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D aan , N . an d R ingelberg, J. (1969). F u rth e r stu d ies o f th e positive a n d negative p h o to tactic
reactio n s o f D aphnia m agna S tra u s. N e th . J . Z o o l. 19: 525— 540.
D artn all. H . J. A . (1974). A ssessing th e fitness o f visual pigm ents fo r th e ir p h o tic environm ents. In:
A li, M . A . (ed.) Vision in Fish. P lenum P ress, New Y o rk , p. 543— 563.
D ouglass, J. K . (1986). T h e o ntogeny o f light a n d d a r k a d a p ta tio n in th e c o m p o u n d eyes o f grass
sh rim p , Palaem onetes pugio H o lth u is. P h .D . thesis, D u k e U niversity.
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385— 399.
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F o rw ard . R. B.. J r., C ro n in . T . W ., S te a m s, D . E. (1984). C o n tro l o f diel vertical m igration:
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to th e u n d e rsta n d in g o f d iu rn al vertical m ig ra tio n . N eth. J. Sea. R es. 2: 319— 406.
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105— 114.
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verticales jo u rn aliè res. A rch. Z o o l. exp. Gen. 64: 387— 542.
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P ro ce ed in g s o f th e 21st E M 8 S
G d a n s k . 14— 19 S ep tem b er 1986
p. 67— 7 5
1989
P L IS S N 0078-3234
P o lish A c ad em y o f Sciences — In s titu te o f O c ean o lo g y
Vlaams Instituut voor de Zee
Flanders Marine Institute
R ESPIR A TIO N D U R IN G H Y PO X IA O F T H E SH R IM PS
CRANGON CRANGON AND PALAEM O N AD SP ERSU S
FR O M B R A C K ISH W ATER
L. H a g e r m a n 1, A. S z a n i a w s k a 2
1 M arine Biological L ab o rato ry , S tra n d p ro m en ad en . DK.-3000 H elsingor, D en m ark
2 Insty tu t O ceanografii, U niw ersytet G dariski, al. Czolgistów 46, 81-378 G d y n ia, P o lan d
A B ST R A C T
The respiration o f Crangon crangon a n d Palaem on adspersus w as m easu red u n d e r n o rm o x ic
an d hypoxic conditions. All experim ents w ere ru n fo r at least 24 hours (S, 20%o; T , 20°C).
In norm oxia Crangon show s a clear diel resp irato ry p attem : highest resp iratio n d u rin g early
night an d early m orning, low est in d ay tim e. T his resp irato ry rhythm can be co rre la te d to th e diel
activity p attem . T he resp irato ry rh y th m icity d isa p p ears when oxygen tension decreases below ca
100 to rr d u e to a n increase in the average activity o f Crangon (a response to av o id the hypoxic
conditions). Crangon can m ain tain its ro u tin e resp iratio n independent o f oxygen ten sio n in the
m edium (Pw0 3) dow n to a P wO , o f 30 to rr, th e so-called Pc. T he typical diel m ean ro u tin e re sp ira ­
tory level for in term o u lt Crangon (w.w. 0.25 a n d 1.1 g) are 0.9 a n d 0.2 ml 0 2g _ ’h _1 respectively
w hen sa n d is supplied a s su b stra te a n d 1.6 a n d 0.65 m l 0 ;g _1h _1 w hen n o su b s tra te is su p p lied .
M oulting h as n o t been n o ted d u rin g th e hy p o x ia resp iratio n experim ents.
Palaemon adspersus show s no diel resp irato ry rhythm icity un d er th e given exp erim en tal co n d i­
tions, possibly a result o f a “w ro n g " su b stra te in th e experim ental cham ber. Palaem on can m a in ­
tain its respiratory independency d o w n to a PwO , o f 45 to rr. Typically Palaem on w ith a w et w eight
o f for instance 0.7; 1.0 a n d 2.1 g h as a resp iratio n o f 0.65; 0.41 an d 0.13 ml 0 , g - 1 h -1 respectively.
T he results are discussed in relatio n to the ecology o f the species a n d specially in relatio n to
hypoxic conditions increasingly o ccu rrin g in shallow eu tro p h icated w aters.
IN T R O D U C T IO N
The increasingly eutrophicated brackish w ater areas are the h a b ita t o f
a shrim p stock, which is im p o rta n t from both an ecological as well as an
economic viewpoint. In D anish w aters both Crangon crangon and Palaemon
adspersus are locally subject to an intense fishery. One consequence o f e u tro ­
phication is im paired oxygen conditions in the shallow w ater, resulting in sh o r­
ter or longer periods o f low oxygen concentrations. M ost o f these periods
occur in the sum m ertim e (M iljostyrelsen, 1984) when due to high w ater tem pe­
rature the solubility o f oxygen is already lowered and the m etabolic rate o f
69
68
m ost organism s is higher. Since th e m etab o lic ra te o f th e anim als is influenced
and because species show different degrees o f tolerance to changed oxygen
conditions, such periods o f low er oxygen co nditions m ay change the balance o f
the whole ecosystem.
O ur knowledge of the resp irato ry rate ( M 0 2) of brackish w ater anim als is
now reasonably good due to im proved techniques a n d b etter understanding o f
the physiological processes in the organism s. H ow ever, m ost results published
deal w ith sh o rt tim e experim ents, i.e. von O ertzen (1984), H agerm an an d W e­
ber (1981), van D onk an d de W ilde (1981) a n d only few e.g. B utler et al. (1978)
record the M 0 2 fo r extended periods. As the respiration is likely to vary w ith
time it is im portant (i) to record M 0 2 a fte r th e anim al has recovered from
handling and acclimated to the new enviro nm ent an d (ii) to record M O , long
enough to reveal any rhythm icity. T hese p recau tio n s m ake it necessary to use
a flow -through system for recording the M O ,. Such a system is, however,
always better th an a closed one (von O ertzen, 1984).
The purpose o f the present p a p e r is th u s to record th e M O , o f Crangon
crangon and Palaemon adspersus u n d e r n o rm oxic an d hypoxic conditions for
a t least diel periods.
R ESU LTS
R e s p i r a t i o n ( M 0 2) a n d w e t w e i g h t
T he relations between routine M 0 2 a n d wet weight for Palaemon adspersus
(n = 16) and for Crangon crangon (n = 12) during day-time are shown in Fig. 1.
2 .0 .
1.5.
1.0 .
'a
0 .5 .
E
O
•2
100
Oxygen consum ption m easurem ents (M O ,) w ere perform ed in a flow through
system. The shrim p was placed in a resp iration ch am b er (8 cm long, 2.5 cm
inner diam .) and sand o r a piece o f n e t was ad d ed as substrate. T he oxygen
tension o f the w ater entering an d leaving th e cham b er w as m o n ito red co n tin u ­
ously by two R adiom eter E 5047 electrodes connected to tw o R adiom eter
PH M 71b acid-base analysers a n d a G o erz S ervogor 220 recorder. T h e therm ostatted w ater was pum ped via a C ole-P alm er peristaltic pum p and tygon
tubing was used. T he flow th ro u g h th e ch a m b er w as ad ju sted to give a M 0 2 o f
around 15% o f the available oxygen, w h at norm ally m ean t a flow o f 1.2— 3.2
m l-m in- 1 , depending on the size o f the shrim p. C o n tro ls were run before and
after each experim ent w ith su b strate in the ch am b er b u t w ithout the anim al.
T he shrim p was allowed to recover from handling a n d to get used to the
respiration cham b er fo r a couple o f h o u rs b efore th e a ctu a l experim ent started.
A plastic sheet prevented the shrim p from receiving any visual stim uli from the
surrounding. Each experim ent was run, unless otherw ise stated , for at least 24
hours. Lower, stable, PwO , were o b ta in e d b y blow ing N , th ro u g h the reservoir
water.
400
1000
3000
m g w e t w e ig h t
M A T E R IA L A N D M E T H O D S
Crangon crangon and Palaemon adspersus were collected in August— Septem­
ber in the Rockilde-and Isefjord area, N orth ern Sealand, (S, 20%o; T, 15— 20°C)
on sandy Zostera covered b o tto m s a t d epths o f 0.5— 1.5 m by m eans o f
a sm all handoperated trawl. In the la b o ra to ry the shrim ps were stored (S,
20%o; T, 20°C) for at least 3— 4 days before experim entation. M ysids were
provided as food.
200
Fig. I . Crangon crangon (o) a n d Palaem on adspersus ( • ) . R o u tin e o xygen co n su m p tio n
rate as a fu n ctio n o f wet w eight (M O , = aW b) (log— log scale). S, 20%o; T , 20°C.
The general expression M 0 2 = aW b relating respiration and weight was for
Crangon M 0 2 = 0.19 W 0-78 (r = 0.80) an d for Palaemon M 0 2 = 0.28 W 0-85
(r = 0.84). The coefficient b is well within the range of the general coefficient
for all crustaceans (b = 0.81; W inberg, 1950). The constant a gives a species
difference o f 0.09 and this difference and the difference in b are responsible for
the higher weight specific M 0 2 for Palaemon. As is seen from Fig. 1 Palaemon
h ad a M O , 0.15— 0.3 ml 0 , g - l h -1 higher than for Crangon o f sim ilar w eight.
D ie l n o r m o x ic M O,
Palaemon show ed very sm all diel deviations from the 24-hour m ean M O ,
(Fig. 2) and thus no clear diel respiratory pattern was evident for this species.
Crangon, on the other h and, (Fig. 2; 4 individual 24-h cycles chosen fo r each
species, the rest showed the sam e p attern, n = 12 for Palaemon, n = 12 for
Crangon) showed a clear diel respiratory rhytmicity where lowest M 0 2 occurred
during the day (September— October: 8 to 17 o’clock). This M 0 2 was generally
ca. 15% lower, in extrem e cases 40% lower, than the 24 h o u r m ean. T he
highest M 0 2 w as consequently found during night-time w ith values up to 30%
higher than the corresponding 24 h o u r mean. The shrim ps did m oult occasio­
nally during the norm oxic experim ents, increasing the M O , du rin g the ecdysis
w ith around 16%.
71
70
C ra n g o n
30 9b
50 9b
£
7 ~ \ —i
•S
c
60 9b
c»
fi
P O , 7 0 o/o
Tim « of th a day
6
12
T im e of th e day
Fig. 2. Crangon crangon a n d P alaem on adspersus. D iel v a ria tio n s in M O , fo r four
specim ens o f each species u n d e r n o rm o x ic co n d itio n s. M O , expressed a s in teg rated
p ercentage h o u rly d ev iatio n s from the 24 h o u r m ean. S, 20%o; T , 20°C.
Fig. 3. Crangon crangon. E xam ples o f diel v ariatio n s in M O , u n d e r hypoxic con d itio n s
( % sa tu ra tio n ). xMO, expressed as in teg rated percentage h o u riy d ev iatio n s fro m the 24
h o u r m ean. S, 20%o; T , 20°C.
D ie l h y p o x ic M O,
A t a w ater oxygen sa tu ra tio n (Pw0 2) o f less th a n 70% , the diel respiratory
pattern o f Crangon was d isturbed an d only m inor deviations from the 24 h o u r
m ean occurred (Fig. 3). A covariance analysis also show ed th a t the 24 h o u r
m ean in P 20 , 70, 50, and 35% sa tu ra tio n was higher th an the corresponding
norm oxic 24 h o u r m ean M 0 2 (p < 0.01). In P w0 2 70% M 0 2 increase was 10 to
50% (from for instance 0.25 to 0.37 ml 0 2g -1 h - * for a Crangon of 0.56 g w.w.).
In Pw0 2 50% the increase was betw een 20 to 90% (from for instance 0.26 to
0.49 ml 0 2g - 1 h -1 fo r a Crangon o f 0.47 g w.w.) an d in P w0 2 30 to 35% it was
16 to 50% o f the 24 h o u r m ean norm oxic M O ,. T his M O , change corresponds
to the change found earlier w hen Crangon changes its activity level from ro u ti­
ne to swim m ing (H agerm an an d Szaniaw ska, 1986). U nder hypoxic conditions
longer th an 24 h o u rs (up to 72 hours) M 0 2 rem ained at the sam e level as
during the first diel period. M o u ltin g did n o t occur during the hypoxia experi­
ments.
M 0 2 u n d e r h y p o x i a ( d e f i n i t i o n o f th e P J
T he possibility to m ain tain M O , independent o f P w0 2 w as tested in a series
o f short-term experim ents w here the Pw0 2 was low ered 5— 10% per hour. The
percentage change from th e n orm oxic M 0 2 fo r th e tw o shrim p species are
show n in Fig. 4. Both Crangon an d Palaemon show ed a considerable ability to
co n tro l th eir resp iratio n irrespective o f the PwO ,, Crangon dow n to a Pw0 2 o f
©
s %
P, 02, **>
Fig. 4. Crangon crangon (o) a n d Palaemon adspersus (•). R elative changes in ro u tin e
M O , w ith decreasin g oxygen ten sio n o f th e w ater. T he big half-filled d o t a t P wO ,
100% indicates th e relativ e M 0 2 for settled anim als. S, 20%o; T, 20°C.
20% , Palaemon dow n to a P w0 2 o f 30% . This ability seem ed independent o f
size, even if, o f course, the M O , varied according to the size o f the individual.
Both Crangon an d Palaemon show ed an increase in M 0 2 w hen P w0 2 decreased
from norm oxia.
In Fig. 4 this increase is n o t so evident, because in these short-term experi­
m ents the specim ens were n o t allow ed to settle dow n long enough in the respi
ration cham ber. Settled anim als have a respiration ca. 25— 30% low er th a n
non-settled anim als (H agerm an an d Szaniawska, 1986). T hus the starting point
in Fig. 4 a t PwO , 100% should be placed lower for settled anim als w hich w ould
m ake th e hypoxia M 0 2 increase evident.
73
This increase occurred for Palaem on in the Pw0 2 80— 30% , for Crangon in
the P w0 2 50— 30% . F o r Crangon this Pw0 2 corresponds to the em ersion res­
ponse (Hagerman and. Szaniawska, 1986) and for Palaemon to the range where an
increased locom otory activity has been noted (H agerm an an d W eber, 1981;
von Oertzen, 1979).
D IS C U S S IO N
In the present w ork it w as fo u n d th a t Palaemon adspersus always h ad
a higher M 0 2 than sim ilar sized Crangon crangon. In n atu re both species are
subjected to roughly sim ilar variations in the environm ental conditions (S, T, O),
b u t inhabit different biotopes. T he burying Crangon alw ays is associated w ith
sandy bottom s while Palaemon inhabits algae an d plants and thus does not
have the ability to bury in a su bstrate. Bridges a n d B rand (1980) n o te d general­
ly low er M O , in burrow ing crustaceans com pared to non-burrow ing species.
T his is probably also valid fo r o th er groups, for instance de W ilde (1973)
found reduced M O , in the b uried bivalve M acom a com pared to epifaunal
bivalves. R eduction in M 0 2 has also been noted intraspecific in buried com pa­
red to non-buried horseshoe crab s (Johansen a n d Petersen, 1975) a n d in buried/non-buried Nephrops norvegicus (H agerm an and U glow , 1985) and recent­
ly in Mesidoihea entomon (H agerm an a n d Szaniaw ska, in prep.).
A typical routine respiratory level fo r interm oult Crangon (w.w. 0.25 and
1.1 g) und er norm oxic co nditions are 0.9 an d 0.2 m l 0 2g - I h -1 in the present
investigation. E arlier w o rk o n th e resp iratory rates o f Crangon has show n
values slightly higher th an those found here (H agerm an, 1970) o r in the same
range (van D o n k a n d de W ilde, 1981). T he differences are often caused by the
different techniques used, fo r instance H agerm an (1970) used a closed respira­
tory system and the experim ents were short-term (a couple o f hours) where
handling and o th er types o f stress caused a respiration higher th an th a t show ed
by an anim al adap ted to a long term experim ent, preferably not less th an 24
hours. It has been show n also by v o n O ertzen (1975) a n d by K ristensen (1983)
th a t the M 0 2 for Gammarus and and fo r Nereis were m uch higher in short-term th an in long-term experim ents. In this connection the necessity o f a co r­
rect substratum m ust be stressed if a reliable value o f M 0 2 is to be obtained. If
Crangon has no suitable substrate, in n atu re corresponding to displacem ent to
a rocky o r h ard substrate, th e diel lo co m o to ry rhythm icity breaks dow n (H a­
germ an, 1970) and the respiratory ra te increases to 1.6 an d 0.65 ml O , g - 1 h -1
fo r th e specim ens m entioned above. R eg n au lt an d L agardère (1983) observed
th a t an increased noise level (co m p ared to the b ackground noise always prevai­
ling a t sea) increased the m etabolic rate in Crangon w hen m easured both as
am m onia excretion and as oxygen consum ption. A com m on response to all
kinds o f disturbancies is certainly a m odification o f the activity level an d rhyth­
micity resulting in an increased M 0 2.
Typical routine M O , for Palaemon adspersus in the present investigation
were (wet weight 0.7, 1.0 and 2.1 g) 0.65, 0.41 and 0.13 ml 0 2g -1 h -1 un d er
normoxic conditions. These values can be com pared to earlier findings o f M 0 2
of 0.15 for an individual o f 2.5 g w et w eight (H agerm an and W eber, 1981) and
1.35 pi 0 2g D W _1h _1 as found by von O ertzen (1984) u n d er sim ilar environ­
mental conditions.
In Crangon the diel norm oxic M O , can be correlated to th e diel locom oto­
ry pattern (H agerm an, 1970), i.e. M 0 2 shows peaks during the same diel pe­
riods as Crangon is active. Crangon is a voracious predator, feeding m ainly ju st
after daw n an d before dusk (Pihl a n d R osenberg, 1984). T he locom otory acti­
vity of Crangon is m ainly influenced by exogenous factors (H agerm an, 1970)
and as the m etabolic cost o f sw im m ing is low (H agerm an and Szaniawska,
1986) the noted respiratory rhythm icity is probably endogenous in origin, ref­
lecting digestive and endocrine processes in the animal.
F o r Palaemon adspersus the absence o f a pronounced diel rhythm icity will,
despite a nocturnal locom otory activity (H agerm an and 0 s tr u p , 1980) reflect
endogenous processes at, how ever, a higher rate th a t in Crangon. The latter
can in the respiration cham ber bury in the sand, while Palaemon always will be
more exposed to external stim uli an d thus keeps its m etabolic state at a m ore
vigilant level. Sm it (1965) noted in goldfish an increased M O z th a t was not
reflected in activity changes an d H alcrow and Boyd (1967) show ed that exci­
ted, but not m oving, Gammarus had a higher M 0 2. In addition m easurem ents
of heart rate has show n a large excitem ent dependent variability (Dyer and
Uglow, 1978) in shrim ps an d crabs an d these authors also show ed th at heart
and ventilatory rates, an d thus M 0 2, decreased when specim ens buried.
It is now generally accepted th a t m ost crustaceans th a t in nature m ight be
exposed to v ariation in external PwO , have developed an ability to m aintain
respiratory independence during hypoxia. Such an ability is also found in som e
molluscs where T aylor and B rand (1975) found an increase in the ability o f
bivalves to m aintain oxygen independence w ith increasing size during hypoxia.
Fam m e and K ofoed (1980) noted th at the M O , of M ytilus to a certain extent
was independent o f the Pw0 2.
The PwO , where oxygen consum ption independence no longer can be m ain­
tained is usually defined as the so-called Pc (Newell, 1979; H erreid, 1980). In
the present investigation the M 0 2 o f Palaemon and Crangon rose during prog­
ressive hypoxia, specially w hen a P w0 2 o f ca 45— 60 to rr was reached. T his is
interpreted as an increase in the activity w hen the anim al perceives an d /o r
reacts to the decreasing Pw0 2 a ro u n d 75— 60 torr. Below this the M O , is still
high due to the stress caused by the still decreasing Pw0 2. T he Pc, as defined
above, is for Crangon obviously a t a very low Pw0 2 (ca 30 to rr). How ever, this
point reflects only the PwO , w here oxygen independence can no longer be
m aintained, i.e. where the active M 0 2 is decreasing. O xygen supply to the
tissues for routine/basal m etabolism is still adequate and the P c is in this sense
thus n o t critical as the anim al can sustain these conditions for extended pe-
nods. However, when the oxygen supply fo r the basic m etabolic needs is distur­
bed, i.e. when anaerobic m etabolism is initiated, a real critical p o int („P,actate” )
is reached. This level is ca 15 to rr in Crangon (H agerm an and Szaniawska,
1986) where, as a result o f an aerobiosis, lactate is beginning to accum ulate. In
Crangon, a species show n to be to leran t to hypoxia, the trad itio n al Pc and the
p oint where anaerobic m etabolism accelerates,are very close to each o ther but
there is an evident difference in the level o f M O r H ow ever, under in ap p ro p ria­
te experim ental conditions o r w ith d isturbed anim als the traditional Pc will
certainly m ove upw ards (H erreid, 1979; B utler et al., 1978; Dali, 1986). A nalo­
gically to this the Pw0 2 for the trad itio n al P c in Palaemon was ca.45 to rr while
the level where an anaerobic m etabolism was found to increase was 30 torr.
Both values are higher th a n for Crangon but well in agreem ent with the ecology
of the species.
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