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Encephalization

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DOI: 10.1002/9781118584538.ieba0155

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Encephalization
RALPH L. HOLLOWAY
Columbia University, USA
Encephalization can be defined as the amount of
an animal’s brain mass relative to its total body
mass, or the process of increase in brain size
over time. The relative size of an animal’s brain
is often taken to be some sort of measure of its
intelligence. This general idea goes back at least
to Aristotle (Norvig 2003), but its quantification
was first realized by Snell in the late nineteenth
century (Snell 1891). It is a relativistic concept,
whose values depend entirely upon the dataset
that is used.
Encephalization is sometimes expressed as a
simple brain–body weight ratio, but this fails to
account for the fact that, other things being equal,
small animals tend to have relatively larger brains.
One way of taking this allometric relationship
into account is to express relative brain size as
an encephalization quotient (EQ). In order to
calculate EQs for a species, three measures are
necessary: (1) brain size (either mass or volume),
(2) body mass, and (3) an equation based on some
theoretical expectation or empirical dataset that
has multiple members of some taxonomic group
above the species level, for example, the Order
Primate. Two major examples of such datasets are
those of Martin (1981), which is based on all then
known species of primates, and that of Jerison
(1973), based on all mammals. The data for body
and brain weights are plotted against each other,
with the body weight as the abscissa (x-axis) and
the brain as the ordinate (y-axis). This is usually
done in logarithmic units. Without the log units
the data scatter will follow a curved distribution,
becoming steeper with higher values. A linear
regression is then calculated for the entire scatter
or some subset, or imposed on it out of theoretical
considerations, and the EQ is expressed for each
species as a ratio of observed brain size to that
predicted from the overall regression.
The International Encyclopedia of Biological Anthropology. Edited by Wenda Trevathan.
© 2018 John Wiley & Sons, Inc. Published 2018 by John Wiley & Sons, Inc.
DOI: 10.1002/9781118584538.ieba0155
Thus, when Jerison examined the polygon of
log values for his dataset, he placed a line with
a slope of 2/3 through the data which had been
derived from basal mammals and then drawn
through his polygon of values for all mammals,
yielding the following equation:
Brain weight = 0.12 × body weight0.667
When Martin (1981) plotted all of the brain and
body weights for primates, his formula became:
Brain weight = 0.28 × body weight0.76
In Jerison’s formula, the 0.667 ratio suggests an
area-to-volume relationship, whereas Martin’s
0.76 exponent points to a relationship with basal
metabolic rates, which is the 3/4 power of body
mass within eutherian mammals and other,
similar mid-level taxonomic groupings (Kleiber
1947).
While these data appear relatively recent, EQ in
Jerison’s formula thus becomes:
EQ = brain weight ∕ 0.12 (body weight)0.66
Some examples of EQs calculated for different
animals are: human = 7.4, dolphin = 4.14, chim-
panzee = 2.5, rhesus monkey = 2.1, elephant =
1.13, dog = 1.2, squirrel = 1.1, and cat = 1.0 (from
Wikipedia contributors 2016).
While interesting, these make little sense for
understanding the human EQ relative to other
animals. If one takes the world average of human
brain weight as 1,330 g and body weight as 65,000
g (65 kg), and assumes that if there is zero body
weight there is zero brain weight, plotting the
logs of these values gives an equation such as:
EQ = brain weight ∕ 1.0 × body weight0.6409
This equation immediately provides the percent-
age value of the human EQ calculated as 100 per-
cent.
The following quote from Holloway, Yuan, and
Broadfield (2004, 13–14) should indicate the rel-
ativity of these relative brain measures.
2 E N CE P H A L I Z AT I O N
… when the EQs calculated by a particular
equation are divided by the human EQ value
(whether Jerison’s, Martin’s, Holloway and
Post’s 1982, etc.), the results can be expressed
as a percentage of human value. Rank order
correlation of these results provides a correlation
of at least 0.9, indicating a close but not perfect
agreement. Thus the rank of EQs for some ani-
mals changes depending on the database used. We
prefer the homocentric equation simply because
it gives in percent an immediate EQ of the aver-
age modern human value. However, we wish to
make the point here that EQs do not evolve. EQ
values may increase or decrease for an animal
line depending on the equations used, but what is
changing are brain/body weight variables, which
may or may not be under selection pressures at
any given time.
The application of these equations to particular
fossil hominids requires an accurate estimation
of body weights, and in a given living population
these vary considerably. For example, based on a
data set of 48 chimpanzee brain and body weights,
the EQs (using Holloway and Post’s Homocentric
equation) varied between 29% and 57% of the
modern human value of 100%, while for 23
gorillas, the EQs varied between 17% and 37%.
Using the Danish brain weight sample discussed
in Holloway (1980), and culling out extreme body
weights, adult human EQs varied between 70%
and 145%, the average and modal value being
100%. Consequently we should expect some of
our early hominids to overlap with chimpanzee
EQ values.
As one should expect, accurate EQs from the
fossil record will depend on accurate assessments
of both brain and body size, the latter not being
directly measured but inferred from modern
data from various species of primates, partic-
ularly modern humans. It should also be clear
that EQ values within a species have little if any
significance, and are best used in comparisons
between larger taxonomic units than the species.
While true in a broad sense that there is some
relationship between EQ values and intelligence,
it is almost impossible to measure intelligence
between different animal species which have
evolved their own species-specific adaptations
through evolutionary time, which clearly involves
relative amounts of brain and body mass, as
well as different neural organizations, reflecting
different neurogenomic parameters.
Most recently, the goal of achieving more
accurate EQ formulas has been taken up by
Grabowski, Voje, and Hansen (2016) who argues
that corrections for phylogeny and taxonomic
skewing provide an equation with an exponent
of roughly 3/5, or 0.6. All of the earlier equations
discussed above were derived from common least
squares regression techniques, and Grabowski
et al.’s (2016) arguments, as well as those by
Isler et al. (2008), indicate that the quality of
data, reduced axis techniques, and phylogenetic
weighting will show statistically significant dif-
ferences in the resulting exponents, for example,
between 0.6 and 0.76. The question of whether
such correcting is necessary has been questioned
by Martin and Isler (2010). This author prefers
Martin’s approach because it adheres to the actual
database for primates (without correcting factors
based on taxonomy) and provides an exponent,
roughly 3/4, which has real biological significance
in terms of metabolic relationships between the
brain, ontogeny, and maternal demands (mater-
nal energy hypothesis (Isler et al. 2008); see also
Isler and van Schaik (2006) in support of the
maternal hypothesis).
SEE ALSO: Brain evolution (primate); Brain
growth; Cognition (primate); Energetics;
Functional morphology, skull
REFERENCES
Grabowski, M., K. L. Voje, and T. F. Hansen. 2016. “Evo-
lutionary Modeling and Correcting for Observation
Error Support a 3/5 Brain–Body Allometry for Pri-
mates.” Journal of Human Evolution 94: 106–16.
Holloway, Ralph L. 1980. “Within-Species Brain–Body
Weight Variability: A Reexamination of the Danish
Data and Other Primate Species.” American Journal
of Physical Anthropology 53 (1): 109–21.
Holloway, R. L., and D. Post. 1982. “The Relativity of
Relative Brain Measures and Hominid Evolution.” In
Primate Brain Evolution, edited by E. Armstrong and
D. Falk, 57–76. New York: Plenum.
Holloway, R. L., M. S. Yuan, and D. C. Broadfield. 2004.
“Brain Endocasts: Paleoneurological Evidence, Vol-
ume 3.” In Human Fossil Record, edited by J. Schwartz
and I. Tattersall. New York: John Wiley & Sons.
 E N CE P H A L I Z AT I O N
Isler, K., and C. P. van Schaik. 2006. “Metabolic Costs of
Brain Size.” Biology Letters 2: 557–60.
Isler, K., Kirk E. Christopher, J. M. Miller, G. A.
Albrecht, B. R. Gelvin, and R. D. Martin. 2008. “En-
docranial Volumes of Primate Species: Scaling Anal-
yses Using a Comprehensive and Reliable Data Set.”
Journal of Human Evolution 55: 967–78.
Jerison, H. J. 1973. Evolution of the Brain and Intelli-
gence. New York: Academic Press.
Kleiber, M. 1947. “Body Size and Metabolic Rate.” Phys-
iological Reviews 27: 5411–541.
Martin, R. D. 1981. “Relative Brain Size and Basal
Metabolic Rate in Terrestrial Vertebrates.” Nature
293: 57–60.
Martin, R. D., and K. Isler. 2010. “The Maternal Energy
Hypothesis of Brain Evolution: An Update.” In The
Human Brain Evolving: Paleoneurological Studies in
Honor of Ralph L. Holloway, edited by D. Broadfield,
M. Yuan, K. Schick, and N. Toth, 15–35. Blooming-
ton, IN: Stone Age Institute Press.
Norvig, Peter. 2003. Artificial Intelligence: A Mod-
ern Approach. Upper Saddle River, NJ: Prentice
Hall/Pearson Education.
Snell, O. 1891. “Das Gewicht des Gehirnes und des
Hirnmantels der Saugetiere in Beziehung zu deren
geistigen Fahigkeiten Sitzungsberichte.” Gesellschaft
für Morphologie und Physiologie in München 7:
90–94.
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Wikipedia contributors. 2016. Encephalization Quo-
tient. Accessed October 2, 2017. https://en.wikipedia.
org/w/index.php?title=Encephalization_quotient&
oldid=801948431.
FURTHER READING
Boddy, A. M., M. R. McGowen, C. C. Sherwood, L. I.
Grossman, M. Goodman, and D. E. Wildman. 2012.
“Comparative Analysis of Encephalization in Mam-
mals Reveals Relaxed Constraints on Anthropoid
Primate and Cetacean Brain Scaling.” Journal of Evo-
lutionary Biology 25: 981–94.
Dubois, E. 1897. “Ueber die Abhangigkeit des
Hirngewichtes von der Korpergrosse bei den
Saugetieren.” Archiv für Anthropologie 25: 1–28.
Martin, R. D. 1996. “Scaling of the Mammalian Brain:
The Maternal Energy Hypothesis.” News in Physio-
logical Sciences 11: 149–56.
Stephan, H., H. D. Frahm, and G. Baron. 1981. “New
and Revised Data on Volumes of Brain Structures in
Insectivores and Primates.” Folia Primatologica 35:
1–29.