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Physiological condition of fishes has been defined as the gross nutritional state (Love 1970) and
the level of reserve nutrients, particularly fat, present in the body (Gershanovich et al. 1984). Consequently, chemical body composition of an individual fish should characterize its physiological
condition and, in general, its health. Furthermore,
this physiological status determines the individual's ability to compete successfully (e.g., through
optimal foraging and reproduction), sustain
growth, maintain and repair tissues, and cope with
stresses induced by environmental changes. Energy reserves in fish are primarily expressed as
visceral fat (Love 1970). Changes in chemical body
composition generally reflect storage or depletion
of energy reserves.
Quantitative analyses of primary body constituents of fish have been reported for numerous
marine (Vinogradov 1953) and freshwater species
(Jacquot 1961). Generally, live-weight, whole-body
composition offish is 70-80% water, 20-30% protein, and 2-12% lipid; however, extreme values
for these components may fall well outside these
ranges (Weatherley and Gill 1987). Several studies
have shown significant changes in whole-body
composition or in the composition of specific organs or muscle tissue due to age, diet, feeding frequency, migration, ration, season, sex, starvation,
and temperature (Chang and Idler 1960; Brett et
al. 1969; Groves 1970; Savitz 1971; Niimi 1972;
Elliot 1976; Craig 1977; Grayton and Beamish
1977; Millikin 1982; Weatherley and Gill 1983).
Condition, as applied to fish population ecology, has been described as an indication of fatness,
general well-being, or gonad development (among
other traits) of an individual or a group of individuals (Le Cren 1951). To put it simply, condition is the relative plumpness or well-being offish
(Wege and Anderson 1978). These definitions are
characterized by indices based on empirical
weight-length relationships. The relative condition factor (Kn; Le Cren 1951) provides a measure
for comparison of individual fish weight to a predicted weight derived for that population or subgroup, whereas relative weight (W r ; Wege and Anderson 1978) measures the variation between
individual fish weight and a length-specific standard weight (Ws) for the species in question. Presently, the Wr index is in wide use among fisheries
management agencies (Murphy et al. 1991), primarily because of the comparative attributes of
the index among species and between individuals
of different lengths within a species. Anderson
(1980) suggested that a Wr of 100 (individual
weight = standard weight) may reflect ecological
and physiological optimality; however, little empirical evidence of the relationship of these factors
to Wr has been shown.
The measurement of physiological condition,
determined by comparison with a standard weightlength relationship, may provide reliable estimates for determining body composition of live
fish. Typically, body composition of fish is assessed by chemical proximate analysis, which is
509
510
Experimental design. We obtained 30 juvenile striped bass (mean weight, 184.1 g 33.3
SD) and 30 hybrid striped bass (543.2 g 89.0)
from the San Marcos National Fish Hatchery and
Technology Center, San Marcos, Texas, and
stocked them in 109-L aquaria equipped with siphon drains. Stable conditions were maintained
by a closed recirculation system (total volume,
4,357 L) that delivered water to each aquarium at
a rate of about 1 L/min; partial exchange (50%)
with well water occurred every 3 d. Total ammonia, nitrites, alkalinity, hardness, dissolved oxygen, and pH were measured weekly. Water quality complied with standards suggested for striped
bass culture (Bonn et al. 1976; Lewis and Heidinger 1981; Rogers et al. 1982). Water temperature
was maintained at 24-26C with a chiller unit.
This temperature has produced optimum growth
of juvenile striped bass under laboratory conditions (Cox and Coutant 1981). Fluorescent lighting, controlled by an automatic electric timer, provided a light: dark cycle of 12:12 h.
Fish were fed a pelleted grower diet (Bioproducts, Warrenton, Oregon) in both experiments (4mm-diameter pellet for striped bass; 9-mm-diameter pellet for hybrid striped bass). This diet, a
semimoist extruded feed formulated to produce
rapid growth of salmonids, meets the known nutritional demands of juvenile striped bass (Bonn
et al. 1976; Millikin 1982, 1983; Zeigler et al.
1984). All fish were conditioned for 2 weeks by
feeding them at a rate of 2% (0.05 g) of dry body
weight per day (BWPD). After the pretreatment
period, five randomly selected replicates were each
fed one of six ration levels (0.25, 0.50, 0.75, 1, 2,
511
termined by the macro-Kjeldahl technique (AOAC gain for each experiment (Brown 1957). Total
1984) and converted to total protein equivalents growth efficiency (Et) was calculated biweekly as
(percentage crude protein = percentage nitrogen
x 6.25). Crude fat, crude protein, and ash were
expressed as dry-weight percentages. We calculated gross energy content of whole bodies on ash- G is weight gain (g) and / is dry-weight consumpfree dry-weight samples with standard conversion tion (g) (Warren and Davis 1967). The instanfactors recommended for fat (39.54 kj/g) and pro- taneous (specific) growth rate (IG) was detertein (20.08 kj/g) (Brett and Groves 1979).
mined as
Our threshold of statistical significance was P
< 0.0001 unless noted otherwise. The z-test did
not reveal any significant differences for proximate component replicates for individuals (P >
0.05), so means for duplicates of individuals were WT is the final weight at time T, and Wt is the
used in all analyses. Variances associated with ra- initial weight at time t (T t was about 14 d),
tion treatments were homogeneous, so untrans- derived from the rate expression
formed values were reported. Tests for differences
among ration treatments for final Wn percentage
water, crude fat, crude protein, ash, and visceral YT is the final weight and Yt is the initial weight
fat were based on analysis of variance (ANOVA) (Brown 1957).
Growth efficiency and instantaneous growth rate
and Tukey's W-procedure (Lentner and Bishop
1986). We evaluated residual plots for regression assessments were based on Wr values derived for
analyses to select best-fit models; all relationships the interval associated with the growth period and
evaluated in this study produced linear models. on Wr values for a lag phase of one interval before
Parameters were derived for calculation of pre- and after calculation of growth characteristics. We
diction intervals for new data (Neter et al. 1989). determined the absolute change in Wr (for a biWe calculated relative weight (Wr\ Wege and weekly period) for evaluation against growth charAnderson 1978) at the beginning of each experi- acteristics in the aforementioned manner.
ment and at the end of each growth interval as
*-
Wr = -^x 100;
512
Striped bass
82i
78
74
70
66
62
75
90
105
120
6(T
75
90
105
120
75
90
105
120
75
90
105
120
28i
25;
22
19
16
13;
75
90
105
120
10-_
60
70i
66
62
58
54
50
60
75
90
105
_
60
120
6
1
5
75
75
90
90
105
105
120
60
75
90
105
120
120
36i
30
24
18
12
6
0'
60
75
90
105
120
Relative weight
FIGURE 1.Scatter plots depicting the relationships of percent crude fat, visceral fat, crude protein, ash, and
water to final relative weight for individual juvenile striped bass and hybrid striped bass. Crude fat, crude protein,
and ash are expressed as percent of dry body weight. Visceral fat is expressed as percent wet body weight.
513
TABLE 1.Variables for least-squares regression analysis of striped bass and hybrid striped bass relationships for
crude fat, visceral fat, crude protein, absolute protein (g), ash, water, gross energy (kj/g), relative growth (%), and
the change (A) in total length (mm) versus final Wr (P < 0.0001). Crude fat, crude protein, and ash are expressed
as dry-weight percentages. Visceral fat and water are expressed as wet-weight percentages. Absolute protein is based
on wet weight of striped bass (310 mm) and hybrid striped bass (369 mm). The sums of squared deviations for Wr
are 2,431 and 1,687 for striped bass and hybrid striped bass, respectively. Means for Wr are 93 and 86 for striped
bass and hybrid striped bass, respectively.
Linear equation
SE of Wr coefficient
fi
Striped bass
Crude fat - -36.766 + 0.661 Wr
Visceral fet = -15.064 + 0.240 Wr
Crude protein = 95.735 - 0.407 Wr
Absolute protein - -73.973 + 1.428 Wr
Ash = 44.706 - 0.313 Wr
Water = 100.070 - 0.308 Wr
Gross energy - 9.416 + 0.076 Wr
Relative growth - -306.566 + 4.337 Wr
A total length = -123.211 + 1.858 Wr
3.180
1.222
2.116
8.548
1.734
1.751
0.833
28.033
11.738
0.064
0.012
0.043
0.173
0.035
0.036
0.018
0.675
0.240
0.790
0.770
0.762
0.708
0.739
0.729
0.727
0.675
0.685
0.064
0.022
0.046
0.314
0.034
0.047
0.020
0.204
0.133
0.656
0.615
0.512
0.676
0.673
0.585
0.632
0.602
0.374
514
nonfatty fish (e.g., Gadidae) show an inverse relationship between protein and water. Based on
these criteria, striped bass and hybrid striped bass
may be classified as fatty fish because both displayed inverse linear crude-fat-water relationships and positive linear crude-protein-water relationships (Figure 2) in this study. This increase
in water content was due to extensive cellular
shrinkage with a concurrent increase in extracellular fluid (Love 1980). The highest water value
(81%) and lowest crude-fat value (5%) were observed for a striped bass; ash (27%) and crude
protein (68%) values were also highest for this
individual. Values of an individual striped bass
for the other condition extreme were 67% water,
33% crude fat, 53% crude protein, and 13% ash.
Striped bass
36
30
a? 24
5 1B1
3 12
6j
0
58 62 66 70 74 78 82
30
24
18
12
6
0^
58
66 70
74
78 82
62
Q- 58
58
54
o so62
66 70
74
78 82
36
50
58 62 66 70 74 78 82
36
30
*T 30
12
6
0
58
62
66]
62
58
.1^*^
70
9 70
? 66
I
515
62
66 70
74
78 82
24
18
12
6
0
58
Water (*)
62
66 70
74
78
82
FIGURE 2.Scatter plots reflecting the relationships of crude fat, crude protein, and ash to water for individual
juvenile striped bass and hybrid striped bass. All components (except water) are expressed as percent dry weight.
ment. This may have been because they were held estimating body composition and gross energy in
in a pond before the study. We observed general juvenile striped bass and hybrid striped bass. Reincreases in crude fat, visceral fat, and Wr with serve energy (visceral fat) predicted from Wr proincreasing ration for both striped bass and hybrid vides a measure of overwintering fitness and perstriped bass; concurrent decreases were noted in haps of suitability for stocking. If Wr can acceptably
water content, crude protein, and ash.
predict stored energy in mature fish, it can be exWe conclude that the Wr index appears to pro- tended to traditional fall population assessments
vide a viable alternative to proximate analysis for to predict overwintering fitness of the general POP-
TABLE 2.Variables for least-squares regression analysis of striped bass and hybrid striped bass relationships for
water versus crude fat, crude protein, and ash (f < 0.0001). All independent variables are expressed on a dryweight percentage basis. The sums of squared deviations for water are 317.19 and 244.79 for striped bass and
hybrid striped bass, respectively. Means for water are 71.53 and 6S.79 for striped bass and hybrid striped bass,
respectively.
Linear equation
Crude fat * 165.206 - 1.968 water
Crude protein = -23.564 -1- 1.141 water
Ash = -48.775 + 0.902 water
Crude fat = 108.106 - 1.215 water
Crude protein = 1 1.440 -f 0.642 water
Ash - -30.186 + 0.694 water
r*
Striped bass
2.054
2.026
1.524
0.115
0.114
0.086
0.912
0.782
0.798
0.168
0.120
0.084
0.659
0.514
0.719
516
TABLE 3.Means for proximate components, visceral fat, and final Wr derived from striped bass and hybrid
striped bass feeding treatments. Mean comparisons are based on Tukey's HP-procedure; means followed by the
same letter within effect rows are not significantly different (P > 0.05).
Ration level (% of body weight per day)
Effect
0.25
Water
Crude fat
Crude protein
Ash
Visceral fat
Final Wr
77 y
12 x
65 x
22 x
0.2 w
76 x
Water
Crude fat
Crude protein
Ash
Visceral fat
Final Wr
72 y
17y
59 x
21 y
1.4 y
76 y
0.50
0.75
Striped bass
72 z
71 z
23 y
26 y
60 y
58 yz
14yz
16y
1.5 x
2.5 y
92 y
92 y
Hybrid striped bass
70 yz
69 yz
23 z
25 z
57 yx
55 yz
17z
18yz
2.4 yz
3.6 y
83 yz
86 yz
56 yz
15yz
2.0 yx
95 yz
68 z
32 z
54 z
13z
3.8 z
103 z
70 z
28 yz
56 yz
14yz
2.7 y
98 yz
68 yz
28 z
55 yz
16z
3.7 y
89 z
67 yz
28 z
54 yz
16z
3.8 y
92 z
66 z
27 z
53 z
15z
4.5 y
89 z
71 z
26 y
517
518
Warren, C.E., and G.E.Davis. 1967. Laboratory studimpoundments. American Fisheries Society, North
ies of the feeding, bioenergetics, and growth offish.
Central Division, Special Publication 5, Bethesda,
Pages 175-214 in S. B. Gerking, editor. The bioMaryland.
logical basis of freshwater fish production. Black- Wicker, A. M., and W. E. Johnson. 1987. Relationwell Press, Oxford.
ships among fat content, condition factor, and first
year survival of Florida largemouth bass. TransacWeatherley, A. H., and H. S. Gill. 1983. Protein, lipid,
tions of the American Fisheries Society 116:264water and caloric contents of immature rainbow
271.
trout, Salmo gairdneri Richardson, growing at difZeigler, T. R., L. C. Woods, and J. Gabaudan. 1984.
ferent rates. Journal of Fish Biology 23:653-673.
Striped bass feeds and feeding. Pages 151-176 in J.
Weatherley, A. H., and H. S. Gill. 1987. The biology
P. McCraren, editor. The aquaculture of striped bass:
of fish growth. Academic Press, Orlando, Florida.
a proceedings. University of Maryland, Sea Grant
Wege,G.J.,andR.O.Anderson. 1978. Relative weight
Publication UM-SG-MAP-84-01, College Park.
(Wr) a new index of condition for largemouth bass.
Pages 79-91 in G. D. Novinger and J. G. Dillard,
Received May 8, 1990
editors. New approaches to the management of small
Accepted December 3, 1990