Fish Bioenergetics:

Concepts and Procedures in Quantifying Consumption and Growth

Jay Hodgson
Fall 1998

Bioenergetics modeling is a powerful tool that quantifies the life history

and ecology of an individual to the population and to the ecosystem. Fish
bioenergetics models have many capacities, many of which lay outside the scope
of undergraduate study. Examples include measuring interactions between
trophic levels, calculating nutrient recycling and contaminant accumulation, and
identifying potential bottlenecks in population ecology. This exercise focuses on
quantifying the amount of food a fish consumes to the resulting growth over time.
These two variables compose the core parameters in fish bioenergetics
modeling. You will be using the Fish Bioenergetics 3.0 model for personal
computers that was developed by the Sea Grant Institute of the University of
Wisconsin. This program serves as the international standard for fish modeling.
Granted, many of you are probably not familiar with the concept of bionergetics
modeling, and the theories and mechanisms may seem overwhelming.
However, you need not worry. This exercise will outline the basic concepts and
gradually build your knowledge of modeling to the point where you can apply the
model to answer specific ecological questions.

Objectives of this exercise:

 Understand the mathematical components of the bioenergetics model
 Learn to use the bioenergetics model on a personal computer
 Understand how to apply the model to answer ecological questions

Introduction
The bioenergetics modeling strategy originated nearly twenty years ago,
and has proven useful in answering specific ecological questions. The fish
model was platformed for personal computers ten years ago, with the newest
edition being released in 1996. Although the model is user-friendly, and can
calculate answers with a few clicks of a mouse, learning to enter variables and
calculating outputs is not the main objective of this exercise. Academics today
(including the natural sciences) have become dependent on the ease of
computer-aided applications, where the core processes are often overlooked and
forgotten, which can lead to gross misuses of the intended application. Not too
many years ago before personal computers were common, ecologists had to
calculate their results the “long way,” which was based upon the actual core
processes. Computer results are still the same results as the “long way,” but it is
still important for ecologists of today to understand the methods of calculation.
 Once these mechanisms are truly understood, the results can be properly
applied to answer specific questions. To understand the core processes of
bioenergetics modeling, one must first understand what a bioenergetics model is.

What is a bioenergetics model? (General definition)

A bioenergetics model is a series of mathematical models that relates the
amount of food consumed by a fish to the resulting growth over a specific amount
of time. More specifically, a bioenergetics model is an energy balancing function
in which total energy (joules) consumed by a fish is balanced by total
metabolism, waste losses, and growth. Any one of these variables can be
predicted from the remaining series. Once again, to better understand what a
bioenergetics model is, one must first understand what a mathematical model is.

What is a mathematical model?

In the simplest terms, a mathematical model is an equation that relates
two or more variables. Although you may not have known it, most of you have
been using mathematical models for many years. The area of a rectangle is
mathematical model:

(width)(length) = area

Building upon this model, you can calculate the volume of a box:

(width)(length)(height) = volume

Unfortunately, most mathematical models (hence the word model, and not
equation) are not this simplistic in format. Most models involve substituting a
series of variables into a series of equations.

Example 1
The volume of a gas with constant pressure is directly proportional to the
temperature. If the gas occupies 100 m3 at 175, find a mathematical model
expressing the volume as a function of temperature, and calculate the volume of
the gas at 140.

Solution:
Let f(x) cubic meters be the volume of the gas at temperature x degrees.
Therefore,

f(x) = kx

where k is constant and taken from the definition of a direct proportion. Because
the volume of the gas is 100 m3 at 175, replace x by 175 and f(x) by 100 and
obtain
 100 = k(175)
k = 4/7

Substituting this back into the value of k,

f(x) = (4/7)x

You have now successfully created a mathematical model for the specific
situation. To find the volume at 140,

f(140) = (4/7)(140)
= 80 m3

Bioenergetics modeling uses constants to balance the equations and solve for an
unknown.

Checkpoint
1. Mathematical models are important in the natural sciences, especially
chemistry and physics. Other than bioenergetics modeling, give an example
where mathematical models may be used in the biological sciences.

The structure of the bioenergetics model

As described earlier, the bioenergetics model is an energy balancing
function, accounting for all energy gained from consumption and lost through
total metabolism, waste material, and growth. The model is firmly based in the
first law of thermodynamics (because energy inputs must equal energy outputs):

Einternal = Einput – Ework

The derivation of the bioenergetics model is mathematically rigorous. Therefore,

we will work our way through the design of the model.

General form of the model:

consumption = metabolism + wastes + growth

(Einput = Einternal + Ework)

The fish must consume a specific amount of joules to meet all of the
requirements of actively maintaining a functioning and growing body. The food
consumed by the fish (Einput) is balanced by the energy expenses of metabolism
and waste materials (Ework), and the remaining energy is spent for growth
(Einternal).
More specifically:

consumption = (respiration + active metabolism + specific dynamic action)

+ (egestion + excretion) + (somatic growth + gonad production)

where:

respiration is the energy expended for routine metabolism

active metabolism is the energy expended while actively moving and foraging
specific dynamic action is the energy expended to digest and assimilate prey
egestion is fecal waste
excretion is nitrogenous waste
somatic growth is non-sexual cell growth
gonad production is the production and growth of sexual reproducing cells

In notation form:
C = (R + A + S) + (F + U) + (B + G)

In addition, the model can be worked in reverse to calculate growth from

consumption, since all of the variables are interrelated. After a few
manipulations:

growth = consumption – metabolism – wastes

(B + G) = C - (R + A + S) - (F + U)

However, this exercise only focuses on the consumption calculations.

Consumption is often too difficult to actually collect field data for, while growth
measurements are quite easy. The model is useful because one can easily
estimate consumption to answer ecological questions without conducting tedious,
and often difficult, procedures involving daily stomach sampling, provided that
growth is known. The variables of respiration, active metabolism, specific
dynamic action, egestion, and excretion are separate mathematical functions that
lay outside the objectives of this exercise. In other words, they are already
programmed into the computer model and you do not have to worry about them.
The main objective of this exercise is not to calculate every variable in
bioenergetics modeling, but to see to the mathematical relationship between
consumption and growth.
Although the above form of the consumption model is mathematically
accurate, the present form of the equation poses a problem. The above form of
the model predicts the maximum consumption of the fish under ideal conditions,
but very rarely in natural settings do fish forage at their maximum rate. There are
a variety of reasons for this. One example is water temperature. Remember,
fish are ectothermic organisms, and their internal temperature depends on the
external environment. Fish metabolisms are temperature dependent, and the
model must be adjusted according to this function. Preserving the integrity of the
above model, new variables can be factored in to account for foraging rates
 below maximum. This is very similar to the mathematical model example
described earlier. First, to better organize this process, a slight change in
notation is required:

Cmax = (R + A + S) + (F + U) + (B + G)

Cmax is the maximum consumption of the fish based on known growth,

metabolism, and waste parameters. This is independent of temperature. The
below equation will calculate the actual consumption, since this is more important
in answering ecological questions:

Cactual = (Cmax)(p)f(T)

where p is the proportion of maximum consumption and f(T) is the temperature

dependence function. The temperature dependence function is another series of
equations that will be overlooked during this exercise. P (more commonly called
p-value to avoid confusion in notation) can range from 0 to 1, where 0 represents
no feeding, and 1 represents feeding at the maximum rate. The model
calculates p to balance the equations to relate the amount and quality of food
consumed to the resulting growth. Although not explicitly factored into the
equations, environmental conditions, such as the proportion and quality of prey,
influence the rate and the amount of food the fish consumes, and ultimately how
much the fish grows over that period. You will examine these interactions as well
in the exercises described below. P is the variable given the most attention from
ecologists using the model because consumption varies with the size of fish, and
does not necessarily demonstrate foraging responses under varying
environmental conditions. P, on the other hand, represents a foraging rate, and
can be used to answer foraging questions, regardless of fish size.

Checkpoint
2. Why are bioenergetics models useful for fish ecologists?
3. Why are ecologists mainly concerned with the consumption calculations from
the model?
4. Explain why p is important, and how it might be used to answer ecological
questions.
Using the computer model
To start the program, go to the Start menu, scroll up to Bioen 95, and
then over to the right until you reach Fish Bioenergetics. Once the program has
started, click on the box with the picture of fish. Then, from the following menu,
select open an existing simulation. Double-click on the samples folder, and
then double-click on perch.run. You will get a screen like this:

You already now the core mechanisms involved in the bioenergetics modeling
strategy. Now, you are going to take a look at the computer interpretations of
these mechanisms.
 Click on the temperature icon from the user input data box. You will get
a screen like this:

The days of the year are represented in column A, and the temperature (Celsius)
is in column B. The start (1) and end day (365) of the simulation must be
represented in each of the following menus after this one. If you wish to create a
simulation longer than one year, start at day 366 to represent day 1 of the
sequential year. Click OK when finished here.
Now, click on diet proportions. You will get a screen like this:

Notice that the start and end days are the same as in the temperature menu.
The days in between (you will notice there are less days represented in this
menu than the temperature menu) are not as important because the model
interpolates any data points between the start and end days. But you should use
all data that you possess to account for any seasonal variations. Columns B and
C are prey types and proportions. The sum of each row from both of these
columns must equal 1 when added across. Click OK when finished here.
 Now click on the prey energy icon. You should have a screen like this:

Once again notice the start and end days. This menu is the quality of the prey;
more specifically, the energy density of the prey in joules per gram. It is very
important that you have the same prey categories (and spelled identically) in this
menu as you had in the prey proportions menu. Click OK when finished here.
In case you are wondering, this exercise is skipping the predator energy menu
because it is not important to this exercise. This is used to alter the metabolism
of the fish. You will also be skipping the mortality menu because it is for
population-based modeling, also not important to this exercise.
 Notice the word update with a bright red back ground in the P estimate
and Bioen run columns in the Results box. Now you are ready to actually run
the model for the first time. Click on the S icon with the yellow background in the
tool bar at the top of the screen. You will get this menu:

Make sure input/out energy as joules is selected. Estimate p-value by

selecting fit to end weight. This will calculate Cactual from the growth fish the
experienced during this time period. De-select input predator energy,
spawning, and population mortality since you are not concerned with these
outputs of the model. Click OK when finished here. A user input data files
menu appears. This menu will be important later during this exercise because it
will allow you easy access to edit each of the menus that were just covered.
Simply click OK to proceed for now.
 Now the user input parameters menu appears. Make sure the first day
and final day are the same as the other menus you have just been working with.
The species start weight and species final weight are the growth parameters
used to calculated consumption.

This menu will be important later on when you are required to change the growth
parameters of the fish. Remember how you got to this menu. Click OK when
finished here to get to the next menu.
Click on the P icon with the red background in the tool bar. This will
update the P estimate column. Now click the R icon with the green background.
That will give you a screen like:

Select constant P-value and click OK to proceed.

 Your next screen should look like this:

This is the “results” window of the model. If you did not already notice, this is the
same window you were shown when you started the model. However, the
results are different because you did not calculate mortality or factor in the
predator energy density. The model also has an advanced spreadsheet and
graphing package to display results, but you will use this window for now. Make
a note of the p-value and consumption results. Notice the small difference
between the end weight and consumption results in the p estimate and bioen
run columns. This is from differences in rounding from calculations based on p.
The two columns are displayed to serves as a “check,” and to show the validity of
the model calculations. For your purposes, use the results from the p estimate
column.
Your next step is to alter some of the model parameters to see the
respective effects on p and consumption.
 Click on the S icon again, click OK (you already set up these parameters),
and then click edit under the temperature menu (this is the menu that gives you
“easy access” to other menus that you control). You are going to increase the
temperature throughout the year. Enter the numbers shown below:

Click OK to proceed. Repeat the P and R icon procedures as described earlier.

Notice the changes to p-value and consumption.
Now that you have seen the effects of temperature, you should take a look
at changes in diet. First, change the diet proportion to include a third prey type.
Select the diet menu the same way you just selected the temperature menu.
Enter the numbers as shown below:

Remember that the proportions must equal 1 when added across. Click OK to
proceed.
 Since you have just updated the prey proportions menu, you now have to
update the prey energy density menu. The same prey types must be
represented in both menus. Update this menu as shown below:

Click OK to proceed. Repeat the same P and R icon procedures as described

earlier. Notice the changes in p-value and consumption. Can you begin to see a
pattern forming?
Now, it is time for you to do a little work on your own. Change the prey
energy density to the following joules per gram:

invertebrates = 1000
fish = 1200
others = 1300

Run the model as described previously. Notice the changes in p-value and
consumption. A pattern should be evident to you now.

Checkpoint
5. What happens to the p-values as the quality of prey increases? Does it
increase or decrease? Why do you think this is? (Hint: assume your daily
consumption is 3000 calories. A quality diet may include three
cheeseburgers, each at 1000 calories apiece. A less optimal diet may include
ten salads, at 30 calories apiece. Would you be eating at a lower or greater
rate when consuming the higher quality diet of cheeseburgers?)
6. What happens to the consumption as the quality of diet increases? Once
again, use the cheeseburger example. If each cheeseburger is 100 grams,
and each salad is 50 grams, did you consume more or less mass if you ate
the higher quality diet?
7. Explain some of the ecological impacts based on the results from questions 5
and 6. (These examples assume that growth rate remains constant. The
 next example will demonstrate the relationship between p and
consumption when the growth rate is altered).
8. Furthermore, in your trial model run, temperature had little effect on changing
the p-values and consumption. This may be due to your changing the
temperature only slightly. Explain why you think or do not think a more
drastic change in temperature will effect the p-values and consumption. If
you are unsure, go back into the model and change the temperature values
and recalculate.

Now you are going to alter the growth rates. Enter the following values:

species start weight = 60 grams

species final weight = 100 grams

Run the model and make a note to the changes in p and consumption. Then
enter these values:

species start weight = 60 grams

species final weight = 200 grams

Checkpoint
9. What conclusions can you make between the growth rates, p, and
consumption? If the fish grows faster over a given period of time does this
increase p and consumption? What does this suggest about the effects of the
quality of the diet?

Now, you are going to create a simulation of your own. Close out the model
and restart it, selecting create a new simulation. Select cohort, then
largemouth bass as the next fish of study. Click on the S icon to get started.
This next exercise will be without an outlined procedure, but it should be easy
now that you have applied the model a couple of times. When editing each of
the following menus, select manual entry, and then edit. That will give you the
spreadsheet format menus. Use the following parameters in your analysis of a
period of 365 days:

Temperature:

Day Temp
1 20
175 30
365 20
 Prey proportions:

Day 1 Day 175 Day 365

Bugs 0.25 0.0 0.25
Fish 0.25 0.5 0.5
Leeches 0.25 0.25 0.25
Others 0.25 0.25 0.0

Prey energy density:

Bugs 2000
Fish 4000
Leeches 3500
Others 2500

Bass growth rate:

Species start weight 250 grams

Species final weight 350 grams

Checkpoint
10. What is the p-value for this simulation? What is the consumption? Would
p increase or decrease if the proportion of fish increased and the proportion
of bugs decreased in the diet? What would happen to consumption? If you
are unsure, use the model to answer these questions. This is an excellent
example of the capacity of the model to answer specific ecological questions.
11. Although you did not run the model to predict growth, explain why you think
or do not think growth will increase if the quality of diet increases.

Critical thinking
12. Throughout this exercise, the model has been described to examine
ecological questions. How might the bioenergetics model be used to answer
limnological questions? (Hint: cascading trophic interactions).