et al., 1983;
Larson, 1986b). As a percentage of DW, gonads of A. aurita contain 12-17x C and
3-5x N (Schneider, 1988b), while in contrast planula larvae contain 39% C and 9%
N (Schneider & Weisse, 1985). Schneider (1988b) also observed that in years of low
planula larvae production by A. au&a, the C:N ratio of the larvae was 6.2, indicative
of a lipid-dominated composition, whereas when larvae prqduction was high, the C:N
ratios fell to 4.5-4.7. The protein, lipid, and carbohydrate content of the reproductive
organs of gelatinous predators, however, is poorly known, although the only available
published data are for A. aurita(Schneider, 1988a).
In the present study, the importance of gonads in reflecting the nutritional and re-
productive condition of the whole medusa was apparent, whereby the trend of declining
total organic content of gonads with size was reflected in those of the whole animal.
Umbrellar tissue by contrast showed little or no variation in biochemical composition
 with size. The total organic content of,4. uuriru was between 2.7 and 5.8 times greater
in isolated gonads than umbrellar tissue. This difference was attributed primarily to
differences in protein content (1.9-6.7 s) and carbohydrate content (1.6-5.9 X) between
the two tissues. Data in the literature suggest that protein forms the major biochemi-
cal fraction of marine invertebrate eggs, followed by lipids and then carbohydrates.
Ecklebarger & Larson (1988) provided ultrastructural evidence that oocytes derived
yolk through both autosynthesis, and heterosynthetic incorporation of high molecule
proteins, possibly vitellogenin. In addition three types of storage product were observed
in vitellogenic oocytes, namely membrane-bound yolk bodies, glycogen, and lipid drop-
lets.
Protein content was greatest in gonads from 60 to 110 mm medusae, before declining
in larger animals. Growth and maturation of medusae is most rapid in this size group,
and occurred during May and early June, coinciding with the spring peak of mesozo-
oplankton abundance in Southampton Water (Lucas, 1993). The abundant food supply
is reflected in the biochemical composition of the gonads, resulting in mobilisation of
storage compounds and gamete production. It is unclear at what size sexual maturity
occurs in A. aurzla, that is when medusae contain eggs and planula larvae in brood sacs
on the oral arms. The minimum size for laboratory-maintained individuals was 55 mm
(Spangenberg, 1965), and although medusae containing planula larvae in the brood sacs
were generally >65 mm in Southampton Water, it is possible that smaller medusae
were sexually mature, as eggs were clearly visible in the gonads of individuals 45 mm
(Lucas, 1993). In larger (> 120 mm) medusae, the decline in biochemical content, in
particular protein content, may be explained by the transfer of eggs to the brood sacs
and subsequent loss in condition of “spent” medusae once the larvae have been shed
into the water column, and the reduction in zooplankton stocks and energy reserves
used for the growth and maintenance of gametes. According to Schneider (1988a), the
biochemical content of the oral arms of fertilized A. aurita medusae (i.e. those con-
taining planula larvae) was found to be similar to that of the gonads. However, the
gonads from fertilized medusae had a lower biochemical content than those from un-
fertilized medusae, because of the transfer of fertilized eggs from the gonads into the
brood pouches on the oral arms.
The temporal trends in the biochemical composition of A. aurita observed in the
present study were highlighted through the analysis of isolated gonadal tissue, with
protein content in particular displaying the greatest changes. Such data provide valu-
able information towards the understanding and interpretation of the reproductive
processes and energy flows of this important predator of the Southampton Water
zooplankton community.

Acknowhdgements

This research was carried out as part of a N.E.R.C. studentship, number GT4/89/
ALS/34. I would like to thank Mrs. L. Wall for her assistance in the laboratory, and
to Dr. J. Williams for reading earlier drafts of this manuscript.
 C.H. Lucas /J. Exp. Mar. Biol. Ecol. 183 (1994) 179-192 191

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