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Use and storage
Jean-Pierre
ABSTRACT
at rates
Starch,
appropriate
matching
sugars,
body’s
stores
is regulated
carbohydrate
are
fat
tissue
mass,
which
one
eats.
oxidation
so small
J Clin
be
Nutr
alter
and
to
because
free
oxidize
as
WORDS
acid
con-
Fat intake
and
in determining
fat
as
one
1995;61(suppl):952S-9S.
body
fat content,
respiratory
Dietary
carbohydrate,
weight
maintenance,
quotient,
obesity,
dietary
fat, glycogen,
macronutrient
balance,
exercise
The
distributed
with
fatty
their
other
energy
most
to
These
the
which
must
by
the
nervous
use
therefore
are
are used
used
central
cannot
they
fuels
ATP
the
cells
energy,
in which
metabolic
However,
The human
body
adjusts
in the relative
proportions
dietary
form
regenerate
specialized
needs,
of the
are the main
activities.
a few
glucose.
ferences
provide
the body.
efficiency
metabolic
and
meet
fats
acids
throughout
similar
body’s
tem
and
and
fatty
sysacids
be supplied
to
with
readily
to considerable
of canbohydnate
and
fat
difin
the diet, even when
major
changes
in the carbohydrate-to-fat
ratio
occur
(1). Because
the fraction
of total dietary
energy
provided
by protein
is relatively
small
and relatively
constant,
and
because
stant
amino
protein
acid
termined
The
the
body
spontaneously
maintains
content
by adjusting
amino
intake,
body
weight
maintenance
by the
intake
regulation
many
factors
some
are
are
and
utilization
of body
weight
in preventing
inherited,
and
others
a nearly
is a complex
are
and
problem
or causing
solve
is to review
olism
for
ents play
9525
and
the
a better
in the
dietary
main
variables
related
features
understanding
maintenance
(2).
aim
of carbohydrate
J Clin
evaluation
dietary
of
in
various
carbohydrate
forms
of edible
and
carbohydrates
tubers.
They
energy
needs,
except
in affluent
is a major
source
of energy
almost
the
when
and
fat
as much
particular
processing
important
rate of
provide
are
the
the bulk
Western
and where
societies
sucrose
energy
as starch
(3). It is helpful
problem
that the organism
has
carbohydrates.
For
first be broken
exert a powerful
that
starch
carbohydrates
to
to
to be
down
into monosaccharosmotic
effect
and it is
intestinal
absorption
be able
hydrolysis.
Furthermore,
to keep up
absorption
with
the
must
be completed
before
tenial
fermentation;
otherwise,
such
as those experienced
after lactose
ingestion
by individuals
low intestinal
lactase
activity.
Meals
commonly
supply
carbohydrates
g carbohydrate.
glucose
there
This
present
in the
is
much
rapidly
cells.
transferred
into
state
are
blood
glucose
expended
glucose
of glucose
must
Glycogen
incorporate
rise
1
trapped
fat.
carbohydrates,
and
glycogen.
much
be expended
molecules
in the
cells
mol
The
(4).
Two
they
ATE
glycogen.
are thus
made
of the
associated
effectively
without
creating
hydrophilic
with
of glycogen
density
moles
into
complete
oxidation
of
of the energy
content
were
In view
is
energy
in the
of the glucose
it as glycogen.
large;
they
effect.
glycogen
oxidation
most
glucose
to store
are very
in which
of an osmotic
of glucose
must be
conversion
to glycogen,
primarily
uptake
and glycogen
synthesis
are
the secretion
of which
increases
concentrations
to
effects,
the 15-25
g
5 g is in the
than
spillage
circulation
10 g/h,
to bac-
side
only
and
the
into
only
taken
up must be stoned by
in liver and muscle.
Glucose
greatly
stimulated
by insulin,
when
more
Because
consumes
subjected
of which
hyperglycemia
transferred
postprandial
are
are unpleasant
body,
blood
itself.
To avoid
into the urine,
glucose
to
de-
and
of this
to
and
fat
article
nature
‘3
stores
g
of
water/g
is thus
only
Nuir
From
the
University
of
Massachusetts
Medical
School,
Department
of
Worcester,
MA.
metab-
of the roles these macronutniof stable
body
weights
and
Am
stabiliza-
important
obesity-
to lifestyle
The
the
of
by grains
absorbed,
they must
ides. Small
molecules
I
socioeconomic
for
weight
is
Because
36 ATP are gained
during
the
one molecule
of glucose,
2/36, or ‘“5%
con-
acid
oxidation
is primarily
of carbohydrate
involved
basis
effect
abundant
provided
supplies
understand
free
glucose
understanding
storage
most
starches
50-150
Introduction
Carbohydrates
at which
Such
regulation.
Carbohydrate
with
and
the
weight
therefore
KEY
rational
about
body
of human
where
fat
in adipose
much
a
composition
mech-
fatty
fat oxidation.
are important
occur.
on
cumulative
to changes
body
to
tends
arguments
pertaining
to the
by fat intake.
Ad-
occurs
time
the
tion
mix
in the diet.
regulatory
influencing
providing
of a fuel
that
events
than
substantially
to
the
of the
later
carbohydrate
oxidation
through
evolution.
Fat
by
rather
oven
sensitivity,
concentnations
has
Am
lead
can
provide
distribution
to intake
balance
insulin
glycogen
fat
the
adjusting
developed
primarily
economy,
of fat oxidation
in the
centnation,
habitual
about
of efficiently
intake
have
errors
triglycerides
the macronutnient
glycogen
justment
and
preserve
homeostasis,
most
must be stored
to be mobilized
in
1995;61(suppl):952S-95.
2
Supported
3
Address
University
by NIH
reprint
requests
of Massachusetts
Printed
grant
in USA.
DK
33214.
to I-P
Medical
© 1995
Flatt,
Center,
American
Worcester,
Society
for
Biochemistry,
MA
Clinical
01655.
Nutrition
Downloaded from ajcn.nutrition.org by guest on August 29, 2014
oxidation
body’s
To
to bring
on average
anisms
capable
to carbohydrate
how
and fat3
Flatt
bulk of dietary
energy.
glucose
and fat absonbed
The
of carbohydrate
USE
“4.2
kJ/g
1 kcal/g),
(::
of energy
glycogen.
imposing
that can
Glycogen
typically
‘‘4%
definite
conveniently
concentrations
after
STORAGE
limits
fast,
and
OF
on the
be carried
are highest
an overnight
(5).
AND
amount
in the
in the
up to 8%
AND
of tissue,
form
liver,
after
CARBOHYDRATE
of
ie,
carry
making
weighs
“‘1.5 kg, hepatic
glycogen
to “‘120
g. Glycogen
amounts
in
bile
and
and
intestinal
muscle
reesterified
necessary
muscle
much
lower
and
deliberate
to raise
them
much
above
accounts
of glycogen
that
for
20-30%
stored
in the
Total
glycogen
the day
glycogen
the amount
maintenance
effective
carbohydrate
Marked
liver
to ensure
trations,
which
of
depletion
adequate
requires
can
creted
amount
form
on body
and vary
and
much
times
thus
be
size and
substantially
on
exertion.
greater
degraded
lipases.
into
into
the
circulation.
few
hours
femned
These
coating
after
cells,
the
an enzyme
produced
of adipose
tissue
causes
a threat
to the
blood
glucose
release
of
to
ability
of
concen-
5 g glucose/h
(or
are
most
fatty
acids
able
as a metabolic
(10),
fatty
fat
up by the
the
in which
fuel.
Dietary
in adipose
a
be trans-
chylomicrons
must
lipase
be
(LPL),
endothelial
cells
acids
produced
by LPL
cells
and
reestenified,
only
the
pool
of
to reach
form
in
appearance
To
by the
The
se-
in the
to remain
milky
by lipoprotein
abundantly
escaping
deposition
out
lumen,
are solubilized
consumed.
of the
carried
taken
fraction
for
triglycerides
of
and
mesentery
them
its
been
the
cells,
droplets
plasma
has
capillaries.
from
the
lipid
help
by pancreatic
intestinal
allowing
the
meal
a function
promptly
small
give
a fatty
consumed
in 1 d, and
within
a desirable
range
oxidation
tiny
the
glucose
draining
of lipoproteins,
They
into
the
vessels,
of chylomicrons.
a thin
with
absorbed
by
the lymphatic
by
are
triglycerides
hydrolyzed,
carbohydrate
These
and
survival
(4).
intestine
acids
and
to allow
deprivation
in the
to fatty
to accumulate
enough
of food
emulsified
The
than
circulating
the
because
to four
in adults
is
then
animals
fatty
acids
fatty
a
albumin-bound
are
acids
made
are
avail-
thus
targeted
tissue.
100 mg/mm)
to replace
the glucose
removed
by the nervous
system.
Because
the influx
of nutrients
from
the intestine
occurs
through
tunity
to replenish
the
carbohydrates.
carbohydrates
glucose
relative
reach
roles
vary
glycogen
carbohydrate
that after
depending
the
consumed
a typical
meal
is converted
to
oxidized
during
in
Conversion
the
the
of
of depletion
meal,
and
the
in the
muscle,
liver
whereas
postpnandial
type
the
one-third
oils
and
fats
containing
energy
three
precursors
solubility
maining
large
in
fat
a small
during
that
not
aqueous
can
make
active
whether
be stored
up
of capillary
meates
the
This allows
adults
ingestion
of
500
quotient
(RQ)
the
unusually
Thus,
primarily
The
body’s
33.5
blood
influence
present
their
and
nerve
or
of
conversion
Two
water.
can
of
of
ex-
terminals
reduces
per-
‘‘85%
of which
is fat.
to be stored
per gram
fat
glycogen
maximal
as the
cost
of
on
from
wt.
into
then
a fuel
remains
temporary
to
accu-
rates
of
total
their
(as
close
car-
glycogen
usual
4-6
g/kg
This
requires
deliberate
of large
amounts
of carbo-
need
will
frequently
that
to use
be
are
lipogenesis
fat
factor
be
can
stores
high-carbohydrate
is an important
consumed
as
RQ
body’s
conditions
readily
however,
the
are
stores
substantial
the
usual
capacity,
are
as well
overlooked,
intake
body
1)
loads
glucose
glucose
such
induce
conclusions
their
metabolic
their number
(9). A delicate
the
raised,
under
carbohydrates
dissipation
of
that
fat,
(Figure
d (13).
1)
below
missed
To
g/kg
important
to food,
that
use
reducing
into
2-3
observations:
far
most
layer
fact
absorbed
1.0,
at a rate
oxidation
the
the
above
carbohydrate
of the glycogen
ovenconsumption
for
fat
after
consumed
rise
expansion
be considerably
sustained
to 02
not
synthesized
occasional
in time
8-10
being
of
Even
respiratory
converting
glycogen.
to >
are
exclusive
1 by
hours),
must
acids
rate
by
bohydrate
and
fatty
In
an insignifi-
(1 1).
does
fat.
a hepatic
nonprotein
metabolism)
large,
mulations
cess
triglycerides
adipocytes
many
me-
in small
bound
1.0
wt
of CO2
to build
little
plays
diets
produced
nearly
in Figure
hydrates
substrate
by itself
than
by a thin
but once formed,
only their size
vessels
in-
tissue,
These
adipose
tissue,
kJ (8 kcal)
energy
(on
of
to the
conversion
mixed
ratio
fat
animals
very
is primarily
the
niassive
shown
stores
this
concomitant
glycogen.
body
lipogenesis
the
that
handled
allows
contains
g carbohydrate,
and
(12).
fat
feed
consuming
exceeding
amount
de novo
(ie,
indicates
for
their
Quantitatively,
in
is
starvation
without
adipose
cytoplasm.
pand
or shrink
considerably,
cannot
be reduced
by fasting,
network
(5).
role
which
are
compartments,
form a central
fat droplet
(much
larger
the body’s
other
cells)
that is surrounded
metabolically
of
fat
into
when
such
to
esteni-
critical
even
by carbohydrate
accounts
for
(or 5% of
Because
do
body’s
fats
but
(4).
unnoticed
and
cells
animals
molecules
periods
triglycerides
the
metabolically
amounts,
the
acids
and
into
of carbohydrate
humans
process
of
remainder
The glycerol
triglycerides
deprivation)
in water,
plants
fatty
providing
carbohydrate
concentrations
by
of glycerol.
of long-chain
content),
glucogenic
severe
adult
cant
hours.
accumulated
fied to one molecule
10% of the weight
their
Conversion
up fat reserves
of
to envision
of the carbo-
and
of carbohydrate
of
leads
triglycerides
In
of
degree
(6-8).
It seems
reasonable
one-quarter
to one-third
glycogen
oppon-
storage
The
of
a prime
consumption
only
a fraction
of
substantial
amounts
on the
size
has
after
are stored
in muscle.
The
in the disposition
of a car-
to glycogen
the
liver
this demands
by a meal and
stores,
one-half
Fat
the
reserve
the periphery
and
of liver and muscle
load
hydrate
system,
its glycogen
Usually,
provided
bohydrate
the
portal
retained.
deduced
from
these
of unrestricted
spontaneously
and
ac-
maintained
2) the
turned
into
made
argument
common
fat
belief
can
be
dis-
that
the
high
is a cause
for
greater
energy
diets.
It
should
not
consumption
as a fuel,
in determining
of
so
be
carbohydrates
that
how
carbohydrate
much
of the
Downloaded from ajcn.nutrition.org by guest on August 29, 2014
the
usually
amounts
the
three
stones
adjustment
intake.
glycogen
weight,
of food intake
is in effect
not
of carbohydrate
of glycogen
requires
body
g, depending
consumed,
as a function
reserve
loading
However,
is generally
estimated
to be “200-500
the amount
of carbohydrate
during
body’s
2% (5).
of total
in muscle
liver.
carbohydrate
are
for
substantial
periods
fats
Because
an adult’s
liver
storage
capacity
is limited
are
of fuel
extended
Dietary
953S
it possible
a reserve
during
meals
FAT
954S
FLATT
gluconeogenesis
which
in the
meal
1#{149}1
C
e
glucose
uptake
also
a signal
to restrain
provides
glucose
phorylase
0
09
0.
in muscle
during
muscle
permit
rapid
To
0
a
C
is activated
ATP
the
to
provide
a
5)
05
0
body’s
major
under
Maintenance
30L
ad
.J
0
E
E OS
the
in
because
of carbohydrates.
foods
to
al-
often
As
after
carbohydrate
intake.
glucosunia
is effec-
and
nutritional
these
and
mechanisms
of glucose
maintaining
storage,
effec-
mobilization,
glycogen
glycogen
metabolic
operate
amounts
amounts
when
in
there
an
is free
facilitated
if adjustment
intake
were complemented
of
to diminish
or enhance
food intake
when
and/or
glycogen
reserves
rise on decline
16),
of regulation
intake
from
but
attempts
appears
to operate
day to day in mice
to demonstrate
in
fed
a similar
in humans
have so far been unsuccessful
(1). What
for humans
is that hypoglycemia
induces
an ex-
tremely
free-living
powerful
urge
to find
food,
that
individuals
is such that glycogen
(on
03
rates
reserves
concentraevolution
fuel
occurs
would
be greatly
to carbohydrate
(15,
glycogen
glucose
through
behavior
is known
well
tamed
04
to
as well
form
differing
that
(14). This type
changes
in food
libitum
glycolysis
after carbohydrate
ingestion
by glucose
to the fuel mix
rapidly
of stable
to food
oxidation
excessively
modulating
0#{149}
7
in the
widely
by signals
working
glucose
concentnations
SO
locally
whenever
food
intake
fails
Yet glucose
must
also be
rarely
it is evident
access
glucose
Phos-
primarily
available
to maintain
metabolic
energy
increases
and oxidation
and
appropriate
mange.
C
.
the
in adjusting
tively
120
anaerobic
oxidation
reserve.
of their
avoided
conditions,
E
by
ability
hypoglycemia
tively
150
by catecholamines,
glucose
when
(4).
below
decreases)
decreases
their
in
(or
the
increases)
maximal
diet’s
food
stores
capacity,
and
carbohydrate
in energy
intake
in
are main-
that
increases
content
intake
lead
to
(17-19).
02
0#{149}1
hours
Fat
oxidation
Fat
FIGURE
glucose,
1. Changes
and
in six young
men
carbohydrate
with
in plasma
in response
(white
permission
in the
insulin
bread,
nonprotein
and
free
respiratory
fatty
to a breakfast
jam,
and
fruit
acid
quotient,
concentrations
providing
juice).
8372
i ±
in blood
observed
kJ (2000
SEM.
kcal)
Reproduced
total
oxidation
energy
the form
Because
critical,
effectively
concentrations
erates
the
phosphorylase
glucose
regulatory
prevent
thus
rate
and
regulation
can
economy
intake,
is constantly
mechanisms
hypoglycemia.
elicit the release
of glucose
to degrade
rates
of the
release
glycogen,
used
and
evolved
its availability
that
are
able
Declining
of glucagon,
by
blood
glucose
which
accel-
the liver
by activating
as well as by enhancing
is
to
primarily
and
of carbohydrate
oxidation
themselves
oxidation
balance
is determined
expenditure
that
activity,
Carbohydrate
carbohydrate
fat balance
the
and protein,
the
is determined
amino
acid
to the amounts
readily
selected
and
of the
fat
between
ingested
than
by the amount
body
size
oxidation
and that
unrelated
because
the body’s
fat reserves
on the other
hand,
is a function
foods
gap
in
This may seem surprising
of the facts
that overall
by
be set by indexes
the
of energy
rather
and glucose
consumed,
amounts
and it is not likely
to be influenced
losses
of fat (21), which
are trivial
large
by
amount
of fat consumed
on a given day (20).
at first,
but it is the consequence
substrate
(12).
and
and
physical
mates adjust
fat oxidation
to the body’s
are very
of the fat
large.
content
of foods
consumed,
by short-term
compared
with
fat
Fat
of
gains
or
the body’s
stores.
The lack of direct
tion and fat intake
regulatory
interactions
is demonstrated
by
the
between
fat oxidachanges
in fuel
Downloaded from ajcn.nutrition.org by guest on August 29, 2014
4
high
sustain
adequate
blood
mechanisms
developed
indeed
Because
ii
sufficiently
by a rapid
rise in the RQ
1), the contribution
made
oxidized
0)
-J
be
most
shown
(Figure
E
glycogen,
release
of glucose.
ensure
lowed
0
into
glucose
to make
nesynthesis
that restrict
glucose
replenish
the glycogen
-J
(6, 7). Insulin,
hyperglycemia
its conversion
hepatic
are
contraction
that can sufficiently
tions
at all times,
0.7
and
concentrations
as by oxidation
..C
amino
acid and lactate
state acts to prevent
by promoting
blood
10
from
postpnandial
USE
utilization
induced
sumed
by
together
stimulating
drate,
iment
secretion
but a decrease
described
in
and
a breakfast
affected
by
postpmandial
the
use
addition
of
of metabolic
of the RQ
as a function
(22).
The
effect
ever,
because
with
such
attenuates
(but
does
and
postprandial
fat
of
rise
and
concentrations
75
g carbohydrate
is essentially
fat
a dose
fat
are
not prevent)
RQ,
a decrease
by
0
and the
evolution
20
fact
slow
that
dietary
deposition
in
other
although
of fat
how-
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Figures
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and its effect
on raising
As
amount
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the
diet.
without
the
need
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(46).
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goal
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goal
the
body’s
metabolism
and amino
acids
this
expansion
is more
as are
steady
of the
readily
state
adipose
achieved
when
a
diet with a high carbohydrate
content
is consumed
because
it is
obviously
easier to burn as much fat as one eats when one does
not eat a lot of fat. Exercise
provides
additional
leverage
it causes
a greater
the
specific
increase
in the oxidation
the body will spontaneously
and carbohydrate
balances,
sense to direct voluntary
Furthermore,
concentrating
restrict
that
acids,
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regulate
because
control
fatty
The
to influ-
to
weight
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and
maintenance
A varied
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intensity
selection
as much
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Food
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weight
Macronutrient
because
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periods
activity
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Weight
in
of weight
as in animals
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physical
oxidize
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are
appetizing
as well
must
after
which
in glycogen
at a higher
societies
but
spontaneous
average
to
of
they provide
macnonutnient
fats, and protein
all contrib-
highly
palatability,
intake
for
state
but
fat content,
of
changes
the steady
them
conditions
spontaneously
tissue
mass,
compensate
of energy.
libitum,
the
(superman-
causes
under
by
in affluent
ad
in
fat balance
goal
than
nearly
of fat than
and effectively
mainbut does not (or cannot)
as accurately
(14-24),
it makes
efforts
at this relatively
soft target.
on the fat balance
provides
a more
one
based
on
overall
energy
exchanges.
further
food intake than when faced with unappealing
foods
(42). This tends to raise the upper limit of the range within
which
glycogen
concentrations
are habitually
maintained.
In
addition,
the ubiquitous
availability
of foods, by inducing
eating between
meals,
can have the effect of keeping
glycogen
concentrations
high. Given the ensuing
curtailment
of fat oxidation, fat will accumulate
until expansion
of the adipose tissue
mass is sufficient
to bring about a rate of fat oxidation
com-
Awareness
mensurate
with
maintenance
this effect
to become
established.
if one considers
that the
lowering
blood cholesterol
concentrations
and in minimizing
the development
of arteriosclerosis
(48, 49). To these reasons
for restricting
fat intake one can add the further argument
that
gen is maintained
reserves
fat intake,
permitting
is not precisely
are maintained
the steady
of weight
There
is much
room
for
range
within
which
glyco-
determined
far below
state
their
because
level
glycogen
of saturation
under conditions
of unrestricted
access to food (12,
influence
of the palatability,
variety,
and accessibility
teristics
tions,
of the
and hence
sideration
when
affluent
facilitate
Effect
The
food
supply
on
on fat oxidation,
accounting
habit
generally
weight,
is generally
need
for the high
societies,
as well
weight
control.
ofphysical
habitual
as when
glycogen
setting
into
up conditions
of engaging
not well
of
steady
adipose
that
low-fat
have
diets
because
limiting
fat
saturated
fats)
appears
state
balance
tissue
to
nary
stores
a particularly
provide
about
the
to satisfy
the possible
carbohydrates.
decline
the
of
foods
intake
more
high
bulk,
led
glucose
the
a key
can
diet’s
be
hunger,
with
disadvantage
is low.
on
of caution
inherent
known
for
smaller
of relying
a word
that
concentrations
in
expected,
fat content
advantages
con-
measure
to be achieved
It is now
in blood
(particularly
to be
maintenance
reasons,
when
consideration
be included
absorbed
of weight
fuel
In the
in
high-
should
to rapidly
a slight
tempo-
is a physiological
event that can lead to initiation
of food intake
(50, 51). In
animals
consuming
a feed of fixed composition,
the time
elapsed when this event occurs is proportional
to the size of the
activity
quite effective
or in inducing
sumption
carbohydrate
con-
and
associations,
metabolic
13). The
charac-
of obesity
density,
foods
dietitians
long ago to recommend
low-fat
diets for weight
control
(47). Recommendations
to reduce
fat intake have indeed been increasingly
promulgated
in dietary guidelines
published by government
agencies
and medical
on health-related
the
even
concentra-
to be taken
incidence
energy
of the fact that fatty
in substantial
in limiting
loss of excess
understood
why
physical
the accumulation
adipose
tissue
increments
activities
is
of excess
(43). Yet it
in food
intake
elicited
by exercise
are sometimes
sufficient
to ensure weight
maintenance
(ie, in a physically
active individual),
but sometimes not (ie, when someone
initiates
a physical
training
pro-
preceding
meal
(50).
It may
carbohydrate
influx from
from a glucose-removing
Rapidly
absorbed
this signal
carbohydrates
to occur
view
of the high
elicit
and
not only
degree
the rapid
rates
well
be
linked
to
the
the gut, when the liver has
state to a glucose-releasing
would
earlier,
of postpnandial
of peripheral
be
expected
but also
more
insulinization
glucose
end
of
to shift
state.
to cause
sharply,
that
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in
they
in-
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a relatively
ably
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consumed
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pable
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into fatty
but to eat
uals aiming
ductive
in
impact of
effective
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difficulty
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fat
oxi-
most
reduction,
complemented
by efforts
increasing)
carbohydrate
select
foods
high
can
be
not
entirely;
based
in
(8).
that
a reduction
a long-term
adipose
ca-
tissue
weight
control
a reduction
foods
occurs
in fat intake
to limit (at least
intake, to reduce
to the extent
sugar intake,
intake
P, Sj#{246}strdm L. Carbohydrate
and
quantitative
ethanol
reduces
fat but
It is of course
feasible
to reduce
energy
intake.
Selecting
foods
not
Sargent
and
D
RA,
in humans.
obesity.
Carbohydrate
in humans.
I Clin
Am
Kuwajima
liver
Nutr
deter-
1985;41:1132-45.
Diabetes
1971;20:785-99.
in man:
Metabolism
Rev
speculations
1978;27:1853-65.
of hepatic
glucose
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