How to simulate this endotherm

Consider a 250 g generalised mammal where operative environmental temperatures vary

from 10 to 40 oC. As an endotherm, assume that it generates heat endogenously whenever
its body temperature falls below 37.5 oC. Assume that its thermal conductance is typical
for mammals of this size and that it can vary this thermal conductance six fold as many
mammals can.

1. Open “Calculators.xls” and click on “Allometric M calculator”. Enter mass (250)

into one of the appropriate calculators presented. Using SMR according to White
et al. (2006), SMR is 1.168 W when Tb is 38 oC, but you could use other
calculators.

2. Click on “Ko calculator” and enter 250 to get a minimal thermal conductance of
 0.1355 W/oC for mammals according to Withers (1992).

3. Click on “Respiratory variables” and enter 250 to get a minute volume of 98.71
ml/min for mammals according to Withers (1992).

4. Open “Optemp.xls” and enter data into St St Start. While these data will define
the reference equilibrium (see User Guide and paper for explanations) and should
therefore eventually be in the Eq End column, we need to use “Tlc to balance”
and this requires data to be in St St Start. Assume metabolic rate is basal (1.17
W) and thermal conductance is minimal (0.1355 W/oC), minute volume is as
calculated (99 ml/min) when metabolic rate is basal and increases in direct
proportion with increases in metabolic rate. So the proportionality factor is 1.
Let’s have a cooling effect of the nasal turbinates, present in mammals, of say 5
o
C. Assume an arbitrary humidity of 30 % when air temperature is 20 oC.
Obviously, use respiratory minute volume (as opposed to total evaporative water
loss) and link minute volume to metabolic rate. We haven’t decided on an
operative temperature yet, but we don’t have to. The program can do that for us.
Any endotherm thermoregulating according to the Scholander-Irving model
should decrease its thermal conductance with decreasing environmental
temperature until this reaches its limit (maximum insulation). Then, the
endotherm will increase its metabolic rate with further decreasing temperatures to
maintain its body temperature (unless it goes torpid). The environmental
temperature at which body temperature can be maintained with minimum thermal
conductance and basal metabolic rate is known as the lower critical temperature
and “Optemp.xls” can calculate this using the button “Tlc to balance”. Enter the
mean body temperature (38) and press the button to obtain Tlc of 30.26 oC.
5. Make operative temperature = the above result and press “start” to solve this
stage (i) simulation and define what will become the endpoint equilibrium and

therefore the reference equilibrium (see User Guide and paper for more
explanation).

6. Before using “swap” to move these inputs, use the Eq End column to define a
starting point because, after all, we’re starting our mammal in an environment at
10 oC. To maintain its body temperature in environments colder than Tlc it needs
to increase its metabolic rate while keeping its thermal conductance at a
minimum (assuming it does follow the Scholander-Irving model). By how much
is the question. While there is a calculator, “M to balance” that could help, to
work it needs the data in the St St Start column and it needs minute volume.
Unfortunately, minute volume varies with metabolic rate, so we don’t know what
this will be either. Not to worry, minute volume is calculated automatically for all
simulations subsequent to the first when the “Link minute volume to metabolic
rate” option is ticked. So the easiest option is to try a range of metabolic rates (in
Eq End) together with minimum thermal conductance, the same humidity
parameters and operative temperature of 10 oC and find a solution that balances
for a body temperature of 38 oC.
7. A metabolic rate of 3 to 4 times basal would be a good bet, so enter basal (1.17)
for the nominal metabolic rate and the factors 3 and 4 to define the range. Enter
other inputs as before except for operative temperature that should be 10 oC.
Don’t bother with minute volume because whatever you enter will be overwritten
by the value calculated by “Optemp.xls”. In this case, it will be 99, the same as in
St St Start, because the nominal metabolic rate is the same. The actual value used
will be adjusted for each of the nine solutions ranging from 3 × 99 to 4 × 99
ml/min.

8. A solution for body temperature of 38 oC lies somewhere between 3.50 × and

3.63 × nominal metabolic rate, so adjust the factors accordingly and re-run (press
“end”) to get a solution of 37.99 oC (which is close enough) for line 8. This
shows that the actual metabolic rate to maintain this body temperature is 4.228
W. If you want, you can place 4.228 into the nominal rate in Eq End, make the
range from 1 to 1 (or anything else you fancy) and press “End” again to see
minute volume calculated out as 357.8 ml/min. You’ll get a very slightly
different body temperature, but this difference is immeasurable in any practical
sense. The actual evaporative heat lost (0.438 W) was already calculated.
9. Assuming you didn’t play around and still have the inputs and outputs as
depicted above, press “swap” to get all the inputs into the right places and re-run
the simulations. When you press “start”, you’ll probably get an error message.
This is transitional. For the moment, while the start and the end are exactly the

same, Excel is trying to plot the difference between them on a log scale in stage
(ii). You may notice the little stage (ii) results graph in the top right of screen has
disappeared. We don’t care at the moment, because we are about to change the
end anyway, so just ignore the message, press “OK” on it and then press “end”.

10. Once the inputs are in the right places and the final stage (i) outputs have been
generated (as depicted below), click on stage (ii).
11. Note the reproduced output from stage (i) with line 8 steady state start body
temperature of 37.99. Enter mass (250) and line (8) and press “Use steady state
start for stage (iii)”. Then go to the stage (iii) sheet.

12. Enter the lower set point (37.5), an upper set point (38.5) and Q10 (2.81 from
“Calculators.xls”). Allow the animal some aerobic scope by entering a maximum
M (12 ~ 10 × BMR). Enter thermal conductance values, minimum (0.1355) and
 maximum (= 6*e7 or .183) and effects of turbinal scrolls (std effect = 5). Enter
the range of operative temperature (maximum 1 = 40, minimum 1 = 10) and
length of sim (2880). Press “Define the dynamics for a stage (iii) simulation”.

13. Work through the tabs in the “Set Parameter Functions” form.
14. In the M tab choose “Varies according to a Q10” and “defend Tb from
dropping”.

15. In the Ko tab choose “Maximise approach of Tb to Tpref” and “Adjust Ko to

maintain midpoint of Tpref”.
16. The
he next two are really simple. In the Vi/E tab choose “Maintain” and in the Tue
tab choose “constant Tue” at all Te”.

17. In the Te tab choose “Sinusoidal Function starting at minimum”.

18. Press “Run the stage (iii) simulation” and wait for the simulation to complete.
 Various indicators show the simulation is running including one showing how far
the simulation has
progressed (in sim time
and as a percentage).
When complete, a form,
“Summary Statistics and
Analysis?” appears giving
you several options to
calculate statistics for part
or all of the output. You
may choose to ignore it by
clicking “No” or “Exit”. If
you do select “Yes” fill
out start and finish time
and select the stats you
want. (For explanation of
leeway, see the User
Guide.pdf.).

And here are the results displayed in the “Lg Graphs” worksheet: they are also

displayed in “(stage (iii)” at a smaller scale. For most of the two days of the
simulation, the endotherm maintained its body temperature between 37.3 and 38.1 oC
through a combination of variations in thermal conductance (when operative
temperatures were thermo-neutral) and increases in endogenous metabolic heat
(when operative temperatures were lower). Only at high operative temperatures did
the body temperature deviate from homeothermy. The combination of high
environmental heat load and (increased) metabolic heat was too great for the animal
to maintain Tb despite dumping heat at the maximum rate allowed by its thermal
conductance. (Increasing maximum thermal conductance from 6 × to 8 × minimum
reduces the maximum Tb during the simulation by about ½ oC). Metabolic heat
increases when Tb increases because of the effect of body temperature on metabolic
rate, generally a 2 to 3 fold increase for every 10 oC increase in temperature. Thus
while metabolic rate influences body temperature by providing some of the heat in
the system, body temperature influences metabolic rate through the Q10 effect: there
is a feedback loop between the two. This feedback loop is incorporated at the heart of
“Optemp.xls” and is one of the reasons “Optemp.xls” can be used for both
ectotherms and endotherms in a variety of circumstances. While still there, the
feedback loop is masked in endotherms by their ability to generate extra heat over
and above the Q10 effect, either to keep warm in cold places or, perhaps, to run
cooling mechanisms in hot places.

Clearly, our hypothetical animal in this environment needs cooling mechanisms to

avoid Tb > 38.5 oC in the circumstances modelled. While “Optemp.xls” does not yet
include sweating, it does include panting at least in a rudimentary way. Panting is
merely a dramatic increase in minute volume initiated, perhaps, at a metabolic cost.
Enter a value for “high Vi” and “metabolic cost” in the inputs part of the spreadsheet.
I chose about 20 × Vi when M is basal (20 × ~ 99 = 2000) because experience has
shown me this is the smallest factor sufficient to bring Tb within about ½ o of the
upper set point, and 0.1 W (a substantial increase over BMR) as a pure guess. In the
“Vi/E” tab of the “Set Parameter Functions” form, select “Increase at high Tb” and
then run.

The output shows a substantial increase in evaporative heat loss leading to more
pronounced homeothermy with Tb being reduced below Te during the hottest part of
the day. Interestingly, the lower Q10 effect of this reduced Tb on M more than
compensates for the increase in metabolic rate expected from a substantial cost of
panting.

Congratulations! You have simulated a homeothermic endotherm in a fluctuating

environment. And, with a bit of luck, you have thought of a few questions about
endothermic thermoregulation along the way! Now play with the programming to
either break it, or to generate more questions. If you do break it, or you have any
questions or suggestions, let me know. Remember though, there is a more
comprehensive user guide that contains many of the details omitted here.