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Hartman Brandt 1995

Hartman Brandt 1995

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Tranxaciitmx
oftheAmerican Fisheries Society
124:347-355,
1995
© Copyright by
the
American Fisheries Society
1995
Estimating
Energy
Density
of
Fish
KYLE
J.
HARTMAN'
Chesapeake Biological Laboratory,
The
University
of
Maryland SystemPost
Office
Box 38,
Solomons,
Maryland
20688-0038,
USA
STEPHEN
B.
BRANDT
Great Lakes Center,
Buffalo
State College1300 Elmwood Avenue.
HC
215,
Buffalo,
New
York
14222,
USA
Abstract.—
The
use of bioenergetics models in fisheries ecology and management has increased
rapidly in
recent
years,
but
application-specific information
on
energy content
of
fish
and
theirpreyhaslagged behind.Webelieve thisis
because
the
process
ofdirectly measuring energy density
is
very time consuming.
In
this paper
we
present
and
evaluate
a
series
of
general empirical models
that
predict energy density (J/g
wet
weight)
from
fish
percent
dry
weight. Data were gathered fromthe literature, obtained from cooperating investigators, and measured directly. Least-squares modelswere derived for all
species
combined and for
orders,
families,
and
species.
All
models were linear;however, logarithmic transformation was
necessary
to normalize residuals in the combined model.
All
models
with
more than
four
data points were
highly
significant
(P <
0.002)
and had
coefficientsofdetermination
of
0.76
or
greater.
The
model
for all
species
combined
(N
=
587,
r
2
=
0.95)wasJ/g wetweight=
45.29
DW
1
-
507
;
DW is thepercentdryweightof the
fish.
At alltaxonomiclevels, energy density models showedastrong positive relationship between energy densityandpercent
dry
weight. This relationship,
if
corroborated, should allow
the
estimation
of
seasonal
and
ontogenetic changes in energy density based solely on percent dry
weight
data.
The use of
bioenergetics models
in
fisheries sci-ence
and
ecology
has
increased rapidly
during
thelast
10-15 years. Early models relied heavily onthe work of Winberg
(1956),
Kitchell et al. (1974,1977),
and
Brett
and
Groves
(1979).
Recently,
bio-energetics models have been used to answer spe-
cific
management
and
ecological questions.
For
example,
models have been used
to
establish
the
stocking rates
of
predators (Stewart
et al.
1983;StewartandIbarra
1991),
to examine the potential
effectsof
global warming
on
fish
production
(Hill
and
Magnuson 1990), to evaluate predation as a
factor
in
recruitment
failure
(Hartman
and
Margraf
1993),
to estimate bioaccumuiation of organic pol-
lutants
in
fish
(Barber et al.
1991),
and to examine
how
patchiness
in
thermal
and
prey
fields
mightaffect
fish
growth (Brandt
et al.
1992; Brandt1993).Despite
the
widespread
use of
bioenergeticsmodels
in
fisheries,
information
on one key
input
variable,
energy density (J/g)
of
predators
and
prey,
has
been limited. Although much
has
been
published
on the
energy density
of
fish
on a dryweight
basis, most data have been presented with-
out a
measure
of the
wet-to-dry weight
of the
fish.
1
Present
address:
Great Lakes Center,
Buffalo
State
College,
1300
Elmwood Avenue, HC 215, Buffalo, New
York
14222, USA.
Because bioenergetics models commonly use
field-derived
growth estimates measured
in wetweight,
energy density data must
be
placed
in wet
weight unitsto beapplicableformodeling.
There
are
changes
in the
energy density
of
fish
both with
ontogeny
(Wissing1974;Craig 1977; Minton
andMcLean
1982)
and season (Craig 1977; Adams etal.
1982;
Flath and Diana
1985).
Within a
species,
the
energy content
can
vary
by
water body (Pierce
et al.
1980; Strange
and
Pelton 1987).
If
results
obtained
from
the use of
bioenergetics models
are
to be accurate, estimates of energy density of pred-ators
and
prey
are
required
for
specific seasons,
fish
ages,
and
ecosystems.Bomb calorimetry
is an
accurate method
formeasuring the
energy content
of
aquatic organisms
(Cummins
and
Wuycheck
1971) and is
preferableto estimates based on proximate analysis (Craig et
al.
1978).
However, bomb calorimetry
is a
time-
consuming
process that requires
0.75-1.0
h/sam-ple (Parr Instrument Co. 1960). Flath and Diana
(1985)
observed
a
negative relationship between
the
water content
of
alewives
Alosa pseudohar-engus
and the caloric content (wet weight basis),
which
is due to the inverse relationship betweenpercent
lipid
and
percent body water
in
fishes
(Craig et al.
1978;
Flath and Diana
1985).
We be-
lieve a
relationship between
the
percent
dry
weight(100-dry weight/wet weight) and energy density
347
 
348
HARTMAN AND BRANDT
(J/g
wet weight)
exists
for
most
fish
species,
and
we hypothesized that a
general
model
fitted
to thisrelationship
across
taxonomic groupings
would
permit estimation
of
energy
density
of
most
fish.
We
further identified
the limitations of this
rela-
tionship
by examining the effects of using
energy
density
models
on
bioenergetics
modelresults.
Methods
Calorimetry.—
Striped
bass
Morone
saxatilis,
bluefish
Pomatomus
saltatrix,
weakfish
Cynoscion
regalis,
and bay anchovy
Anchoamitchilli
were
collected
from
the
Chesapeake
Bay, Maryland, be-
tween June
1990
and
February
1992.
Fish
wereplaced
in
bags
and
iced
(striped bass,
bluefish,
and
weakfish)
orfrozenin
water
(bay
anchovy)
to
slow
decompositionand
prevent
water
loss.
In the lab-oratory,
fish
were weighed wet
0.1
g) and
ovendried to constantweightat
70°C.
Each
fish
wasthen
weighed and the percent dry weight
(DW)
for
each
fish
was
calculated.
Foreach
species,
11-26
individuals
werese-
lected
for
calorimetric
analysis
with
standard Pan-oxygen bomb techniques
(Parr
Instrument
Co.
1960;
Craig
1977).
Individuals
were
selectedtocover
the
entire
range
of DW and
fish
sizes
we
recorded
for
each
species.
For
each
fish
(several
individuals
for bay anchovy), the dried
carcass
was
homogenized,
and two
1.0-g
subsamples
werecombusted
to
measure
energy
content.
If the sub-
samples
differed
by
more
than 10% of the
mean
energy
density
(J/g),
a
third
subsample
wascom-busted.
The
mean
of 2-3
subsamples
was
used
for
estimates
of
energy content
for
those
fish.
General
models.—
Indeveloping a general mod-
el for
predicting
the
energy
density of
fish,
we
used
data
from
our
research,
from
peer-reviewed liter-ature (since
1973),
and
from
cooperating
investi-
gators.
Alldata
were
treated as
independent
datapoints,
regardless
of the
taxonomic
level.
Only
en-
ergy contents
for dryweight
were used
becausesome
data were
not
reported
as ash
free.
In
all,
587 data points representing 35
species,
21
fam-
ilies,and 8
orders
were
included
in thegeneral
model data set (Table1).For the
purposes
of
dis-
cussion,
"combined
model"
refers to the energy
density
model
derived
from
all
fish.
Each order
model
included
data
from
fish
of
that
specificor-
der.
Similarly,
each
family
or
species
model
con-tained
only
data
from
fish
of a
single
family
or
species.
Least-squares
regression
modelsweregenerated
with
energy density as the
dependent variable
and
percent
dry weight is the independent
variable.
Theempirical modelforenergy densitywasED
=
a
+
6-DW,
(1)
where DW is the
percent
dry
weight
of the
fish,
ED is the
energy density(J/g
wet weight) and
a
and
b
are
empiricallyderived
constants. For the
combined energy density
models,
logio transfor-mation
of
data
was required to normalize the re-siduals.
Thus,
the
model
for all
species
combined
was
ED
=
a-DW*.(2)Although
a general relationship between energy
density
and percent
water content
was
expected,
differences in
handling
and
processing
of
samples
amonginvestigators probablyintroduced
errors
into the
estimates.
Our
initial
criterion
for
model
acceptance
was
arbitrarily
selected
as a
coefficient
of determination
(r
2
)
greater
than
0.75.
Prior to model development, three
randomly
se-lected data
sets
(with
N
>
15) were set
aside
for
testing
the
models.
Once
models
were
accepted
they
were tested
with
the
data
sets
for bay anchovy
(Hartman
1993),
rainbow
trout
Oncorhynchus
mv-
kiss
(From and
Rasmussen
1984),
and
yellow
perch
Percaflavescens
(Hayesand
Taylor
1994).
These
independent data
sets
were
not included in
any
other energy
density
regression models.
In-
dependent data
sets
were
fitted
to
model
(1) and
compared
with
the
combined,
order,
and
family
models
by analysis of
covariance
(ANCOVA) to
determine
if
slopes
and
intercepts
among
models
and independent
data
sets
were
different
(a =
0.05).
Bioenergetics simulations.
To evaluate
the
util-
ity
of
general models
of
energy
density for
bio-
energetics
modeling, we ran
simulations
with
es-tablished
bioenergetics
models
for bay
anchovy
(Luo
and
Brandt
1993),
steelhead
(anandromusrainbow trout;
Rand
etal.
1993),
and
yellow
perch
(Kitchell
et al.
1977).
We
used
theHewettand
Johnson
(1992)
software
to
implement
the
bio-
energetics
models for
these
species
and used a
standard set of data for
seasonal
weights
(Table
2)and
temperatures
(Figure
1)
specific
to
each
spe-cies.
Energy
densities of
prey
for each predator
were considered
aconstant
value
that
did not
vary
by
season.
Prey
energy
densities were2,552
J/g
wet weight for bay
anchovy (Luo
and
Brandt
1993),
5,699
J/g wet
weight
for
steelhead
(Rand
et
al.
1994),
and
3,766
J/g wet
weight
for
yellow
perch
(Hewett and Johnson
1992).Toevaluate therelative
importance
of
seasonal
 
ESTIMATING
ENERGY
DENSITY
349
TABLE
I.—Energy
densitymodels
for estimating
energy
content (J/g
wet weight)
from
percent
dry weight (DW)
dataand the
data
sources.
All
models
are of the
form:
J/g wet weight =
a
+
6-DW,
except
model 2,
which
is of the
form:
J/g wet
weight
=
0-DW^,
where
a
and
b
are
empirically
derived
constants.
Model
number
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
2324
25
26
27
28
29
30
31
32
3334
3536
37
38
39
Model
taxonCombined
(linear)
Combined(logic)Clupeiformes
Perciformes
Cypriniformes
SalmoniformesPleuronectiformes
ClupeidaeSciaenidae
Cyprinidae
Percichthyidae
OsmeridaeSalmonidacCottidae
Alosa pseudoharengus
Brevoortia
ryrannusCatostomus commersoniCoregonis
clupeaformis
C
hoyiCottus
cognatus
Cynoscion
regalu
Dorosoma cepedianumEsox masquinongyLuxilus
cornutus
Morone chrysops
M.
saxatilis
Oncorhynchus mykissO. kisutchOsmerusmordax
Phoxinus phoxinus
Pleuronectes
aspefi
Pomatomus saltatrix
Salvelinus
namaycush
Stizostedion lucioperca
Theragra
chalcogramma
Cottus cagnatus
Anchoa mitchilli
PercafiavescensOncorhynchus mykiss
Species
included
Culaea
inconstantFundulus
heteroclitus
F.
majalis
Gobiosoma bosc
Lagodon rhomboides
Leiostomusxanthurus
Micropogon undulatus
Mugil
cephalus
Myoxocephalus
quadricornisParalichthys
dentatusSalmo
trutta
Symphurus
plagiusa
r*
0.950.950.850.880.950.970.91
0.85
0.920.86
0.91
0.%0.%
0.800.930.89
0.970.96
0.91
0.870.97
0.990.93
0.94
0.990.93
0.99
0.97
0.%
0.82
0.91
0.98
0.97
0.97
0.880.87
Data
sets
not
0.77
0.82
0.90
N
587
587
82
80
38348
2482
13
33
19
25
242
10
5325
5
5
37
6
10
4
8110
4
1519
7225
23
20
13
108
19
11
6
a
-3,419
45.29
-2,532
-1,875-1,265
-3,386
-1,832
-2,532-1,936
-981
-2,533
-1,094
-3,632
-1,498
-2,086
-2.695
232
-4,870
-2,424-3,418
-1,997
-4,155
-1,939-1,026
-1,932
-1,460
-2,735-3,207
-1,094-4,237
-1,895
-3,792
-3,809
-2,011-4,210
-3,418
b
375.0
1.507
328.6309.5
262.2
379.0
286.1
328.6
309.9
251.1
349.1
303.2
386.7
306.0
323.7
309.0
143.0
424.9
336.2385.2
319.4
390.9294.5260.5
293.8
313.9
357.5
367.8
303.2
374.9
290.0
372.4
397.9
309.4
401.1
385.2
Data
source
11
1^1,6,8-17
1-4,6,8-17
1,4,
11-13.
15. 16
1,2,9.
17
3,8
8,
11.
13, 14,
16
10,16
4,11-13,15,16
1.16
3,8
1,
1711,
16
5, 13,
1413
4,
11.
13
12.
16
8
13
13.14
13
1
15
8
8
17
I
13
13
11,16
3
10
1
13
2
6
13
included in
combined
model
26
55
35
691
-2,873-2,457
156.3
313.1
347.8
1
7,9
5
in
combined
model,
bat
with
too few
data
pointsfor
analysis
2
111
3
2
1
2
4
2
1
2
9
16
1616
16
16
161613
16
13
16
a
Sources:
(1)
This
study
(2)Buijseand
Houthuijzen
(1992)
(3) Cui and
Wootton(1988)
(4)
Rath
and
Diana
(1985)
(5)
From
and Rassmussen
(1984)
(6)
Harris
et
al. (1986)
b
Formerly
known
as
Limanda aspera.
(7) Hayes and Taylor
(1994)
(8) J.Jonas (personal
communication)
(9) Kelso(1973)
(10)
A. J.
Paul
(personal
communication)
(11)
Rand
el al.
(1994)
(12)
Rippetoe(1993)
(13) Rottiers and
Tucker(1982)
(14) Rudstamet
al.
(1994)
(15)
Strange
and Pelton (1987)
(16)
Thayeretal.(1973)
(17)
Wissing(l974)

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