Original

Energy content
Michael

E May

ify the

This

calorie

acid

calorimetry

acids
for

was
the

to

calculated

the

availability

of

the

of the

energy

metabolism,

with

when

in-

oxidation

carbohy-

free amino
of combustion

characteristic

of

were presented
before,
ofpublished
values and
program

to

do

these

1990;52:770-6.

acids,

indirect

heat
acids

computer

Amino

diets
and

representative

of amino

Am J C/in Nutr

KEY WORDS

to a variety

calculations,

amino

mine,

uric

nitrogen

acids,

tion

calorimetry

acids

ofnutrient
of loss

tion ofthe
lism. The

minus

the

been

bustion

minus

urine

(1). Simple

acids,

such
heat

and

heat

heat

urine,

the frac-

induced

by inges-

pathways
of metabonutrients
is the heat of

of combustion

of products

of me-

sugars,

these

tween

biological

latter

content

will be greatest

is least,
value.

and thus

completely

and

the energy

are

oxidized

acids

urinary

metabolites
account

content

It seems

each

of corn-

when

amino

The purpose
arithmetic
bases

of this article
for computation

food

to outline

the

that

and
the

to carcontent
to carbon
ni-

difference

the heat

proportion

acid

have

and

calculations

other

foods.

and

different,

the

compounds

among
yield

These

products

were

chosen

these

a small

energy

depends

content
on

compounds.
amount

the

The

of urinary

nitrogen-

distribution

of the

sulfur-containing

of urinary

taurine

ni-

amino

along

with

urea,

ammonia,
creatinine,
and uric acid and produce
additional
metabolic
energy by the oxidation
ofthe sulfur to sulfate. These
computations
were summarized
recently
(5), but the tedious
task of computing
the urinary
metabolite
energy content
for
all naturally
occurring
amino
acids was simplified
in the past
by assuming

that

urine

nitrogen

is in only

one

form,

such

as

urea (5).
We have written
a microcomputer
program,
METENERG
(ME May, JO Hill, vanderbilt
University,
Nashville,
TN) that
computes
the metabolizable
energy ofthe amino acids after the
user enters the distribution
ofurine
nitrogen
either as a fraction
or as weight of compound
excreted
per day. Because
the urinary nitrogenous
metabolites
contain
carbon,
hydrogen,
and
oxygen
in addition
to nitrogen,
the volume
of oxygen
consumed
in the catabolism
g ofurine
nitrogen
(O:N),

ofamino
acid
the metabolic

liter

used

of nitrogen

a different

caloric

to computerization
Am iC/in

ammonia.

abundance
in urine of normal
adults (4). Beof combustion
of these compounds
per mole

of gaseous

oxygen

sufficient
energy

in oxidation

of
Nutr

From

the Departments

of Medicine

to produce
1
equivalent
per

of an amino

and Pediatrics,

University,
Nashville,
TN.
2 Supported
in part by a pilot and feasibility

energy

is to review
the chemical
and
of the energy
value of defined
approach

computerized

acids

acid

beof com-

biological

the weight

may

and

fatty

containing

for the

of protein

intuitive

material

nearly

absorption,

compounds

of protein.

chemicals,

as glucose,

are oxidized

several

bustion

heat

and short-chain

after

energy

measured

such

Amino

and

the

of fecal

acid,

water

from

of combustion

ofcombustion.
water,

trogen;

770

secretion

the biochemical
of absorbed

determined

the

as acetic

bon dioxide
is the

lost in the

of intestinal

that are not fully oxidized.

These principles
are well
and the usual factors for caloric content
of foods have

traditionally

dioxide,

feces

foodstuff,
and
energy
content

combustion

tabolism
known,

ofa foodstuffis
determined
by multiple
efficiency
of intestinal
absorption,
the

or its metabolites

into

acid,

are

containing

Introduction

fraction

the

ofamino

The heats of combustion

and heats of formation
of amino
acids were obtained
from standard
references
(2, 3). Heats of
combustion
were calculated
for lysine, ornithine,
proline,
and
histidine
from the heats of formation
of amino
acid, carbon
dioxide,
and water (see Appendix
for step-by-step
description
of calculations).
The biologic
energy content
was obtained
by
subtraction
ofthe residual
heat ofcombustion
contained
in biologic end products
from the heat of combustion.
The major
nitrogenous
excretory
products
were assumed
to be urea, creatbecause
of their
cause the heats

branched-chain

to apply

Methods

trogen

The energy content

factors
including
the

and

ofmixtures

or protein.

of individual

humans.
Although
these computations
we now report the limit ofapplicability
calculations.

acid

substrate

and

content

oxidation

these

mod-

program
to compute
the enand respiratory
quotient
for

by correction

incomplete

nitrocan

experimental

determine

acids

calorie

that

are compared

a computer
equivalent,

The

ofdietary
factor

of amino

when

of amino

fats.

amino acid composition13

l990;52:770-6.

grant

from

Vanderbilt
the Diabetes

Research
and Training
Center,
NIH 5 P60 DK20593-lO,
and by NIH
grants DK38088
and DK26657.
Address reprint requests to ME May, AS 105 MCN Vanderbilt
University, Nashville,
TN 37232-2230.
Received
September
5. 1989.
Accepted
for publication
December
27, 1989.
Printed

in USA.

1990 American

Society

for Clinical

Nutrition

Downloaded from ajcn.nutrition.org by guest on September 9, 2014

and

weight

is used

mixtures

drates

per

is one

compositions

rates. We developed
ergy content,
oxygen
arbitrary

acids

to consider

amino

direct

in the distribution
amino

equivalent

is important

Communications--general

0 Hill

variation
the different

different

of diets of variable

and James

ABSTRACT
gen among

Research

CALORIE

CONTENT

OF

(H02),
and the ratio of gaseous
carbon
dioxide
produced
oxygen
consumed
(RQ) also depend
on the distribution
urine nitrogen
among
the possible
metabolites.
METENERG
computes

these

nitrogen

quantities

distribution

computational

for each
is specified

:
heat

ing amino

to

the

the

urine-

following

and

corn-

by the expression

nitrogen

in compound
i, and

n is the

i, H,
number

urinary
cornacids (includ-

proteins).

metabolizable

as heat

in urinary

nitrogenous

nitrogen

ofcompound

maximum

is computed

urinary

per mole i; i represents

any
nitrogen
derived
from amino

acids

2) The

of

urine

of urine

ofcombustion

of moles nitrogen
pound containing

heat

according

after

(F1,).(HJn1)

F, is the fraction

is the

acid

algorithm:

1) The heat of combustion

pounds
is computed
per mole

where

amino

to
of

energy

ofcombustion,

nitrogenous

ofeach

amino

acid,

amino

minus

acid

residual

compounds.

formulas:
c

ou

(F1,)

>

4)

The
can

chemical
be written

(c1/n1)

(F1,)-(h1/n1)

reaction

for biologic
terms

8) Sulfur

amino
for energy

account

+ X02

oxidation

of an amino

as
+ YCO2

ZH2O

The molecular
compositions
of the natural
amino
acids are
known and the apparent
molecular
composition
of the urinary
nitrogenous
compounds
depends
on the distribution
of urine
nitrogen
as detailed
in reference
3. The stoichiometric
coefficients are computed
as follows:
=

ca

(ha

c#{149}
na

hufla)/2

X(2Y+Z+Oufla)/2
where Y is the CO2 production
in mol/mol
amino acid and X
is the 02 utilization
in mol/mol
amino acid.
5) The RQ for a diet is the ratio of carbon
dioxide produced
to oxygen
consumed
in complete
oxidation
of the diet after
complete
absorption:
RQ

6) The heat equivalent

Z/X

of oxygen

short-chain

carboxylic

METENERG
for use on an
MSDOS.

(Metabolizable

7) The ratio of oxygen

tion (L 02:g N) is
O:N

energy)/(X
utilization

two

DOS

batch

a minimum

files allow

report

gas constant)

nitrogen

file

is 01

Basic 1.0 and compiled

microcomputer
under

of 128 kb RAM

selection

for

the

reports

sections

memory

and

ofeither

molecular

weights

first

report

and

file MOLdirectory
directory.
the name

then

increases

are generated.

read

combustion
of amino
fault
distribution
of

the chemical

acids
urine

formulas

and other
nitrogen

and

substrates,
from
the

heats

of

and the defile MOLE-

CULE.CON.
All numbers
are stored as double-precision
reals
(8 bytes,
14-digit
precision)
to minimize
rounding
errors.
The
main menu allows four choices or exit: enter fractional
distribution

of urinary

with

nitrogen,

O:Nu,

and

nitrogen,

calculate
calculate

enter

amino

energy

mass

acid

of urinary

metabolites

metabolizable

parameters

ofa

energy

food

and

mixture.

Program
flow is directed
by a SELECT
CASE <choice)
statement, which forces return to the main menu before to exit from
the program.
There are two functions
and 32 procedures
that
do the following
operations:
screen manipulation,
1 1 routines
to clear specified
blocks,
print messages,
format
numbers
as
strings for display, and print successive
screens; keyboard
input
routines,

7 routines

routines,
tations;
program

ganic

produc-

to

This feature
allows
multiple
reports
to be generated
in one run of the program.
When the report counter
reaches 99, all subsequent
report files
are named
RQDIO1KA.OUT
until the report files are cleared.
It is the responsibility
of the user to delete old report files by
using appropriate
DOS commands.
The program
is organized
as follows: The first screen displays
the name, copyright
notice, and an option to exit to DOS. The

output

substrates
acids,

acids,

the latter
(2). The

for entry

ofspecified

quantities;

calculation

10 routines,
each for a specific portion
ofthe compuoutput
routines,
4 routines;
and program
flow, main
plus 2 routines
comprising
groups ofthe input, calcuand

The

02) is

to urine

X.gasconstant/14.0067.n5

calculations
compounds.

acids.

was written
in Turbo
IBM-PC#{174}--compatible

It requires

22 amino

Heq

easy extension
of the
of urinary
nitrogenous

one disk drive.

Printer
output
uses ASCII
codes only and
should
be compatible
with most common
printers.
The program files are METENERG.EXE,
MOLECULE.FW.
MOLECULE.MW,
AAFORWT.BAT,
and AAMOLWT.BAT.
The

lation,

(kcal/L

to

The ionization
fraction
of organic
acids is required
to obtain
correct
RQ values
(5), and the program
includes
input of the
ionization
fraction
and the RQ correction
for five common

to 99 as additional

CCUHhUOOUN

acids require
additional
algebraic
terms
release
and oxygen
utilization
in oxidation

of the calculations,
with
any specified
distribution

initialization
CcaHhaOoaNna

771

ofsulfur
to sulfate and the small change in apparent
molecular
composition
ofunnary
nitrogenous
compounds
occasioned
by
the excretion
of taurine.
The above algorithm
is not original (5), but the algebraic
formulation
of the chemical
balance
allows the computerization

of the

>: (F1,)-(o1/n1)

in general

ACIDS

or acyl residue
formula
weights
for the parameter
ECULE.CON.
The program
searches
only the default
for the latter file and puts output
files in the same
Report
files are named
RQDI??KA.OUT.
The 7? in

where c, h, and o1 are the numbers

ofmoles
ofcarbon,
hydrogen, and oxygen
per mole of nitrogen
in compound
i, respectively. In the calculation
ofthe correction
for energy content
of
amino acids, only nitrogenous
compounds
in the urine will be
included
rather than all urinary compounds.
acid

AMINO

routines.

for diets included

5 carbohydrates,

ethanol,

and

18 substrates
program

tains
compositions
(MOLECULE.MW)

4 fats.

in MOLECULE.CON
3 polyols,
5 short-chain

Thermodynamic

were obtained

is supplied
and

with
molecular

and the other

from

two

properties

published

parameter
weights

contains

are
or-

files;
of

for

literature
amino

compositions

one

conacids

and

Downloaded from ajcn.nutrition.org by guest on September 9, 2014

3) The number
ofcarbons
(ca), hydrogens
(he), and oxygens
(on) in the urine per mole nitrogen
are derived
from analogous

FREE

772

MAY

TABLE
Heats

AND

HILL

1
ofcombustion

and biologic

oxidation

ofamino

acids
Metabolizable

Compound

Heat

ofcombustion

Set 1

Set 2

Set 3

kca//g
Alanine

2.875
3.256
3.0 15
3.646
4.207
3.097
4.85 1
6.523
6.524
6.038
4.456
5.493
6.723
5.681
3.308
4.120
6.588
5.859
5.963

I
SD
CV(%)
a

4.781
1.324

For four distributions

and set 4-data

formula
acids

3.425
3.254

ofurine

from Table

weights

ofthe

nitrogen:

3 ofreference

acyl portion

(MOLECULE.FW).

by the DOS batch

for diets composed

tein

amino

natural
diets
the nonprotein

acids,

3.549 ( 14.20)
3.508 ( 14.03)
2.4 19 (9.68)
2.344 (9.38)
4.402 ( 17.6 1 )
4. 182 ( 16.73)
3. 166 ( 12.66)
3.241 (12.96)
2.158 (8.63)
3.487 ( 13.94)
5.986 (23.94)
5.988 (23.95)
5.072 (20.29)
5.385 (2 1.54)
4.426(17.71)
6.296 (25. 18)
5.066(20.27)
2.637 ( 10.54)
3.527(14.11)
5.896(23.58)
5.470 (2 1.88)
5.358 (2 1.43)
4.253(17.01)
1.321 (5.283)

27.7

32.4

urea,

3% ammonia,

(dehydrated)

ofthe

of parameter

amino

files

is ac-

files. The program

as written
is
of modular
components
(pro-.

carbohydrate,

fats),

would
require
modification
nitrogen
was negligible

and

extension

to

that
cornputations
strictly
apply
to the portion
of the diet that is absorbed or taken parenterally;
digestibility
is not specifically
entered in the program.
Because the user is prompted
for the diet
components
by weight, known digestibility
factors (amount
absorbed/amount
eaten) can be accounted
for by entering
Weight
x Digestibility
instead
ofWeight
for each nutritive
component
and entering
Weight
X (1-Digestibility)
plus the weight
of

or assumption
in quantity.
The

dietary
minerals
as Mineral
for the program-this
rect energy
computations
per weight
of the original
compiled

program

to cover

reproduction

Results

and discussion

Table
energy

1 shows
for amino

is available

from

the

authors

allows
diet.

cor(The

for $25

(US)

and shipping.)

the
acids.

heats

of combustion

Metabolizable

3.494 ( 13.98)
3.396 ( 13.58)
2.345 (9.38)
2.307 (9.23)
4.362 ( 17.44)
4. 143 ( 16.57)
3. 133 ( 12.53)
3.173 (12.69)
2.093 (8.37)
3.392 ( 1 3.57)
5.949 (23.80)
5.950 (23.80)
5.005 (20.02)
5.353 (2 1.4 1)
4.352(17.41)
6.266 (25.06)
5.024(20.10)
2.590 ( 10.36)
3.486(13.94)
5.848(23.39)
5.443 (2 1.77)
5.3 16 (2 1.27)
4.201 (16.80)
1.339(5.321)
31.1

2% uric acid; set 2-100%

5% creatinine,

( I 8.28)
( I 5.83)

4.369
3.784
2.7 18
2.689
4.792

( 1 1.37)
( 1 1.25)
(20.05)
-

3.583 (14.99)
3.61 1 (15.11)
2.741 (11.47)
4.2 16 ( I 7.64)
6.886(28.81)
6.888 (28.82)
5.686 (23.79)
6.06
(25.34)
-

7.036 (29.44)
5.937(24.84)
3. 107 (13.00)
4.101 (17.16)
6.419(26.86)
6.039 (25.27)
6.276(26.26)
4.847 (20.28)
1.530(6.402)
31.7

set 3-l00%

ammonia;

urea;

(2) for 100% urea.

Selection

complished
appropriate
and/or

set 1-90%

( 13.70)
( 13.02)
2.252 (9.0 1)
2.26 1 (9.04)
4.3 12 ( 17.24)
4.093 ( 16.37)
3.09 1 ( 12.36)
3.089 (12.36)
2.01 1 (8.04)
3.273 ( 13.09)
5.902 (23.6 1)
5.903 (23.6 1)
4.92 1 ( 19.68)
5.3 13 (2 1 .25)
4.259(17.04)
6.229 (24.92)
4.970(19.88)
2.532 ( 10. 13)
3.434(13.74)
5.787(23.14)
5.409 (2 1.64)
5.264 (2 1.06)
4.136(16.54)
1.342(5.370)

and
energy

metabolizable
is presented

for

four distributions
of urine nitrogen.
The right-hand
column
of
Table
1 is the metabolizable
energy
reported
by Livesey
and
Elia (5) under the assumption
that urea is the sole urinary
end

product.
Minor
slightly different
our calculations
some.

We

differences
would be expected
from the use of
primary
sources,
but the large differences
in
and those previously
reported
was bother-

replicated

the

numbers

of Livesey

and

Elia

by use of

the compositions
ofthe acyl portion
for computation
of nitrogen content
and formula
weights
rather
than the molecular
composition
for each ofthe amino acids. That is, the previously
reported
parameter
values refer to amino acids in protein when
the amino acid data has been expressed
as
Amount

ofamino
in amino

acid
acid

fraction

X acyl

ofprotein

formula

weight

nitrogen
per

mole

nitrogen

Ifamino
acid composition
is expressed
in these terms, the sum
of amino acid weights will be equivalent
to the protein
weight
and we can call this mode of expression
protein
equivalent.
Actually,
the sum ofamino
acid weights obtained
by hydrolysis
of protein
will be greater than the weight of the initial protein
by the amount
ofwater
added to hydrolyze
the peptide
bonds.
There is no convention
stated for expression
of amino
acid
data in standard
tables of food composition
(6). Furthermore,
the protein-equivalent
mode ofexpression
ofamino
acid cornposition
ofproteins
is not applicable
to diets composed
of crystalline

amino

parenteral
tabolizable

acids,

as are

often

used

in experiments

and

in

nutrition.
The previously
published
values for meenergy of amino
acids do not apply to diets corn-

Downloaded from ajcn.nutrition.org by guest on September 9, 2014

Asparagine
Aspartic acid
Cysteine
Cystine
Glutamic
acid
Glutamine
Glycine
Histidine
Isoleucine
Leucine
Lysine
Methionine
Ornithine
Phenylalanine
Proline
Serine
Threonine
Tryptophan
Tyrosine
Valine

Set 4

kca//g(kJ/g)

4.34 1
S. 129
3.488

Arginine

energy5

CALORIE
TABLE

CONTENT

OF

ofoxygen

for amino

Heat equivalen

acids
ofoxygen

Set 1

acid

consum ed

in oxidation5

Set 2

Set 3

Valine

4.630(18.52)

4.668(18.67)

4.631 (18.52)

j:

4.688
0.259

4.757
0.248

4.690 ( I 8.76)
0.24(1.01)
5.4

Asparagine
Aspartic acid
Cysteine
Cystine
Glutamic
acid
Glutamine
Glycine
Histidine
Isoleucine
Leucine
Lysine
Methionine
Ornithine
Phenylalanine
Proline

Serine

SD
CV (%)

Sets described

( 18.75)
( 1.04)
5.5

in Table

equivalent

4.686

( 18.74)

4.671
4.655

(18.68)
( 18.62)

4.630(18.52)
4.798 (19.19)
4.607(18.43)
4.567 (18.27)
5.31 1 (21.24)
.300 (21.20)
4.70(18.28)
4.98(18.39)
4.005 (16.02)
4.696(18.79)
4.642 (18.57)
4.643 (18.57)
4.664 ( I 8.65)
4.790(19.16)
4.666(18.66)
4.618(18.47)
4.692 (18.77)
4.858 (19.43)
4.63 1 (18.52)
4.633(18.53)
4.632 (18.S3)

( 19.03)
(0.99)
5.2

metabolizable

energy

acid

the findings

(7). This

ofthis

paper.

data are ofinterest

for two reasons:
the design of isocaloric, isonitrogenous
research
and therapeutic
diets requires
these computations
ifthe distribution
ofamino
acids is atypical, and the measurement
ofenergy
expenditure
and substrate
oxidation
rates by indirect
calorimetry
or measurement
of energy

expenditure

in free-living

subjects

by doubly

labeled

water

require
proper
values of 0:N,
H02,
and RQ for accuracy.
Table 5 shows the effect ofapplying
these values to the computation ofthe biologic energy content
ofseveral
amino acid mixtures.
variation
in amino acid composition
results in different
energy
contents
despite
similar
weight
percentages
of total
amino
chain

acids.
amino

For

example,

acids

have

solutions
a higher

enriched

caloric

in the

value

than

brancheddo formula-

tions enriched
in alanine
and glycine despite
similar
or lower
weight percentages
oftotal
amino acids. The essential
or indispensable
amino
acids have not only a high metabolizable
energy but also a high oxygen
5 includes
several
enteral

in addition
spondingly

to amino
higher

equivalent
products

acids,

for those

and

that

the

products.

ofurine
contain

nitrogen.
Table
carbohydrate

calculated
O:N

RQ

directly

is corre-

reflects

the

amino

acid composition
ofeach
diet.
It can be predicted
that it would be difficult to balance
both
calories
and nitrogen
among
test diets of a few amino
acids
chosen from the extremes
ofcaloric
density.
The most extreme

TABLE

Oxygen

equivalent

ofurine

nitrogen
used in production
ofurine
nitrogen5

on average

than

do the

nonessen-

acids.

3, the oxygen

with

Oxygen

Tables
2-4 present
the values for free amino
acids rather
than protein-equivalent
amino
acids. The heat equivalent
of
oxygen varies little among the amino acids (CV < 6%, Table 2),
but large variations
among amino acids in the values of oxygen
equivalent
of urine nitrogen
(CV 57%) and respiratory
coefficient (CV 18%) are shown in Tables 3 and 4. As demonstrated
in Table

per g free amino

These

Amino

higher

not

1.

prising
crystalline
amino
acids. We prepared
two parameter
files containing
either molecular
weights ofamino
acids or acyl
residue
formula
weights
for amino
acids, for use with actual
amino
acid weights
or protein-equivalent
weights
of amino
acids, respectively.
There is over twofold variation
in the values
for individual
amino acids, and the essential
amino acids have
tial amino

and

is in agreement

equivalent

of urine

whether
the urinary
metabolite
is urea
lated result was verified
as a general
monocarboxylic
acid. The mixture
and addition
of other caloric
sources
variation
in energy content
of natural
amino
acids usually contribute
only
normal
human
diets.

nitrogen

is the

same

or ammonia;
this calcurule for any monoamino
of multiple
amino
acids
will tend to minimize
diets, especially
because
10-20%
of the energy in

Between
the time of first review
and publication
of this
manuscript,
an Erratum
was published
stating that the metabolizable
energy density
of individual
amino
acids published
in
reference
5 of this paper was expressed
in units of energy per

acid

Set 1

Set 2
L 02/g

Alanine
Arginine
Asparagine
Aspartic
acid
Cysteine
Cystine
Glutamicacid
Glutamine
Glycine
Histidine
Isoleucine
Leucine
Lysine
Methionine
Ornithine
Phenylalanine
Proline

Serine

4.709
2.108
2.308
4.709
7.015
6.615
7.109
3.509
2.709
2.575
I 1.910
11.910
5.509
11.816
4.309
15.9 10
8.709

3.909

Threonine

6.309

Tryptophan

9. 109

Tyrosine
Valine
I
SD

CV (%)
S

Sets described

15. 1 10
9.509
7.154
4.117
57.5

in Table 1.

Set 3
N

4.801
2.200
2.400
4.801
7.105
6.705
7.201
3.600
2.800
2.667
12.002
12.002
5.601
11.906

4.801
2.200
2.400
4.801
7.105
6.705
7.201
3.600
2.800
2.667
12.002
12.002
5.601
11.906

4.401

4.401

16.002
8.801
4.001
6.401

16.002
8.801
4.001
6.401

9.20 1
15.202
9.601
7.245
4.117
56.8

9.201
15.202
9.601
7.245
4.117
56.8

Downloaded from ajcn.nutrition.org by guest on September 9, 2014

4.703 ( 18.8 1)
4.957 ( 19.83)
4.753 (19.0 1)
4.640 ( 18.56)
5.359 (21.44)
5.350 (2 1.40)
4.6 18 ( 18.47)
4.696 ( 18.78)
4. 1 30 ( 16.52)
4.827 ( 19.3 1)
4.67 1 ( 18.68)
4.672 (18.69)
4.726 ( 18.90)
4.8 18 ( 19.27)
4.745 ( 18.98)
4.640 ( 18.56)
4.732 ( 18.93)
4.945 ( 19.78)

Threonine
Tryptophan
Tyrosine

4.627 ( I 8.5 1 )
4.799 ( I 9. 19)
4.600 (1 8.40)
4.563 ( 18.25)
5.318 (21.27)
5.308 (2 1.23)
4.567 ( 18.27)
4.593 ( 18.37)
3.979 ( 1 5.9 1 )
4.693 ( 18.77)
4.64 1 ( 18.56)
4.642 (18.57)
4.662 ( 1 8.65)
4.790 ( 19. 16)
4.663 ( 18.65)
4.6 17 ( 18.47)
4.69 1 ( 18.76)
4.859 ( 19.44)
4.629 ( 1 8.52)
4.631 (18.53)
4.63 1 ( 18.52)

Alanine
Arginine

773

ACIDS

communication

kcal/L (ki/L)

AMINO

g protein

Heat equivalent

Amino

FREE

774

MAY

TABLE 4
Respiratory

quotient

for amino

acids

CO2 produced
per oxygen
consumed in catabolism5

AND

HILL
ERG. The precision
ofthe heat equivalent
ofoxygen
(Table 2)
supports
its use as a measure
ofcaloric
expenditure.
However,
the partitioning
of total oxygen
consumption
into carbohydrate and fat oxidation
requires
correction
of the oxygen utilization by subtracting
urine nitrogen
X O:N
As seen in Table
3, O:N
is highly
variable
among
the natural
amino
acids.
A
problematic
but critical question
is whether
the mix of amino
acids being oxidized
is the same under all conditions
or among
all subjects
in a research
study. Computation
of a correction
for urine nitrogen
from the dietary
amino
acid composition
implies
the assumption
that urine nitrogen
is derived
from
amino acids in the same relative proportions
as in the diet. This
assumption
may be true in a steady state in adult humans
or
nongrowing
animals
but is only one ofseveral
possible assumptions in states of nitrogen
accretion
(growth)
or negative
nitrogen balance
(starvation,
postinjury
or postoperative
catabolism). Other assumptions,
such as that the mix of amino acids
.

Aminoacid

Setl

Set2

Set3

L/L
0.835
0.727
1.011
1.175
0.554
0.588
1.004
0.893

1.000
1.091
1.333
1.333
0.667
0.706
1. 1 1 1
1. 1 1 1

0.833
0.727
1.000
1.167
0.556
0.589

Glycine

0.862

1.143

0.857

Histidine
Isoleucine
Leucine
Lysine
Methionine
Ornithine
Phenylalanine
Proline
Threonine
Tryptophan
Tyrosine
Valine
i

0.906
0.733
0.733
0.714
0.600
0.727
0.85 1
0.819
1.006
0.877
0.871
0.896
0.750
0.824

1.200
0.800
0.800
0.857
0.667
0.909
0.900
0.909
1.200
1.000
0.957
0.947
0.833
0.976

0.900
0.733
0.733
0.714
0.601
0.727
0.850
0.818
1.000
0.875
0.870
0.895
0.750
0.822

SD

0.151

0.196

0.148

Serine

CV(%)
5

18.3

Sets described

in Table

1.000
0.889

20.1

18.0

I.

being

oxidized

TABLES
Calorimetric

is distributed

variables

ofsome

Mixture

cial

roles

in the

modulation

ofa single amino acid among

because
the branched-chain
and valine) may have benefi-

of proteolysis

in severe

catabolic

states (9, 10), and the energy

content
of the branched-chain
amino
acids is high. For example,
the effects of five solutions
on nitrogen
balance
were studied
in traumatized
rats ( 1 1): each
solution
contained
(per L) 50 g glucose
and, in addition,
the
first had 1 5 g leucine,
the second had 15 g isoleucine,
the third
had 15 g valine, the fourth had 15 g alanine,
and the fifth had
an additional
I 5 g glucose.
The diets were thought
to be balanced for energy (assuming
all amino
acids and glucose had a
biologic
energy content
of4 kcal/g), but the actual energy contents of the five solutions
were 275, 275, 266, 238, and 242
kcal/L,
respectively.
The improvement
in nitrogen
balance
seen with the leucine
solution
compared
with the alanine
or
dextrose
solutions
was as likely to have been because
ofthe 14%
greater energy content
rather than special properties
of leucine
(10). METENERG
will allow such computations
to be done
easily

for many

other

special

diets

and

will

allow

those

compu-

tations to be done for different

distributions
ofnitrogen
among
urinary
metabolites
to allow
customization
for genetic
or
pathophysiologic
states
in which
urine
nitrogen
is shunted
away from urea.
The proper interpretation
ofthe measurement
of respiratory
gases depends
on several
parameters
computed
by ME TEN-

same

amino

as the amino

acid

distri-

acid mixtures5

ME

RQ

Heq

02:N

kcal/L

Travasol
AminoSyn

495
556

0.776
0.772

4.669
4.687

5.126
5.769

483

0.772

4.689

6.151

Aminoacidtt

344
282
266
442
27 1
341
4435ff

0.780
0.744
0.780
0.787
0.864
0.761
0.807

4.697
4.708
4.720
4.705
4.800
4.715
4.694

5.876
9.372
6.212
8.460
4.054
9.698
6.885

AA-HBCtt

8.5%t
8.S%t

HepatAmine
FreAmine III 8.S%
Nephramine
S.4%
TrophAmine6%
Indispensable-8.5%
Dispensable-8.S%i

fl

RDA mixture-6.36%55
imbalance
arises for comparisons
diets. This is of great interest
amino acids (leucine,
isoleucine,

the

Nutrisource
Nutrisource

47 1 8

0.780

4.706

7.206

HepaticAidII

1077

TravasorbRenalt
StandardVivonex
VivonexHighN
Amin-Aid

1170
764
758
1769

0.874
0.945
0.977
0.961
0.924

4.864
4.921
4.950
4.920
4.950

6.740
6.471
5.368
5.636
9.534

Traum-Aid

HBC

TravasorbHepatict
Caseclill

Traum-Aid
Isocallill

HN

Ensure11J
Vital HN1
S Values
creatinine,

calculated
for urine
2% uric acid.

997

0.909

4.917

7.633

1108
4605ff

0.937
0.822

4.964
5.397

6.723
6.544

949

0.925

4.912

7.0 14

1073
1064
41 16ff

0.870
0.882
0.940

4.861
4.878

6.755
6.503
6.077

nitrogen:

90% urea,

t Travenol,
Inc, Deerfield,
IL.
:t Abbott Laboratories,
Inc. North Chicago,
American
McGaw,
Inc, Santa Ana, CA.

4.943

3% ammonia,

5%

IL.

II His, lie, Leu, Lys, Met, Phe, Thr, Trp, Tyr, Val: 8.5 g/L each.
#{182}
Ala, Arg, Asn, Asp, Cys, Glu, GIn, Gly, Pro, Ser: 8.5 g/L each.
55

Recommended

dietary allowance

(8). 3.49 g Cys, 8.38 g Ile, 1 1.17

g Leu, 8.38 g Lys, 3.49 g Met, 5.59 g Phe, 5.59 g Thr,

Tyr, and 9.78 g Val in 1 L solution.

tt Sandoz Nutrition, Minneapolis.

if Dry powder (kcal/kg).
Norwich Eaton, Inc. Norwich, NY.
II II Mend Johnson Nutritionals,
Evansville,
#{182}11
Ross Laboratories,
Columbus,
OH.

IN.

2. 10 g Trp,

5.59 g

Downloaded from ajcn.nutrition.org by guest on September 9, 2014

Alanine
Arginine
Asparagine
Asparticacid
Cysteine
Cystine
Glutamic
acid
Glutamine

CALORIE
bution

ofwhole

are oxidized

result

body

protein

in preference
protein

metabolism

can

bohydrate

from

indirect

rates

of

difficult

of different
oxidation

affect

and

fat

end

of the

products

ofrates

acids

acids
would

number

in
acid

obtained

ofthe

oxidation
for the

ofdoubly

labeled

ide production

water

from

results

bonate

pool

from

which

the difference

of deuteriurn

curves

water,

and

in dilution

oxygen-

corn-

carbon

diox-

ofenrichment

to the water

pool

labeled

in

by the deute-

consumption

and

thus

As detailed
amino
acids

tion

and

in RQ

among

fats

to error

carbohydrates

(5).

of energy
expenditure
by doubly
labeled
also usually
includes
the assumption
that
net zero-energy
balance
during the period
may

be considered

part

of the

calculated
dietary
RQ ( 12). The
be appropriate
for the individual
ofoxidized
substrates,

substrates
and this

that

calculated

subject

The

of
is wide
varia-

RQ

equals

the

for the diet

will

only ifthe

is the same as the distribution

assumption
would
be untenable

distribution
of dietary
for sub-

jects altering
weight by deposition
or loss ofbody
tissue during
the period of study.
Urine nitrogen
can also reflect nucleic acid catabolism
and
excretion
of many non-amino
acid nitrogenous
compounds
contained
in natural
diets. Another
major assumption
of the
above discussion
is that urinary
nitrogenous
compounds
are
derived
from amino acids. This assumption
is valid whenever
the nitrogen
content
ofdietary
amino acids is overwhelmingly
large compared
with the nitrogen
content
of non-amino
acid
compounds.
Foods
or experimental
diets that contain
large
amounts
of non-amino
acid nitrogen
cannot
be treated
by the
equations
or program
described
above nor can they be allowed
in studies employing
indirect
calorimetry
for computation
of
substrate
oxidation
rates.
Finally,

we feel

that

proper

design

of amino

In: Sober

handbook

HA, ed.

CRC Press,

ofchemistry.

13th

ed.

1970:

New

York:

7th ed. Ardsley:

Ciba-

1985.

4. Diem K, Lentner
C, eds. Scientific
Geigy Corp 1975:663-72.
G, Elia

M. Estimation

tables.

ofenergy

expenditure,

net carbohy-

by indirect

to the detailed composition

offuels.
Am J Clin Nutr l988;47:608-28.
6. Consumer
and Food Economic
Institute.
Composition
of foods:
dairy and egg products.
Agricultural
handbook
no. 8-1. Washing-

ton, DC: USGovernment

PnntingOffice,

1976.

7. Erratum. Am J Clin Nutr l989;50:l475.

8. National
Research
Council.
Recommended
10th ed. Washington,
DC: National
Academy

dietary
allowances.
Press, 1989.

9. Brennan MF, Cerra F, Daly JM, et al. Report ofa research

shop: branched
chain amino acids in stress and injury.
1986;

workJPEN

10:446-52.

Buse MG. Effects ofbranched

chain amino
acids on protein turnover.
Diabetes
Metab Rev l989;S:227-45.
1 1. Freund
H, Yoshimura
N, Fischer JE. The role of alanine
in the
nitrogen
conserving
quality ofthe branched
chain amino acids in
the post injury state. J Surg Res 1980; 29:23-30.
12. Schoeller
DA. Measurement
of energy expenditure
in free-living
May

ME,

humans

by using doubly

labeled water. J Nutr 1988; 1 18:1278-89.

APPENDIX
Sample

computations

for an individual

amino

acid

1) Urine nitrogen
will be assumed
to be 90% urea,
nia, and 5% creatinine.
The molecular
formulas
of combustion
of these compounds
are
Urea

C1H4O1N2

l5l.Okcal/mol

Ammonia
Creatinine

C0H3O0N1
C4H701N3

70.6

kcal/mol

558.1

kcal/mol

2) The heat of combustion

pounds
is (0.9 x 151.0/2)
558.

1/3)

80.78

kcal/(mol

of urinary
+ (0.05
urine
N).

3) The C, H, and 0 lost in urinary

arec
=(0.9
X l/2)+(0.05
(mol/mol
N).

4)

5% ammoand heats

nitrogenous

70.6/1)

nitrogenous

X0/l)+(0.05

corn+ (0.05

0.5167

h = (0.9 x 4/2) + (0.05 x 3/ 1 ) + (0.05 x 7/3) = 2.0667

(mol/mol
N).
ou = (0.9 X 1/2) + (0.05
x 0/1) + (0.05 x 1/3) = 0.4667
(mol/mol
N).
The remaining
calculations
will be illustrated
with lysine.
The molecular
formula
of lysine is C6
14 N 202
The heat
ofcombustion
oflysine
is not tabulated
but can be calculated from the heat of formation.
The heat offormation
is
the enthalpy
change for the reaction

El

References
I. Widdowson
EM. Note on the calculation
of the energy value of
foods and of diets. In: Paul AA, Southgate
DA, eds. The composition of foods. 4th ed. New York: Elsevier/North-Holland
Biomedical Press, 1978:322-6.
2. Hutchens
JO. Heat of combustion,
enthalpy and free energy of for-

compounds
X4/3)=

acid-modified

research
diets will require
application
of the individual
energy
parameters
rather than assumption
of constant
values for all
amino
acids. We are making
METENERG
available
for this
purpose.

compounds.

2nd ed. Cleveland:

estimation

water decay curves

the subjects
are in a
under study, which

assumption
RQ

in estimation

in Table
4, there
as well as the known

related

6C+

7 H2 +02

The heat ofcombustion

+ N2-C6H14N202

is the enthalpy

change

for the reac-

tion

C6H 14N202
The summation

+ 02

6 CO2 + 7 H2O

#{248}

reaction

6C+7H2+02+N2-.+6C02+7H2O+N2

+ N2

Downloaded from ajcn.nutrition.org by guest on September 9, 2014

ofoxygen

total energy
expenditure.
variation
in RQ among

McGraw-Hill,

10.

num. Oxygen consumption

is then estimated
by assuming
that
the subjects
RQ equals the RQ of the subjects
usual diet. Errors in estimation
of the diet RQ can lead to errors in estimation

acids and

ofbiochemistry.

drate utilization,
and net fat oxidation and synthesis
calorimetry:
evaluation
of errors with special reference

18. The difference

in dilution
ofoxygen-l
8 into the bicar-

the distribution

in addition

estimates

ofamino

S. Livesey

mon end product

carbon
dioxide.
METENERG
allows cornputation
ofbounds
ofvalues
ofO:N
and Heq for differing
assumed proportions
ofamino
acids.
An alternative
technique
for measurement
of energy expenditure in living subjects is determination
ofthe decay of specific
activity

775

ACIDS

B62-4.
3. Dean JA, ed. Langes

is technically

of tracers

AMINO

mation

of car-

expenditure

FREE

Handbook

Errors

ofamino

simultaneously

limited

amino

ofoxidation

energy

OF

acids,

for 0:N

(5). Determination

amino

because

from

of total

amino

values

the estimates

and

calorimetry

multiple

dispensable

to indispensable

in computation

estimating

or that

CONTENT

776
has an enthalpy
products,
(Heat
=

that

is a sum

of heats

of formation

MAY AND
of the

ie,
+ (heat

of formation)L,

6 X (heat

of formation)co2

Substituting
get

the known

of combustion)L5
+ 7

values

X (heat

for heats

HILL
Because food items are usually specified
by weight,
vert this result by division
by the molecular
weight
(146.19)
to 4.933 kcal/g.
6) The CO2 produced
by oxidation
of lysine is

of formation)H20

(6

of formation,

we

7) The

(14
(Heat

of combustion)L,

[6 X 94.38

H2O
-

X 2) mol/mol

0.5167

produced
X 2)/2

2.0667

162.2]

kcal/mol

The metabolizable
energy is the heat of combustion
the heat of combustion
of urinary
products,
ie,
882.7

kcal/mol

Lys

X (2 mol

(80.78
N/mol

kcal/mol
Lys)

of lysine

LYS

consumed

mol/mol

LYS

is

4.9333

mol/mol

LYS

9)

kcal/mol

+ 4.9333

+ 0.4667

of lysine

is

minus

N)
721.14

in catabolism

882.7 kcal/mol
(2 X 4.9667

5)

by oxidation

mol/mol

4.9667

+ 7 X 68.38

8) The oxygen
-

LYS

we conof lysine

Lys

10)
11)

RQ

Heq
O:N

=
=

2)/2 mol/mol

4.9667/6.9000
721.14/6.9000
=

6.900

LYS

0.720
L
X 22.414

X 22.414/14.0067

6.9000

C02/L
=

LYS.

02

43.663
X 2

mol/mol

kcal/L
5.521

02
LO2/gN

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