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Chesapeake Biological Laboratory, The University of Maryland System Post Office Box 38, Solomons, Maryland 20688-0038, USA

STEPHEN B. BRANDT

Great Lakes Center, Buffalo State College 1300 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 their prey has lagged behind. We believe this is because the process of directly 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 from the literature, obtained from cooperating investigators, and measured directly. Least-squares models were 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 coefficients of determination of 0.76 or greater. The model for all species combined (N = 587, r2 = 0.95) was J/g wet weight = 45.29 DW1-507; DW is the percent dry weight of the fish. At all taxonomic levels, energy density models showed a strong positive relationship between energy density and percent 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 science and ecology has increased rapidly during the last 10-15 years. Early models relied heavily on the work of Winberg (1956), Kitchell et al. (1974, 1977), and Brett and Groves (1979). Recently, bioenergetics models have been used to answer specific management and ecological questions. For example, models have been used to establish the stocking rates of predators (Stewart et al. 1983; Stewart and Ibarra 1991), to examine the potential effects of 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 pollutants in fish (Barber et al. 1991), and to examine how patchiness in thermal and prey fields might affect fish growth (Brandt et al. 1992; Brandt 1993). Despite the widespread use of bioenergetics models 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 dry weight basis, most data have been presented without 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 wet weight, energy density data must be placed in wet weight units to be applicable for modeling. There are changes in the energy density of fish both with ontogeny (Wissing 1974; Craig 1977; Minton and McLean 1982) and season (Craig 1977; Adams et al. 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 predators and prey are required for specific seasons, fish ages, and ecosystems. Bomb calorimetry is an accurate method for measuring the energy content of aquatic organisms (Cummins and Wuycheck 1971) and is preferable to estimates based on proximate analysis (Craig et al. 1978). However, bomb calorimetry is a timeconsuming process that requires 0.75-1.0 h/sample (Parr Instrument Co. 1960). Flath and Diana (1985) observed a negative relationship between the water content of alewives Alosa pseudoharengus and the caloric content (wet weight basis), which is due to the inverse relationship between percent lipid and percent body water in fishes (Craig et al. 1978; Flath and Diana 1985). We believe a relationship between the percent dry weight (100-dry weight/wet weight) and energy density

347

348

(J/g wet weight) exists for most fish species, and we hypothesized that a general model fitted to this relationship across taxonomic groupings would permit estimation of energy density of most fish. We further identified the limitations of this relationship by examining the effects of using energy density models on bioenergetics model results.

Methods

Calorimetry.Striped bass Morone saxatilis, bluefish Pomatomus saltatrix, weakfish Cynoscion regalis, and bay anchovy Anchoa mitchilli were collected from the Chesapeake Bay, Maryland, between June 1990 and February 1992. Fish were placed in bags and iced (striped bass, bluefish, and weakfish) or frozen in water (bay anchovy) to slow decomposition and prevent water loss. In the laboratory, fish were weighed wet ( 0.1 g) and oven dried to constant weight at 70C. Each fish was then weighed and the percent dry weight (DW) for each fish was calculated. For each species, 11-26 individuals were selected for calorimetric analysis with standard Panoxygen bomb techniques (Parr Instrument Co. 1960; Craig 1977). Individuals were selected to cover 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 were combusted to measure energy content. If the subsamples differed by more than 10% of the mean energy density (J/g), a third subsample was combusted. The mean of 2-3 subsamples was used for estimates of energy content for those fish. General models.In developing a general model for predicting the energy density of fish, we used data from our research, from peer-reviewed literature (since 1973), and from cooperating investigators. All data were treated as independent data points, regardless of the taxonomic level. Only energy contents for dry weight were used because some data were not reported as ash free. In all, 587 data points representing 35 species, 21 families, and 8 orders were included in the general model data set (Table 1). For the purposes of discussion, "combined model" refers to the energy density model derived from all fish. Each order model included data from fish of that specific order. Similarly, each family or species model contained only data from fish of a single family or species. Least-squares regression models were generated with energy density as the dependent variable and

percent dry weight is the independent variable. The empirical model for energy density was

ED = 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 empirically derived constants. For the combined energy density models, logio transformation of data was required to normalize the residuals. 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 among investigators probably introduced errors into the estimates. Our initial criterion for model acceptance was arbitrarily selected as a coefficient of determination (r2) greater than 0.75. Prior to model development, three randomly selected 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 mvkiss (From and Rasmussen 1984), and yellow perch Perca flavescens (Hayes and Taylor 1994). These independent data sets were not included in any other energy density regression models. Independent 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 utility of general models of energy density for bioenergetics modeling, we ran simulations with established bioenergetics models for bay anchovy (Luo and Brandt 1993), steelhead (anandromus rainbow trout; Rand et al. 1993), and yellow perch (Kitchell et al. 1977). We used the Hewett and Johnson (1992) software to implement the bioenergetics models for these species and used a standard set of data for seasonal weights (Table 2) and temperatures (Figure 1) specific to each species. Energy densities of prey for each predator were considered a constant value that did not vary by season. Prey energy densities were 2,552 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). To evaluate the relative importance of seasonal

349

TABLE I.Energy density models for estimating energy content (J/g wet weight) from percent dry weight (DW) data and 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 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39

Model taxon

Combined (linear) Combined (logic) Clupeiformes Perciformes Cypriniformes Salmoniformes Pleuronectiformes Clupeidae Sciaenidae Cyprinidae Percichthyidae Osmeridae Salmonidac Cottidae Alosa pseudoharengus Brevoortia ryrannus Catostomus commersoni Coregonis clupeaformis C hoyi Cottus cognatus Cynoscion regalu Dorosoma cepedianum Esox masquinongy Luxilus cornutus Morone chrysops M. saxatilis Oncorhynchus mykiss O. kisutch Osmerus mordax Phoxinus phoxinus Pleuronectes aspefi Pomatomus saltatrix Salvelinus namaycush Stizostedion lucioperca Theragra chalcogramma Cottus cagnatus

r*

0.95 0.95 0.85 0.88 0.95 0.97 0.91 0.85 0.92 0.86 0.91 0.% 0.% 0.80 0.93 0.89 0.97 0.96 0.91 0.87 0.97 0.99 0.93 0.94 0.99 0.93 0.99 0.97 0.% 0.82 0.91 0.98 0.97 0.97 0.88 0.87

N

587 587 82 80 38 348 24 82 13 33 19 25 242 10 53 25 5

5

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 in combined model 691 -2,873 -2,457

b

375.0 1.507 328.6 309.5 262.2 379.0 286.1 328.6 309.9 251.1 349.1 303.2 386.7 306.0 323.7 309.0 143.0 424.9 336.2 385.2 319.4 390.9 294.5 260.5 293.8 313.9 357.5 367.8 303.2 374.9 290.0 372.4 397.9 309.4 401.1 385.2

Data source11

1,2,9. 17 3,8 8, 11. 13, 14, 16 10,16 4,11-13,15,16 1.16 3,8 1, 17 11, 16 5, 13, 14 13 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 1 7,9 5 9 16 16 16 16 16 16 16 13 16 13 16

37 6 10 4 81 10 4 15 19 72 25 23 20 13 108 19 11 6 Data sets not included 0.77 Anchoa mitchilli 26 156.3 Perca fiavescens 0.82 313.1 55 Oncorhynchus mykiss 347.8 0.90 35 Species included in combined model, bat with too few data points for analysis Culaea inconstant 2 Fundulus heteroclitus 1 1 F. majalis Gobiosoma bosc 1 Lagodon rhomboides 3 Leiostomus xanthurus 2 Micropogon undulatus 1 Mugil cephalus 2 Myoxocephalus quadricornis 4 Paralichthys dentatus 2 1 Salmo trutta Symphurus plagiusa 2

(7) (8) (9) (10) Hayes and Taylor (1994) J. Jonas (personal communication) Kelso(1973) A. J. Paul (personal communication)

Sources: (1) This study (2) Buijse and 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.

(12) Rippetoe(1993)

Rottiers and Tucker (1982) Rudstamet al. (1994) Strange and Pelton (1987) Thayeretal. (1973) Wissing(l974)

350

TABLE 2.Seasonal wet weight (WW, g), percent dry weight (DW), and predator energy density (J/g wet weight) as calculated from species (BASE), combined (COMB), order (ORD), or family (FAM) energy density regression models.

Predator energy density Day WW

0.09 0.15 0.68 1.40 1.38 1.35 2.00 DW 11.0 12.0 17.0 22.4 22.0 21.4 23.0 BASE 2,410 2,565 3,347 4,192 4,120 4,038 4,289 3,566 COMB

1,678 1,912 3,234 4,904 4,774 4,577 5,104

ORD

FAM

a

Mean 1,083 1,411 3,054 4,829 4,697 4,500 5,026

was the mean of the seven values from the species model. The seasonally averaged model runs were used to show the relative importance of seasonality in energy content that could be missed when average values of energy density are used, as has often been done in bioenergetics modeling (Rice et al. 1983; Beauchamp et al. 1989; Raat 1990; Fox 1991; Wahl and Stein 1991; Hartman and Margraf 1992).

Rainbow trout

1

60 120 180 240 300 365

Comparisons of the bioenergetics estimates of growth from baseline P-values and consumption from growth (discrete P-fits for each run) were conducted for each estimate of seasonal energy density in Table 2. The estimates of consumption and growth from each energy density regression or the seasonally averaged data were then compared with the baseline estimate and expressed as percentage departures from baseline values to evaluate the relative error associated with each regression.

a

h

58 58 63 80 78 100 93

Yellow perch 4.017 4,774 4,774 4,017 4,171 4,937 5,272 4t485 4,954 5,786 6,498 5,581 6,682 5,736

4.709

Results We found significant relationships between percent dry weight and energy density for all models tested. Species models for striped bass, bluefish, weakfish, and bay anchovy showed significant relationships (r2 = 0.76-0.99) between percent dry weight and energy content (Figure 2). The combined model,

ED = -3,419.23 + 374.97 DW,

Database contained no other Engrualidae. Family model not possible because it would contain the same data as the order model.

changes in energy density and the model used to estimate energy density, we ran bioenergetics models for the three species. In the Hewett and Johnson (1992) software, a P-value is calculated as the proportion of maximum ration (a value between 0 and 1) consumed over the run interval, given body size and temperature constraints. A Pfit is the routine in the Hewett and Johnson (1992) model that estimates the P-value over a run interval, given growth, body size, and temperatures experienced by the fish. In our bioenergetics modeling, the year was broken into six periods and a separate P-fit was made for each period. A standard set of percent dry weight values was used to estimate the energy density of each species for species (baseline), combined, order, and family regression models and to estimate a seasonally averaged (average of baseline values) energy density (Table 2). The seasonally averaged energy density

provided a good fit (r2 = 0.95) to the data (Figure 3). However, the combined model suffered from an uneven distribution of residuals that could not be overcome by transformation of ED, DW, or both. Transformation of the data (logjo) normalized the distribution of residuals, but yielded increased variance in residuals at smaller DW values. The log 10 transformed combined model,

ED = 45.29 DW1-507,

also provided a good fit (r2 = 0.95) to the data (Figure 3). Analysis of covariance showed the combined model was significantly different from the independent data for bay anchovy, rainbow trout, and yellow perch. Comparison of the independent rainbow trout data with the order, family, and species models (Salmoniformes, Salmonidae, and rainbow trout) showed no significant differences. However, bay anchovy data had a significantly smaller slope than data included in the order model (Clupeiformes). No other engraulids were included in the database, so a family comparison was not possible

351

30

kBay Anchovy

25

^Yellow Perch

P

20 I 03 15 <D Q. 10 0)

60

120

180

240

300

360

Model Day

FIGURE I.Seasonal water temperatures used in bioenergetics simulations for bay anchovy (dashed line), steelhead (solid line), and yellow perch (chained line).

for bay anchovy. The slopes of the yellow perch and the combined, order, and family models were not significantly different, but intercepts (adjusted means) were significantly different. Intercepts and slopes of most models were sim-

ilar, generally with negative intercepts and slopes averaging 375 (Table 1). Across all models, slopes ranged from 143 (white sucker Catostomus commersoni, model 17) to 424.9 (lake whitefish Coregonis clupeaformis, model 18). Intercepts ranged

10000 8000

6000 4000 'S) ^

>>

bay anchovy

ED = 691 + 156.3 DW

striped bass

ED = -1460 +313.9 DW

2000 10000

bluefish

ED = -3792 + 372.4 DW

weakfish

ED =-1997+ 319.4 DW

15

20

25

30

35

40

15

20

25

30

35

40

FIGURE 2.Relationships between percent dry weight (DW) and energy density (ED = J/g wet weight) for bay anchovy, bluefish, striped bass, and weakfish. Regressions were fitted from species models.

352

2000

10

15

20

25

30

35

40

45

50

FIGURE 3.The linear combined model (solid line) of tish energy density (J/g wet weight) given by the equation: J/g wet weight = -3,419 + 375.0 DW; N = 587 data points and r2 = 0.95. The log|0 combined model (dashed line) of fish energy density (J/g wet weight) is given by the equation: J/g wet weight = 45.29 DW1-507; N = 587 data points and r2 = 0.95.

from 4,870 (lake whitefish) to 691 (bay anchovy, model 37). Overall, slopes ranged from -62% to 13% of the mean, and intercepts ranged from -42% to 120% of the mean.

Bioenergetics Simulations Comparisons of bioenergetics model results with baseline values showed that use of the combined model to estimate energy density provided estimates close to those from the species (baseline) values for bay anchovy, steelhead, and yellow perch (Table 1, models 37, 39, and 38, respectively). Bioenergetics estimates of growth that used the combined regression models to estimate energy were from -8.3% (bay anchovy) to 0.6% (yellow perch) of baseline values. Estimates of consumption from the combined energy density regression were from 1.6 to 4.1% of baseline estimates (Table 3). Bioenergetics model estimates of consumption and growth that used family or order models to estimate energy density had similar precision as those that used the combined model. For bay anchovy, the order model was within -1.4% for growth and 4.6% for consumption, which was similar to the combined model results of -8.3% for

growth and 4.1% for consumption. Results for steelhead and yellow perch were very similar (less than 2% difference) between order, family, and combined energy density regressions (Table 3). These data suggest that the combined energy density regression can substitute for a lower-taxon regression for bioenergetics model applications. The bioenergetics simulations demonstrated the importance of seasonal changes in energy density in bioenergetics model estimates. In every bioenergetics simulation comparison (except for consumption estimates for bay anchovy) the combined energy density regression gave results closer to baseline estimates than the seasonally averaged energy density. With seasonally averaged energy densities, simulated growth of bay anchovy was 13.7% higher than baseline, and estimates of consumption by the three species were as much as -13.3% lower than baseline (Table 3). These results show the potential importance of seasonal changes in energy density upon bioenergetics model results and suggest that seasonal averages of energy density should be used with caution. Discussion The general and species models are robust es-

353

(family, order, or combined) met the acceptance criterion (P < 0.002) with r2 values of 0.76-0.99. The r2 values of the general models compare favorably with those reported for linear functions of the same relation for alewife (Flath and Diana 1985) and bloater Coregonus hoyi (Rudstam et al. 1994). There is a general relationship between changes in energy density and changes in water content across taxa. We found that slopes rarely differed significantly among model comparisons. However, the lack of significance could be due in part to a high degree of variability in the data. The ANCOVA indicated that real differences across species or taxonomic groupings appear to be in the intercepts (or adjusted means) for the models. Explanations for differences in intercepts could be variation in amounts of bones (Cameron 1985; Rottiers 1993), in basic lipid levels across fish taxa (Cummins and Wuycheck 1971), or in procedural differences across the 17 sources of data sets. Taxonomic differences in percentages of calcified structures or basic differences in lipid levels would account for the significant differences in intercepts between general regressions (combined, order, and family) and species regressions. Because bone has a relatively low energy content, species with more bony parts would have a lower energy density per gram dry weight. Fish species with higher basic lipid levels would have higher intercepts because of the higher energy content of lipids relative to other dry matter, such as bone or muscle (Craig 1977). The significant differences in intercepts and slopes among species models may indicate that there are species-specific differences in the energy density relationship. However, despite these apparent species-specific differences, our evaluation of the energy density models for use in bioenergetics models showed that the order, family, and combined models were adequate for describing seasonal changes in energy content in fish. Bioenergetics model runs that used combined regressions for energy density were within -8.3 to 0.6% of baseline growth and consumption estimates for the three species modeled. These generalized energy models were much closer to baseline values than were the estimates of growth and consumption estimated with a seasonally averaged (constant) energy densitya method commonly used in bioenergetics modeling (Kitchell and Breck 1980; Rice et al. 1983; Bevelhimer et al. 1985; Boisclair and Leggett 1989; Hartman and Margraf 1992). The bioenergetics modeling simulations

TABLE 3.Comparisons of consumption and final weights from bioenergetics models for bay anchovy, steelhead, and yellow perch with energy density estimated from species models as the baseline estimates. Energy density regressions for combined, average, order, and family models were used to estimate consumption and final weights in separate bioenergetics model runs with the Hewett and Johnson (1992) model. Values are percentage deviations from baseline model predictions of consumption from weight and final weights from baseline P-values.

Energy density regression Combined Average Order Combined Average Order Family Combined Average Order

a

-8.3 13.7 -1.4

Consumption

-1.0 1.9 6.3

Specific f-tit for consumption 4.1 -3.8 4.6 1.6 -13.3 1.4 1.7 3.9 -12.7 2.7

Bay anchovy11

Steelhead

-1.4 8.3 -0.6 -0.8 -0.2 2.0 -0.2 -0.2

A family model for bay anchovy (Engraulidae) was not possible because no other energy density data on engraulids was used in the models. h A family model for yellow perch (Percidae) was not possible because it contained the same data as the order model (Perciformcs).

presented in this paper point out the need for seasonal energy density information and also support the validity of using general models to predict energy density from percent dry weight information. Used carefully, the energy density models should provide accurate estimates of energy density variables in field applications of bioenergetics models. Fish are routinely saved for diet and growth information in bioenergetics applications and their percent dry weights may be readily available (at little additional cost) as inputs to the regression models. Previous error and sensitivity analyses have shown bioenergetics models to be relatively insensitive to energy density variables (Rice et al. 1983; Stewart et al. 1983; Bartell et al. 1986). Therefore, the general energy density models that incorporate seasonal and ontogenetic changes in energy may be used with minimal risk. We recommend that, wherever possible, direct measures of energy density be made for use in bioenergetics models. However, for many applications and particularly for energy estimates of incidental prey species, the general models presented here should prove adequate. Recognizing that species-specific

354

differences in energy density relationships exist, we recommend that researchers use the model of the lowest practical taxon (species before family, order, or combined) whenever possible. However, in the absence of more specific regressions, the combined model should provide reasonable seasonal and ontogenetic changes in energy density as input parameters for bioenergetics models.

Acknowledgments Preparation of this paper would not have been possible without the cooperation of J. Jonas, B. Lantry, A. J. Paul, P. Rand, and T. Rippetoe, who provided us with data on fish energy density. P. Blanchette provided calorimetry equipment and instruction. Calorimetric bombing of bay anchovy was conducted by R. Miller and calorimetric bombing of weakfish and bluefish was provided by the chemistry department of the Potomac Electric Power Company's Chalk Point generating station. We thank J. Luo, V. Kennedy, P. Rand, D. Rottiers, and an anonymous reviewer for their comments on this manuscript. Support for this study was provided by the Electric Power Research Institute's Fellowship Program in Population Biology (EPRI-09-07), the Sport Fishing Research Institute (SFRP-90-21), the Maryland Sea Grant Program (NA90AA-D-SG063 R/F-65), the National Science Foundation's Biological Oceanography Program (OCE-9116071 and OCE9415740) and the Environmental Protection Agency's Chesapeake Bay Program.

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Received August 30, 1993 Accepted December 13, 1994

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