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Seasonal Variation in Energy

Seasonal Variation in Energy

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Journal of Fish Biology
(2001)
59,
380–389doi:10.1006/jfbi.2001.1649, available online at http://www.idealibrary.com on
Seasonal variations in the energy density of fishes in theNorth Sea
J. P

*‡

J. R. G. H

*Danish Institute for Fisheries Research, North Sea Centre, P.O. Box 101, DK-9850Hirtshals, Denmark and 
Fisheries Research Service, Marine Laboratory, Victoria Road,Aberdeen, AB11 9DB, Scotland 
(
Received 30 September 2000, Accepted 2 May 2001
)
The energy density (
D
, kJ g
1
wet mass) of saithe
Pollachius virens
, haddock
Melanogrammusaeglefinus
, whiting
Merlangius merlangus
, Norway pout
Trisopterus esmarki 
, herring
Clupeaharengus
, sprat
Sprattus sprattus
, sandeel
Ammodytes marinus
and pearlsides
MaurolicusMuelleri 
, from the North Sea, increased with total length,
L
T
. However, there was not alwaysa significant (
P>
0·05) linear relationship between
L
T
and
D
. Seasonal di
ff 
erences in
D
wereobvious in mature fish, while geographical di
ff 
erences were insignificant. For all species therewas a highly significant correlation (
P<
0·0001) between the percent dry mass of the fish (
D
S
)and
D
. A general relationship was established for gadoids and sandeel
D
=
3·1492+0·3459
D
S
and herring
D
=
4·6395+0·4170
D
S
. Thus seasonal and size-specific data on
D
neededfor bioenergetics and gastric evacuation models can be determined simply from
D
S
, which isconsiderably less costly and time consuming than calorimetry or proximate analysis.
2001 The Fisheries Society of the British Isles
Key words: ash content; energy density; dry mass; season; fish length.
INTRODUCTION
The use of bioenergetics models in fisheries science and ecology has increasedrapidly during the last 10–15 years (Brandt & Hartman, 1993)and there is increasing interest in the use of models to study interactions between seabirds,other top predators and fishes(Hislop
a
). Often these models arebased on energy transfer between predator and prey. In such models seasonalchanges in energy density of predators and their prey are important variables.Energy transfer is not used in methods such as multispecies virtual populationanalysis (MSVPA), which has been used to assess some North Sea fish stocks(Sparholt, 1990). Instead, food consumption is estimated by combining fielddata on a predator’s stomach content mass with experimental data on therelationship between gastric evacuation rate and stomach content (Bajkov, 1935;Elliott & Persson, 1978;Pennington, 1985). There is uncertainty which is the best model to describe gastric evacuation (Hislop
b
)althoughexponential models are generally used. However,Andersen (1998,1999) has shown that gastric evacuation of gadoids can be described independently of mealsize by a square root model and that in whiting
Merlangius merlangus
(L.) preysize has no e
ff 
ect on the rate of gastric evacuation. The gastric evacuation rate
‡Author to whom correspondence should be addressed. Tel.: +45 33 96 32 00; fax: +45 33 96 32 60;email: jp@dfu.min.dk3800022–1112/01/080380+10 $35.00/0
2001 The Fisheries Society of the British Isles
 
in whiting fed natural prey is, however, a
ff 
ected by the energy density of the preyitems (Andersen, 1999). UsingAndersen
s (1999)model,Pedersen (2000)found that the speci
c daily rations of whiting were signi
cantly di
ff 
erent between yearsand groups of whiting, depending on the energy density of their stomachcontents. Thus seasonal energy density data are also required for the applicationof the gastric evacuation model. However, information on the energy density of predator and their prey suitable for use in bioenergetics models is sparse.Although many data are available (Cummins & Wuycheck, 1971;Harris & Hislop, 1978;Hartman & Brandt, 1995)they can be misleading, because they do not take into account that there are changes in energy density of 
shes both withontogeny (Craig, 1977) and season (Kelso, 1973;Foltz & Norden, 1977;Hislop
a
). Therefore, if results obtained from models using energy data areto be accurate, determinations of energy density of predator and prey arerequired for seasons and
sh size or age.The present study is an attempt to expand former studies and to determine byseasons and
sh size the energy density of eight species for which there arequanti
ed predator-prey interactions in the North Sea.
MATERIALS AND METHODS
For determination of energy density, ungutted specimens of the piscivorous speciessaithe
Pollachius virens
(L.), whiting, haddock
Melanogrammus aegle
 fi
nus
(L.), and theirmain
sh prey, herring
Clupea harengus
L., sprat
Sprattus sprattus
(L.), Norway pout
Trisopterus esmarki 
(Nilsson), sandeel
Ammodytes marinus
(Raitt) and pearlsides
Maurolicus muelleri 
(Gmelin), were collected by research and commercial vessels in thenorthern and central North Sea during the period 1996
 – 
1998. The
shes were collectedquarterly, Q1=January, February or March; Q2=April, May or June; Q3=July, Augustor September; Q4=October, November or December. At sea, the
shes were sorted intolength classes (total length,
L
T
), whose boundaries varied with species and
L
T
(Table I).Each
sh or batch of 
sh in the case of small individuals was wrapped in a polythene bag,to minimize water loss, and stored in a domestic freezer at
20
C. To minimizedesiccation the
shes were processed within 6 months after capture. However,
shesshowing evidence of desiccation were not used.All the saithe, whiting, herring, sprat, Norway pout and pearlsides were processed atthe Danish Institute for Fisheries Research (DIFRES), North Sea Centre, Hirtshals,Denmark. Prior to determination of energy density the samples were warmed at roomtemperature to a partially thawed state, which prevented loss of water and blood from the
shes. The
shes were roughly chopped and homogenized. Fishes >20 cm
L
T
werehomogenized in a Tecator Mill 1094 homogenizer for 4 min and subsequently in a Bu
¨
chiMixer B-400 for 15 s, while smaller specimens were processed
rst in a domestic foodblender and then in the Bu
¨
chi mixer. Loss of water during processing was insigni
cantfor all size classs. Maximum loss (0
·
5% wet mass) was observed when
shes >20 cm
L
T
were chopped. For all size classes masses derived in this manner were therefore assumedto be reliable and not in
uence by loss of water.Two samples of the homogenate weighing
c
. 20 g each were oven-dried to constantmass at 60
C (48 h) for the determination of dry mass (
D
) content and afterwardsheated for 48 h at 600
C for determination of the ash mass. Two other samples wereoven-dried and two 1
·
0
 – 
1
·
3 g
D
subsamples were combusted to measure energy densitywith an IKA-Calorimeter C 7000. If the subsamples di
ff 
ered by >1
·
5%, the variation wasabove the precision of the calorimeter and a third subsample was combusted. Theaverage of the two to three subsamples was used for estimates of energy density forthose
sh.The haddock and sandeel samples were processed at the Marine Laboratory,Aberdeen, Scotland. Homogenates were prepared using procedures similar to those
   
381
 
T

I. Energy density (kJ g
1
W
) of the common
sh species in the North Sea by sizeand quarter (sample size in parentheses). Bold and underlined
gures are fromHislop
a
)andHarris & Hislop (1978).Medio and ultimo refer to
sh caught between1
 – 
5 February and after 21 March, respectivelySpeciesSize class(mm)Quarter1 (medio) 1 (ultimo) 2 3 4Saithe 200
 – 
249 4
·
1 (4)Saithe 250
 – 
299 4
·
7 (6)Saithe 300
 – 
349 4
·
7 (10)Saithe 350
 – 
399 4
·
2 (20) 4
·
5 (20) 4
·
8 (20) 4
·
9 (20)Saithe 400
 – 
449 4
·
6 (20) 4
·
4 (18) 5
·
1 (20) 4
·
8 (20)Saithe 450
 – 
499 5
·
0 (19) 4
·
0 (20) 5
·
3 (20) 4
·
8 (20)Saithe 500
 – 
599 5
·
5 (9) 4
·
0 (9) 5
·
6 (10) 5
·
2 (10)Saithe 600
 – 
699 5
·
3 (10) 3
·
5 (9) 6
·
8 (10) 6
·
2 (5)Saithe 700
 – 
799 5
·
0 (7) 4
·
8 (8) 6
·
0 (10)Saithe 800
 – 
999 5
·
3 (10) 6
·
3 (10) 6
·
2 (2)Saithe 1000
 – 
11 999 5
·
3 (2) 6
·
0 (4)Haddock 100
 – 
119 3
·
9 (20)Haddock 120
 – 
149 3
·
6 (21) 3
·
3 (10) 4
·
2 (10) 3
·
6 (10)Haddock 150
 – 
199 3
·
7 (22) 3
·
9 (10) 4
·
1 (10) 4
·
1 (30)Haddock 200
 – 
249 4
·
3 (14) 4
·
3 (10) 4
·
2 (10) 4
·
9 (14)Haddock 250
 – 
299 4
·
7 (11) 4
·
0 (10) 4
·
6 (18) 4
·
9 (12)Haddock 300
 – 
349 4
·
7 (14) 4
·
5 (25) 5
·
0 (10) 5
·
2 (23)Haddock 350
 – 
399 4
·
7 (14) 3
·
6 (5) 5
·
5 (11) 5
·
5 (18)Haddock 400
 – 
449 4
·
5 (11) 3
·
8 (7) 4
·
9 (11) 5
·
1 (11)Haddock 450
 – 
499 4
·
4 (5) 5
·
3 (2) 5
·
3 (2)Haddock 500
 – 
549 4
·
7 (1) 5
·
6 (5) 5
·
3 (1)Haddock 550
 – 
599 3
·
6 (1)Herring 40
 – 
49 4
·
1 (14)Herring 50
 – 
59 4
·
2 (35)Herring 60
 – 
79 3
·
9 (39)Herring 80
 – 
99
4
·
6
4
·
5 (20)
4
·
6
Herring 100
 – 
119 4
·
7 (20) 4
·
6 (2) 4
·
4 (5)
4
·
6
Herring 120
 – 
149 4
·
4 (21) 4
·
5 (50) 5
·
2 (20) 6
·
3 (23)Herring 150
 – 
199 4
·
4 (20) 4
·
4 (50) 10
·
1 (20) 7
·
1 (20)Herring 200
 – 
249 6
·
5 (20) 5
·
7 (41) 11
·
0 (20) 8
·
5 (12)Herring 250
 – 
299
8
·
5
4
·
9 (20) 11
·
9 (20) 8
·
8 (20)Herring 300
 – 
349Sprat 43
 – 
93 6
·
7 (2)Sprat 104
 – 
137 10
·
9 (3)Sprat 115
 – 
125 11
·
5 (20)Sprat 110
 – 
119 6
·
4 (6)Sprat 120
 – 
129 5
·
8 (10)Sprat 130
 – 
139 5
·
9 (10)Sprat 140
 – 
149 5
·
6 (6)Pearlsides 45
 – 
72 7
·
3 (50)Norway pout 40
 – 
49 3
·
8 (50) 3
·
9 (20)Norway pout 50
 – 
59 3
·
8 (50) 3
·
9 (20)Norway pout 60
 – 
79 3
·
9 (20)Norway pout 80
 – 
99 4
·
5 (1) 4
·
8 (50)Norway pout 100
 – 
119 6
·
2 (43) 4
·
0 (20) 4
·
2 (20) 5
·
7 (18) 5
·
2 (20)Norway pout 120
 – 
149 6
·
4 (14) 4
·
1 (20) 4
·
4 (20) 6
·
3 (20) 7
·
0 (20)Norway pout 150
 – 
199 4
·
7 (35) 4
·
0 (20) 5
·
7 (20) 7
·
0 (20)
382
.
  
.
.
.


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