Energy in the Biosphere

Heterotrophs
(Most of the material in this lecture are from Vaclav Smil’s books on energy)
Heterotrophs
● Heterotrophs have to get their energy by eating the food produced
by autotrophs such as plants.
● There are two ways living beings use to extract energy from food.
The first one is anaerobic fermentation and the second one is
aerobic oxidation. In the first process no oxygen is involved while
the second one is driven by oxygen.
● All species belonging to the animal kingdom are aerobic
Heterotrophs.
● Fungi are also heterotrophs (some aerobic & the others
anaerobic & some can be both depending on circumstances).
● Bacteria could be either autotrophs or heterotrophs.
Heterotrophic bacteria could be either aerobic, anaerobic or both. 2
Examples of heterotrophs & autotrophs

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Heterotrophic Metabolism
● First steps of heterotrophic metabolism:
○ Break down complex carbohydrates into their constituent simple sugars
(glucose, fructose, galactose etc.)
○ To convert lipids (fats) into glycerol and fatty acids.
○ To break down proteins into their constituent amino acids.
● The energy released during the oxidation of nutrients gets stored in
ATP molecules. This energy is available for building new biomass for
growth or repair and or for doing internal as well as external work by
the conversion of ATP to ADP.
● The maximum overall efficiency of anaerobic metabolism is 30%
while it reaches 60% for aerobic metabolism for converting
monosaccharides (simple sugars) or fatty acids to energy. 4
Basal Metabolic Rate (BMR)
● BMR is the minimum power needed to energize the vital internal organs of a
heterotroph.
● It is measured at complete rest several hours after any food intake (because
even digestion increases BMR) and in a temperature controlled
environment. O2 consumption or CO2 generation is what is usually
measured to determine BMR.
● Some formulae for the BMR of humans: The Mifflin St Jeor Equation (from
Wikipedia): P stands for BMR, m is mass, h is height, a is age & s is +5 for
males and −161 for females.

● Another one which uses the lean body mass (LBM in kg & P in
kcal/day: The Katch-McArdle Formula (P in kcal/day):
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 BMR of animals

● A rough formula for mammals is BMR = 3.52m0.75 where m is

the body mass in kg and the BMR will be in watts. The
exponent 0.75 (but not the prefactor!) is also applicable for
some birds, some ectotherms and many microorganisms.
● Ectotherms do not control their core body temperature using
internally generated heat. Popularly known as
“cold-blooded”. In contrast to those animals which control
their temperature using internally generated heat goes by the
name endotherms.
● For carnivorous mammals the exponent is close to 0.9. 6
BMR of animals - 2
● The exponent in the BMR equation is much lower (just below
0.5) for desert rodents and this minimizes their energy needs and
water consumption in a hostile environment.
● While the BMR of many endotherms and ectotherms may have
the same exponent, endotherms have to pay a high cost for their
temperature control (37–38 °C for mammals, 38–42 °C for birds)
by having a BMR as much as 20–40 times higher than similar
sized ectotherms.
● High BMR indicates a high food requirement and the endotherms
have to feed themselves at frequent intervals to
survive.
● We eat thrice a day. A crocodile/snake feeds once a week! 7
Kleiber’s line (from V Smil)

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Kleiber’s line for endotherms & ectotherms

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Specific BMR
● Specific BMR (SBMR) = BMR/(body mass) is also of some interest.
This should go as w-0.25 (w is body mass).
● The smallest shrew (a rodent) has an SBMR 100 mW/g, a nearly five
orders of magnitude heavier cow requires only 1 mW/g. This limits
the mass of the smallest endothermic organisms, because
otherwise, animals lighter than shrews would have to feed
constantly in order to compensate for high rates of heat loss
because of the large surface area to mass ratio.
● The low SBMR of large mammals makes it possible for them to
survive for several days without food. Some of the large mammals
(such as bears) can also lower their BMR and hibernate for months.
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Specific BMR

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BMR of Birds
Empty circle – migrants
Filled circle - non-migrants

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Metabolic Scope
● The overall energy expenditure of most of the heterotrophs
is only a small multiple of their BMR. For sedentary humans it is 1.2 while for
the most active ones it does not exceed 1.9.
● Vigorous fliers and roaming mammals need more energy for their activities.
About 2.5 times BMR for rodents and more than four times BMR for birds that
hunt while flying.
● Metabolic Scope (MS) is the ratio of short-term (10’s of min) maximum energy
expenditure to BMR. Running, swimming and flying speeds and endurance
are limited by MS.
● For mammals MS is about ten, for some fish and most birds about fifteen and
less than five for reptiles and amphibians.
● Songbirds (4.5g-100g) can develop a maximum power of 150 mW/g, small
rodents: 50 mW/g, iguana (a large lizard): < 5 mW/g. 13
Energy Costs of Locomotion
● Canids (wolves, dogs, jackals) have the highest MS: > 30.
They are the best sustained runners of the animal world: Wolves can
pursue their prey non-stop for twenty minutes at 60 km/hr!
● Comparisons of energy costs for the three forms of locomotion - running,
swimming, flying - show that swimming, because of
water’s buoyancy, is the easiest way to move: typically it requires less
than one-tenth the energy of running. (Not for us! For natural swimmers.)
● Running needs more energy than flying for body mass up to 1 kg
● For heavier fliers the power needed for flight rises faster than the
power that can be delivered by the muscles. There are very few flying
birds heavier than ten kilograms. A peacock weighing up to a maximum of
6kg flies only very short distances. 14
Bird migration (energized by stored fat)

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 Human Energetics
● Energetics is the study of energy and its transformations in
physical, chemical or biological systems.
● We ingest three kinds of macronutrients: carbohydrates
(sugars and starches), lipids (fats) and proteins.
● We also need a large number micronutrients: the various
minerals (Ca, K, Mg, Fe, and Cu, Se and Zn), and vitamins (A, B complex, C, D, E
and K) and many more undiscovered ones!

The picture is from

https://www.difference.wiki
Note that the idea of
macronutrients shown here is
that of the western world.
Indians would include much
more grains and legumes.
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Carbohydrates
● Carbohydrates have been the staple for pre-industrial societies.
● Carbohydrates come from the following sources:
○ Cereal grains (rice, wheat, barley, rye, corn, sorghum)
○ Legumes (beans, peas, lentils, soybeans, and chickpeas)
○ Tubers (potatoes, yams, and cassava)
○ Fruits and some vegetables
● The digestible energy in these carbohydrates largely comes from
complex starches, (or polysaccharides) made up of thousands of
monosaccaride molecules.
● Fruits supply mostly simple sugars.
● All these compounds, whether complex or simple, have energy
density 17 kJ/g. 17
Lipids
● Lipids (fats) are the most energy rich nutrient with 39 kJ/g.
● They are absolutely essential for many functions in the body.
Lipids can be obtained from plant or animal sources. Rapeseed,
olives, soybeans, peanuts, oil palm, coconuts & mustard are
major sources of plant oils; butter (ghee for Indians), lard (pig
fat), and tallow (beef or mutton fat) are the three main animal
fats.
● Meat, fish and dairy products also contain a good amount of fat.
● Lipid consumption was very limited in most pre-industrial
societies but has become excessive in many affluent countries.
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Proteins
● Proteins are used as a source of energy only in emergencies when the other
two macronutrients are in short supply.
● They are the indispensable structural materials used to make new body
tissues & repair old ones.
● We need a balanced supply of essential amino acids from the proteins we
ingest as they are used in the production of enzymes, hormones, antibodies,
cells, organs, and muscles.
● All animal foods and mushrooms supply all the essential amino acids we
need for our sustenance & growth.
● The plant proteins are deficient in at least one amino acid: cereals are
deficient in lysine, and legumes in methionine. Vegetarians & vegans have to
combine various food items to avoid stunted growth and deficiency diseases.
● Energy available from proteins: 17 MJ/kg
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 Item Energy MJ/kg

Energy Content of Some Butter 30

Foodstuffs Ethanol* 29

Note: Adult human energy Sucrose* 17

requirement is about 8 – 10
MJ/day Cereals 15

* Ethanol & Sucrose (in pure Lean Meats 5 - 10

form) are to be consumed in
Fish 3-9
strict moderation.
They are actually slow poisons Potatoes 3-5
which over a period of time can
Fruits 1.5 - 4
damage various organs such as
liver, pancreas, kidneys etc. Vegetables 0.6 – 1.8
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Conversion efficiency of macronutrients
● In healthy people consuming healthy food the conversion efficiency to
energy for the macronutrients are
○ Carbohydrates: 99%
○ Lipids: 95%
○ Proteins: 80%.
● In the numbers given on the previous slide the net available energy was
shown with efficiencies already factored in.
● Metabolized food is converted into new cells and organs with
maximum efficiencies varying from 30% for adults (for maintenance) &
50% for Infants (for growth).
● After puberty energy use is dominated by basal metabolism and
the need of physical activities. The liver, brain and heart
account for more than 50% of the metabolic energy in adults. 21
BMR Distribution in Organs

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Age Dependence of BMR
● The graph shows the variation
for males.
● The numbers match in
childhood but begins to
diverge in adolescence.
● Between individuals of the same
size there is typically 10-20%
variation, sometimes up to 30%.
● There can also be variations in
the BMR of different races/
ethnicities.
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Energy use variations
● Females have lower BMR than males (of the same body mass) because of
less lean body mass.
● Generally BMR accounts for the major energy expenditure for most of us.
● For no activity (except eating and maintaining personal hygiene): 1.2 BMR,
Light labor: 1.6 BMR, Heavy exertion: > 2 BMR
● Thus depending on activity level daily energy requirements can vary from
8 – 17 MJ (2000-4000 kcal).
● The short-term limits are far higher: in trained individuals anaerobic
outputs reach up to 5 kW for a few seconds.
● Longer exertions (10–15 s) can peak above 3 kW for sprinters and
swimmers. Note that in this case the energy generation is mostly an
anaerobic process.
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Human Metabolic Scope
● Healthy adults can work (aerobically) at a metabolic scope well above
ten. For untrained adults the maximum is from ten to twenty, but for
élite athletes it can reach up to twenty five .
● This would be around 1.75 kW of power.
● Our superior temperature control through radiation, convection &
sweating is what makes metabolic scopes as high as 25 possible for us.
● Among mammals we are the leaders in sweating: up to (per minute)
500g/m2 beating our nearest rivals camels (250g/m2) and horses
(100g/m2) by huge margins. This implies we can get rid of body
heat at the rate of 600W (in extreme cases 1.3kW!).

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Human Energetics Measurements

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