between 25–45 years. Thereafter, less child care and
career progression, leading to desk-bound work and
retirement, may further curtail activity. In addition,
BMR falls 2% per decade as lean tissue mass de-
creases. In very old age, incapacity restricts the ability
to be active and PAL levels may be only 1.2. Ill health
at other parts of the life cycle may also affect activity
and TDEE. In many parts of the world, infection not
only causes personal misery but also seriously affects
subsistence and occupational activity. Pregnancy has
the twin characteristics of an increasing body size plus
a customary fall in activity, particularly in the last
trimester. Therefore the total extra cost of pregnancy
may be less than expected, unless the woman does not
reduce activity.
0017 The contribution of genetic factors to differences in
energy expenditure is very difficult to investigate and
whether any genetic differences are actually ex-
pressed in the individual or population depends on a
multitude of factors. Results from twin and other
studies suggest that variations in RMR, thermic effect
of food, level of physical activity, and energy cost of
light exercise have a low to moderate genetic com-
ponent. There are frequent reports in the literature of
low BMR in tropical populations, particularly from
the Indian subcontinent. Whether these arise from
differences in size and composition and nutritional
status, or ethnic differences per se, is controversial.
Some groups have a renowned propensity for fatness,
and a hypothesis of a thrifty genotype has been put
forward to explain massive overweight and obesity
in, for example, Polynesians and the Pima Indians
of Arizona, USA, apparently as they altered their
traditional lifestyle and took up western habits. The
hypothesis suggests that, in response to chronic food
shortages over many generations, individuals with
more efficient energy utilization survived and bred
selectively. Now that high energy intake can be
achieved with low energy expenditure, the superior
efficiency results in high energy stores. To support the
hypothesis, evidence of a lower energy turnover or
increased efficiency is required. In Pima Indians some
families do have lower RMR than others and these
show higher rates of obesity. But, a major problem in
human energetics is how to compare the rates of
energy expenditure in individuals of different sizes,
whether the small, thin individuals in India with ap-
parently low BMR or these large, obese individuals,
where low BMR is also being sought. The problem is
less difficult for physical activity as the net mechan-
ical efficiency (NME ¼ work done divided by energy
expended) is an appropriate index of efficiency. Al-
though NME varies between individuals for reasons
that are not entirely clear, there is no consistent pat-
tern of evidence suggesting that the body can alter
NME in response to varying planes of nutrition in the
short term or over many generations. This attractive
hypothesis of a thrifty genotype spread throughout
the population appears evolutionarily sound but
remains a hypothesis.
0018Owing to the pervasiveness of modern communi-
cation, all societies and populations experienced
changes in lifestyle in the 20th century. For many this
has meant a shift from subsistence lifestyles, where all
material needs were met by 2–3days’ work per week,
to long hours of repetitive, often hard, poorly paid
labor. In the developed economies, the trend has con-
tinued towards increased mechanization, a fall in the
length of the working week, and increased holiday
entitlements, which have reduced the effort in work,
particularly the peak- and high-intensity efforts. The
effect of this on TDEE is difficult to gauge. Surpris-
ingly, analyses suggest there are no differences in levels
of habitual physical activity, as evidenced by PALs,
between the developed and the developing world.
This might be explained, in part, by the curtailment
of physical activity as a behavioral adaptation to the
presence of chronic energy deficiency. It does high-
light, however, the pitfalls of taking as self-evident
beliefs about energy expenditure and physical activity.
There have been few representative national studies of
current-day activity and expenditure patterns, let
alone studies for earlier periods of the last century.
One way round this is to ignore the small effect of
the greater energy stores in modern populations and to
assume that energy intakes reflect energy expenditure.
Again, there are no national representative intake data
from earlier years but the evidence suggests that levels
of energy expenditure have fallen, albeit only in the
last 10–20 years.
Control of Energy Expenditure and
Regulation of Energy Balance
0019There are several types of factors operating at differ-
ent levels. At the primary level, the setting of regula-
tory centers in the brain or mechanisms at the cellular
level may ultimately determine the level of energy
balance. These respond to changes in energy intake
and expenditure, at a secondary physiological, bio-
chemical, or neurophysiological level. Traditionally,
the regulation of energy balance has been seen as
energy intake being controlled to match antecedent
levels of energy expenditure. It has been argued
recently that regulation of energy balance is less ac-
curate when the energy intake is high in fat, as char-
acterizes many contemporary diets, than for diets
with low-fat energy and that this contributes to the
modern rise in the prevalence of obesity. In contrast,
attention has also been paid to the possibility of the
2106 ENERGY/Energy Expenditure and Energy Balance