This paper not to be cjted without the aythors permjssjon
C.M.1990/F:22
Mariculture Commitee
Intemational Council for the
Exploration of the Sea
RESPIRATION'AND BEHAVIOR OF Macrobrachium rosenbergii AT
DIFFERENT LEVELS OF OXYGEN SATURATION
•
by
R.C. Reyes
Biologische Anstalt Helgoland
Notkestraße 31
2000 Hamburg 52
Federal Republic 01 Germany
U. Waller and H. Rosenthai
Institut lür Meereskunde
DOstembrookerweg 20
2300 Kiel 1
Federal Republic 01 Germany
ABSTRACT
Respiration 0110 rnonth-old sibling M. rosenbergii adults belonging to 3 size classes (3. 13 and 44 g)
was measured at various levels 01 oxygen saturation (90, 85, 75, 65, 55. 45, 35 and 25%). Behavior
01 the prawns in the respirometer at various saturation levels was sirnultaneously lilmed using a video
camera. Oxygen consumption of M. rosenbergii was alrnosl on a conslant level lrom 20 to 90%
oxygen saturation. Respiration rales were 0.25 • 0.3 mglglh in animals weighing 44 g. The s naller
individuals (3 and 13 g) had a higher oxygen demand around 0.40 - 0.55 mglg:h. The largest prawns
(44 g) showed the least Irequency 01 aggressive behavior and were comparably quiescenl Ihan the
smaller ones (3 and 13 g).
INTROOUCTION
giant freshwater prawn Macrobrachium rosenbergii (Decapoda, Palaemonidae)
.e of the intensively cultured freshwater crustceans in the world. Since the
i
pi leering work of UNG (1961, 1969) thirty years ago, many aspects of its biology
relevant to aquaculture has been studied.
I
Metabolie aetivity in aquatic animals depends on the dissolved oxygen (00)
avaiJable ;n the environment. Oxygen is therefore one of the most important Iimiting
factor for survival and growth. Only few studies on the optimum threshold level were
done in M. rosenbergii and they could not agree. SCOTT (1988) detected no visible
signs of stress in M. rosenbergii until 00 dropped below 1 mgll. Investigating growth
rates, optimum threshold levels between 1.5 to 5.0 mg/l 00 were suggested
(LLOBRERA and NEILL 1989. ROUSE 1981, SM/TH. SANO/FER and TRIMBLE
1976).
.
For aquaculture the knowledge of optimum 00 levels required by a eultured spadas
is aprerequisite. The main purpose of this study is to acquire first insight into the
resistanee of M. rosenbergii to low 00 levels, respiration of different size c1asses and
their behaviour at various saturation levels.
Page 1
MATERIALS AND METHODS
M:
Ten month old
rosenbergii adults (Irom time of hatching) raised from 01. single brood
up to maturity were arbltranly c1assified for experimental purposes into three size categories. The first group (Grp. S) consisted of 8 small male prawns and weighed 2.9 ±
0.3 9 each (total weight == 23.2 g). The second group (Grp. M) consisted 01 2 males
and 2 females and hOld an average weight of 13.3 ±1.1 9 (total weight = 53.7 g). In the
third group (Grp. B) were 2 blue-clawed males weighing 44.2 ± 11.1 9 each (total
weight == 88.5 g). Only prawns in the intermolt stage were used for experimetal
purposes. Twenty hours prior to measurement, the prawns were fed with artificial dry
feed (Goldflsh Tetra Feed) at 01. ration of 1l!'o body wet weight.
Peristaltic pump
Fie~:Qjr
I---"';@
Fi.~~r
I-----l@
•
Peristaltic pump
Pump
Waste water ........._--.
Animal chamber
D
Oxygen
probe
Computer
Figura 1: Rcspllomctcr uscd
In
the oxygen uplake expemllcnls
•
Figure 1 shows the computer-controlled, c10sed respirometer system used for measuring oxygen consumption. During the experiments temperature of the measuring system were maintained at 28iO.2°C. The saturation level of water flowing through the
system was continously read using an oxygen meter. During the test, the system was
closed and consequently the oxygen saturation of water dropped due to animals
respiration: When the saturation level dropped below the chosen threshold, a
peristaltic pump maintained oxygen saturation at the desired level (± 0.1 %) by
delivering approximately 100% oxygen saturated water from the reservior
(28.21.O.2'C) into the respirometer. Respiration rates were simultaneously computed
Pagc2
from the amount of oxygen delivered with the water, the space of time and the wet
weight of the animals.
During the experiments the saturation level could be changed by hand or automatically by the computer program. Running the automatie mode, the time of each saturation level, the highest and lowest saturation and the value for the stepwise inerease or
decrease of oxygen saturation could be defined. If the new saturation level was lower,
a seeond pump fed water aerated with N2 gas into the measuring system until the new
level was reaehed. If the new level was high er, oxygen saturated water was delivered
into the system. In doing so new saturation level was quickly reached. In the experiments with M. rosenbergii respiration rates were measured at saturation levels of 85,
75, 65, 55, 45, 35, 25, and 90% (recovery), wherein each level was maintained for one
hour.
•
0,80 . , . . . . . - - - - - - - - - - - - - - ,
:E
.Ql
SCl
0,70
0,60
+
0,50
§
0,40
.~
0,30
.~ 0,20
a::
0,10
o,00
-f--,~~~.._r~.......,~~_r_"~....-
o
3
........-1
6
9
6
9
Time (h)
100
.......
•!'
~
80
c:
e
.'"0<1l
:;
iii
<J)
c:
Ql
Cl
S<O
60
40
20
0
0
3
Time (h)
Figure 2: A rcprescnlalive cxample 01 a conlmuous SCIICS 01 oxygcn consumptlon dcfcrmlnatlOns 111 M
Tosenbergii (Mean wcl weighl per pravKI=13.0 g. n=4).
The behaviour of the prawns was video recorded at the different saturation levels
(excluding 90%). Video reeording automatically started at the midphase of each hour
Paqc 3
01 measurements. The time 01 each sequence was set to 10 min, which were interpreted at the end 01 the experiments.
Prior to the start 01 each experiment, oxygen concentration 01 water in the reservior
(7.97 ± 0.21 mg/I, appr. 102'!·o saturation) was determined by Winkle(s method and
the wet weight 01 the test animals was measured. Belore the start 01 the measurement,
the prawns were acclimatized to respirometer conditions for 3 hours at a constant flow
of 100% saturated water maintained at 28.2±0.2'C. After that, the experiments were
started.
RESULTS
A representative example of a continuous series of oxygen consumption determina·
tions is shown in Figure 2. In this particular case, respiration of four M. rosenbergii
adults, each weighing 13.0 9 on the average, was monitored. During the first hour of
the experiment oxygen consumption was 0.28-0.48 mg/g/h at 85% saturation (Figura
2). In the next 4.2 h oxygen saturation was lowered from 75 to 45 % in steps of 10'!W
every hour. At this range, no distinct changes in oxygen consumption were detectable. 5.3 h after the start of the experiment, 35% oxygen saturation was reached. At
this level, oxygen consumption of the test animals increased to values between 0.48 •
0.72 mg/g/h. At the lowest saturation tested (25%, 6.5·7.5 h), respiration rates again
decreased to a comparatively lower level (0.38 ·0.64 mg/g/h). After 8.7 h, oxygen sa·
turation was brought up to 90%. At this level, respiration rates were between 0.42 and
0.50 mglg/h.
0,80
~
-r-------------T
0,70
~ 0,60
.s.
Cl
0,50
§
0,40
.~
0,30
CI:
0,20
0,10
0,00
'n
~
-j--o-.-......-.-..--..-.......--.-....--,r-"""-.-,.-....--.-.-+
o
20
40
60
80
Oxygen saturation (%)
100
•
FIgure 3: Oxygen consu01pllon 01 M losenbelgii (Mean wel weighl per prawn=130 g. n=4) at different
levels 01 oxygen saturation.
In Figure 3, the data obtained in the above described experiment were rearranged.
The oxygen consumption of M. rosenbergii was plotted against the levels of oxygen
saturation. The relationship between respiration and oxygen saturation was alm ost
linear in the range from 85 • 45% saturation. At 35 and 25% saturation, mean
respiration rates increased by 38 and 16%, respeetively. Nevertheless, respiration
rates at all tested saturation levels did not differ significantly from each other at the 5%
level (AN OVA). During recovery (90% saturation) oxygen consumption seemed to be
slightly higher compared to that measured at 85% at the begin of the experiment. This
dlfference can not be interpreted satislactory. because there were only three readings
at 90% saturation (Figure 2).
Pagc4
Weight specific oxygen consumption of the three different size c1asses of M. rosen·
bergii adults in relation to Oxygen saturation level is shown in Figure 4. The relation·
ship between respiration and oxygen consumption in all test groups was almost linear. Over the range of test saturations, respiration rates for every two replicates
amounted to 0.44-0.54 mg/g/h in group S, 0.38-0.58 in group M, and 0.28-0.31 in
group B. These results showed the general trend that respiration rates are usually
declining with increasing body size. Results in Figure 4 indicate that prawns of all the
size c1asses could tolerate low oxygen levels.
0,70
0,60
•
~
.Ql
GroupS (3 gl
0,50
Group M (13g)
Cl
.s
0,40
.~
0,30
§
0
0
~
0
GIOUp B (44g) ~
6.
6-.1.
6.
~
fl
'n.
0,20
Q)
a:'"
0,10
0,00
0
60
40
20
Oxygen saturation
80
100
(~'o)
Flgme 4: Re5f~ration 01 small (Grp. SI, mediurn-sizoo (Grp. Ml and largp. (Grp, B) M./osenbetgH Mulis at d,«".",,1
levels 01 0XY!J'ln saluration. Unes reples"nl lhe average respiratioo rale 01 e",!IY !wo I<"phcates.
60
•
.~" , .c-
50
~o
40
<'il '"
30
~ E
~::
o~
t
..a
E
::J
Z
c
:;j
8c:
Ql
20
10
•
0
25
35
45
-
55
o
o
•
65
75
GrpS(Jg)
Grp 1.11(13 g)
GrpB(44g)
85
Oxygen saturation (%)
Figlle 5: Observed numbers 01 aggressive encounlers al d'«"".'111 levels 01 oxYIJ"n salurabon ,n srnall (Orp. SI.
medlum·sizoo (Grp. MI and large (Grp. Bl M.rosenbefgii adults. Each bar r~l~esenls lhe avera!)'J 01 Iwo ICI~K:alt'S.
Page 5
Figure 5 shows the number of aggressive encounters at different oxygen saturation
levels .The lowest number of encounters (between 1 and 4 for every 10 min
observation) were obtained in the largest prawns (Group B) which were relatively
inactive in all saturation levels tested. Prawns of groups Sand B were relatively more
aggressive. The numbers 01 aggressive encounters fluctuated between 20 and 60 for
every 10 minute observation in groups S, whereas in group M 30 • 40 encounters
were observed.
OISCUSSION
Previous studies on the respiration of M. rosenbergii larvae (STEPHENSON and
KNIGHT, 1980a ), post larvae (STEPHENSON and KNIGHT, 1980b) and juveniles
(NELSON, LI and KNIGHT, 1977) showed that oxygen consumption is a logarithmic
funetion of the body weight. With reference, however, to varying oxygen saturation
levels, MALECHA (1983) reported that below 55 mmHg (6.45 ppm or 75% saturation
at 22"C) respiration is a linear function of oxygen concentration. Results of the present
study indicate such a linear relationship between 90 ·25% saturation at 28°C in all
the three size c1asses of adult prawns tested.
The results give evidence that respiration of M.rosenbergii is independent of the
oxygen saturation levels tested. These findings emphasize a high tolerance of this
species against an oxygen d epleted environment. A similar relationship has been
reported for a deep water Mysid, Gnatophausia ingens, which is a typical inhabitant of
the oxygen minimum layers of the ocean (CHILORESS 1971). In contrast , Euphausia
superba is extremely sensItive to oxygen depletion (KILS 1979). In this species, which
lives in the weil oxygenated surface layers of the ocean, respiration rates increased
up to 85% saturation and then decreased parallel to declining oxygen saturations.
Lower limit for E. superba is around 85% if exposed to it within 12h. The high
tolerance of M. rosenbergii to oxygen depletion is an adaptation to its natural habitat·
lakes, rice paddies, and mangrove swamps with daily fluetuations of oxygen levels.
For animals living in unstable environments (e.g. G. ingens, M. rosenbergit) , the
development of a high tolerance is prerequisite for survival, whereas under constantly
favourable environmental conditions (E. superba) , such adaptations may not be
necessary.
Wlthin the range of saturation levels considered in this study, no visible signs of stress
were ever observed even at 25l!'o saturation. All prawns maintained their upright
position for an hour at this lowest level. According to MALECHA (1983), the lowest
oxygen level wherein prawns can be maintained wlthout stress is between 25 • 30%
saturation (2.25 -2.75 ppm) at temperatures between 25 - 30°C. LOSOROO (1980)
and MALECHA (1983) suggested maintaining 00 ccncentrations above 3 ppm (appr.
40% saturation) for prawn farming. Visible signs of stress may not be immediately
observed in short term exposures to low oxygen levels, but long term exposures may
indicate otherwise. CLARK (1986) studied the effeets of a 17 day-hypoxia (2 ppm) on
molting and survival of adult female Penaeus semisulcatus (53.8±10.4 g) and
observed that the prawns did not molt at all and mortality was high. After oxygen level
was subsequently increased to 5 ppm, mortality ceased and many moltings were
observed. A 40 day experiment in M. rosenbergii juveniles on the effeets of various
oxygen saturation levels (2.5, 3.5, 5.0 and 7.7 ppm 00) indicated that at 2.5 ppm there
was a significant decrease in growth rate, a lower (not statistically significant)
decrease in survival and a significant decrease in food intake. However, no significant
effect of 00 levels on molting and food absorption was detected (LLOBRERA and
NEILL, 1989).
Pa<jC 6
•
•
Contrary to the general description that blue - clawed males are dominant and small
males nondominant (RA'ANAN and COHEN, 1985), the largest prawns (44 g) were
observed to be the least aetive (hardly changing their positions) and aggressive,
whereas the smallest ones (3 g) had the highest frequency of aggressive encounters.
The chance of encountering other prawns increases with the number of individuals
present, but the (arge prawns used in the experiments were observed frequently at
body contaet or at one's reach with the periopods and yet the two usually did not fight
at such proximity. Fluctuations in aggressive encounters within smaJl (group S) and
medium sized prawns (group M) from one saturation level to the next do not, at this
point in time indicate definite trends. Analysis of intensity of encounters, as weil as
respiration and behavior at a particular saturation level within a 24 h period, will be
presented in a future paper•
•
•
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Page 7