Current Biology
Magazine
there is business — they are different
and should be fundamentally driven
by different goals: one, the pure
and unadulterated desire for greater
knowledge and the other, monetary
gain. Branding science puts focus on
making your research appealing, which
is extremely limiting, and — dare I
say? — corrupts the scientific process.
There is a lot of fundamental research
that needs to be conducted that is not
‘sexy’. Such ‘science branding’ has
not yet affected the Chinese Academy
of Sciences and for that I’m grateful.
Do you believe there is a need for
more crosstalk between biological
disciplines? There actually is a lot of it
going on, at least in paleontology, but
more is always better! For example,
I work with several labs that study
the evolution of development, but I
also collaborate with biomechanical
engineers. I love collaborating — there
is no way individuals can be experts
in every field, and by working with
other disciplines you not only expedite
the progress of your research, but
you also often gain new insights and
perspectives that often lead to new
ideas.
What’s your view on social media
and science? For example, the role of
science blogs in critiquing published
papers? Those who can, publish. Those
who can’t, blog. I understand that blogs
can be useful in affording the general
public insights into current science, but
it often seems those who criticize or
spend large amounts of time blogging
are also those who don’t generate
much publications themselves. If there
were any valid criticisms to be made,
the correct venue for these comments
would be in a similar, peer-reviewed and
citable published form. The internet is
unchecked and the public often forgets
that. They forget or are unaware that a
published paper passed rigorous review
by experts, which carries more validity
than the opinion of some disgruntled
scientist or amateur on the internet.
Thus, I find that criticism in social media
is damaging to science, as it is to most
aspects of our culture.
Institute of Vertebrate Paleontology and
Paleoanthropology, Chinese Academy
of Sciences ,142 Xizhimenwai Dajie,
Beijing 100044, China.
E-mail: jingmai.oconnor@gmail.com
R12 Current Biology 26, R1–R21, January 11, 2016 ©2016 Elsevier Ltd All rights reserved
Quick guide
African Pygmies
Paul Verdu
Who are the African Pygmies? African
Pygmies are not a single population. The
Central African populations designated
as Pygmies by Westerners are called
Aka, Baka, Bezan, Efe, Twa, etc. and
very often do not know one another
(Figure 1). The word ‘Pygmy’ derives
from an ancient Greek word referring to a
small size and was first used in Homer’s
Iliad to designate a mythical population
of short stature. In the 1860s, two
Western explorers, Paul du Chaillu and
Georg Schweinfurth, claimed to have
found the mythical Pygmies of Homer
when encountering relatively short-
statured populations in Central Africa. By
the end of the 19th century, more than 20
populations throughout the Congo Basin
were called ‘Pygmies’. During the 20th
century, anthropologists built a complex
system for the binary categorization
of Central African populations as
‘Pygmies’ and ‘non-Pygmies’, relying
on numerous cultural, economic,
linguistic, musicological (Figure 2) and
morphological criteria. Ethnographers
cautiously estimate Pygmy populations
to comprise at most 350,000 people
today. Rather than designating a single
human population, the blanket term
‘Pygmy’ hides in fact a wide cultural,
morphological, and ecological diversity.
Is the term ‘Pygmy’ derogatory?
The word ‘Pygmy’ is sometimes used
derogatorily by non-Pygmies. Using it
bluntly may thus be offensive, which
can easily be avoided by explicitly
detailing the underlying categorization
criteria. Surprisingly, some Pygmies
may use the term themselves. It
sometimes allows them to promote a
status of indigeneity that may benefit
them. Ethnographers often report that
non-Pygmies altogether admire, dread
and despise their Pygmy neighbours.
Is there a Pygmy language? There
are no Pygmy-specific languages, nor
traces of ancient Pygmy languages.
Today, they speak languages mainly
spoken by non-Pygmy populations.
Although ‘Pygmies’ are often opposed
to ‘Bantus’ in the scientific literature
as well as in the public, most Pygmy
populations speak Bantu languages.
Other groups speak Bantoid-non Bantu,
Sudanic, or Ubangian languages (Figure
1). Therefore, African Pygmies are one of
the most linguistically diverse groups of
populations worldwide.
Are Pygmies hunter-gatherers?
African Pygmies are often specialized
in hunting and gathering, but they also
practice agriculture. Likewise, non-
Pygmies are equatorial forest slash-and-
burn farmers, and also hunt and gather,
albeit less frequently than Pygmies.
Exchange of products between
communities is common, resulting in a
shared daily diet, which nevertheless
often differs in the relative amounts of
foraged and farmed foods consumed.
The extent to which Pygmy populations
practice agriculture is highly variable.
Some Pygmy groups, such as the Twa
from Lake Tumba in Western Congo,
specialize in lake-fishing.
Do African Pygmies have a common
ancestor? Population genetic studies
have extensively described Central
African genetic diversity and found high
levels of differentiation among Pygmy
populations, while the non-Pygmy
neighbours were more genetically
homogeneous. This echoed the high
levels of cultural heterogeneity among
Pygmies. Nevertheless, researchers
found that the observed genetic
patterns were highly consistent with
a common origin of all Pygmy groups
roughly 3,000 years ago in Western
Central Africa, and 20,000 years
ago between Western and Eastern
Pygmy groups. Finally, they dated the
ancestral divergence between Pygmy
and non-Pygmy populations to roughly
70,000 years ago, albeit with large
confidence intervals. As the last glacial
maximum, some 25,000 years ago, led
to the fragmentation of the equatorial
forest, population geneticists proposed
that this may have triggered the
geographic fragmentation of Pygmy
populations into Eastern and Western
groups. In addition, the expansion of
agricultural populations from Western
Central Africa into the Congo Basin
started roughly 4,000 years ago,
and was proposed to have possibly
triggered the fragmentation of Western
Pygmy populations. The cause for
the ancestral split between Pygmies
Current Biology

Magazine

and non-Pygmies remains largely N 0 250 500 km

unexplored, however.

Are African Pygmies isolated from

other populations? African Pygmies are
often imagined as hiding in the remote
heart of the equatorial forest. This is not
the case: anthropologists repeatedly
showed that they are not culturally Central African Republic
isolated, let alone hiding. Furthermore,
population geneticists found complex Cameroon
genetic admixture patterns among
Pygmy and non-Pygmy neighbours.
Genetic admixture was found largely Democratic Uganda
asymmetrical from the non-Pygmy Gabon Republic of
gene-pool into each Pygmy population, Congo
Rwanda
although some low levels of Pygmy to
P.R. Burundi
non-Pygmy admixture were also found. Congo
These admixture patterns are strongly
gender-biased towards a non-Pygmy Tanzania
male gene flow, while the non-Pygmy
female contribution is often very
small. Strongly discriminated Pygmy
populations show a stronger admixture
sex-bias compared to populations
more socially integrated, where male- Central African historical
Languages spoken by historical "Pygmy" groups
and female-mediated gene flows are "Pygmy" groups
relatively more even. West Centre East Central-South Central-North
Bezan Twa Asua Niger-Congo Niger-Congo
(Cameroon) (West DRC) (North-East DRC)
Cwa Bantoid Non-Bantu Adamawa-Ubangian
Are African Pygmies short? The short Kola/Gyeli
(Cameroon) (Centre DRC) Sua
(North-East DRC) Tikar Gbanzili-Sere, Ngbaka
height is deeply imprinted in Western Baka Cwa
(Cameroun (Centre-East DRC) Efe Bantu
representations of Pygmies, often Gabon-Congo) (North-East DRC)
A80 C60
Mikaya Central East
ignoring the tremendous variation that (Congo) Twa
(Uganda-Rwanda- C10 C80 Central Sudanic
exists across groups. Adult average Aka Burundi-DRC)
C10 D30 Mangbetu-Asua
(Congo-CAR)
height throughout the historical African Bongo
B30-B50-
J11-J60
Mangbetu
B60-B70
Pygmy populations ranges from 142 (Gabon-Congo)
B20 L30 Mangbutu-Efe
cm among Efe males from Eastern Koya/Kola
(Gabon-Congo) B40
Lese

Congo to 161 cm among Twa males Barimba

from western Congo, with an average
of 151 cm across 23 populations. Thus,
while some African Pygmy populations
are indeed among the shortest human
populations today, height variation
across them often exceeds that of
most other regions. Importantly, some
non-Pygmy populations are also
relatively short (e.g. the Ba.Konjo from
western Uganda measure less than
160 cm on average), but have not been
categorized as Pygmies historically and
genetically resemble other, taller, non-
Pygmy populations. Overall, there is no
discontinuity in adult height between
Pygmies and non-Pygmies. Thus,
categorizing Congo Basin individuals
as Pygmies or non-Pygmies based on
height alone is tricky.
Why are African Pygmies shorter
overall? This has been the subject
of heated debates. Environmental
(Gabon)
Figure 1. Geographical location of historical “Pygmy” groups and their languages.
Map and data adapted from Bahuchet 2012.
hypotheses proposed that the
Pygmies’ short stature could be
adaptive to improve movement in
the dense equatorial forest. Climatic
hypotheses proposed an adaption
to limiting the energetic costs of
inefficient perspiration in the hot and
humid forest environment. Nutritional
hypotheses proposed an adaptation
to limit metabolic requirements in the
equatorial forest where food is scarce.
More recently, demographic hypotheses
proposed that short stature would
indirectly result from an earlier onset
of reproduction being itself adaptive to
high mortality rates in the forest. Until
now, none of these hypotheses has
been convincingly supported, mainly
Current Biology
due to the lack of reliable demographic
and epidemiological data. Importantly,
all these hypotheses assume a priori
that Pygmy populations have evolved
towards a short stature. However,
we do not know how tall ancestral
Central African populations were, nor
do we know the evolutionary path
this phenotype took. Finally, some
anthropologists have proposed that
observed morphological variation in
Central Africa may solely stem from
phenotypic plasticity in challenging
environments without being the result of
adaptive evolution. However, population
geneticists recently have shown that
Pygmy individuals with high levels of
non-Pygmy admixture are also often

Current Biology 26, R1–R21, January 11, 2016 ©2016 Elsevier Ltd All rights reserved R13

Figure 2. Pygmy instruments.
Musical instruments (flute orchestra and drum) of the Nsua “Pygmy” population from Western
Uganda (Bundibugyo district). (Photo: Paul Verdu)
taller. Recent genome-wide association
studies identified possible signatures
of polygenic differential adaptation
between Pygmies and non-Pygmies,
targeting in particular genomic regions
linked to immunity and growth, hence
possibly involved in height differences
determination.
How does the future look for
Pygmies? African Pygmy populations
are far from collapsing. Thus, like any
other human population, they will
keep on evolving genetically, randomly
acquiring new mutations and losing
others due to natural selection and
the lottery of reproduction. Historical
admixture between Pygmies and
non-Pygmies is highly variable across
groups. While we can predict that
admixture may also occur in the
future, it is hard to predict whether
it will intensify or diminish in each
group. Finally, the various Pygmy
populations have experienced
massive cultural changes and
erosions since ethnographers first
started to systematically investigate
their societies in the 1950s. This was
due, directly and indirectly, to major
local and regional changes of the
social, economic, political, and geo-
strategical contexts. Thus, African
Pygmy cultures are already very
different from those of the previous
R14 Current Biology 26, R1–R21, January 11, 2016 ©2016 Elsevier Ltd All rights reserved
generations, and it is likely that they
will continue to change.
Where can I find out more?
Bahuchet, S. (2012). Changing language, remaining
pygmy. Hum. Biol. 84, 11–43.
Hewlett, B.S. (2014). Hunter-gatherers of the Congo
Basin: Cultures, Histories and Biology of African
Pygmies (New Brunswick: Transactions Publishers).
Patin, E., Laval, G., Barreiro, L.B., Salas, A., Semino,
O., Santachiara-Benerecetti, S., Kidd, K.K., Kidd,
J.R., Van der Veen, L., Hombert, J.M.,
et al. (2009). Inferring the demographic history
of African farmers and pygmy hunter-gatherers
using a multilocus resequencing data set. PLoS
Genet. 5, e1000448.
Patin, E., Siddle, K.J., Laval, G., Quach, H.,
Harmant, C., Becker, N., Froment, A., Regnault,
B., Lemee, L., Gravel, S., et al. (2014). The
impact of agricultural emergence on the genetic
history of African rainforest hunter-gatherers and
agriculturalists. Nat. Commun. 5, 3163.
Perry, G.H., Foll, M., Grenier, J.C., Patin, E., Nedelec,
Y., Pacis, A., Barakatt, M., Gravel, S., Zhou, X.,
Nsobya, S.L., et al. (2014). Adaptive, convergent
origins of the pygmy phenotype in African
rainforest hunter-gatherers. Proc. Natl. Acad.
Sci. USA 111, E3596–E3603.
Verdu, P., Austerlitz, F., Estoup, A., Vitalis, R., Georges,
M., Thery, S., Froment, A., Le Bomin, S., Gessain,
A., Hombert, J.M., et al. (2009). Origins and
genetic diversity of Pygmy hunter-gatherers from
Western Central Africa. Curr. Biol. 19, 312–318.
Verdu, P., Becker, N.S., Froment, A., Georges, M.,
Grugni, V., Quintana-Murci, L., Hombert, J.M.,
Van der Veen, L., Le Bomin, S., Bahuchet, S.,
et al. (2013). Sociocultural behavior, sex-biased
admixture, and effective population sizes in
Central African Pygmies and non-Pygmies. Mol.
Biol. Evol. 30, 918–937.
CNRS/MNHN/University Paris Diderot/
Sorbonne Paris Cité, UMR 7206
Ecoanthropology and Ethnobiology,
Paris, France.
E-mail: verdu@mnhn.fr
Current Biology
Magazine
Primer
Antimicrobial
peptides
Ling-juan Zhang and Richard L. Gallo*
Antimicrobial peptides and proteins
(AMPs) are a diverse class of naturally
occurring molecules that are produced
as a first line of defense by all
multicellular organisms. These proteins
can have broad activity to directly kill
bacteria, yeasts, fungi, viruses and
even cancer cells. Insects and plants
primarily deploy AMPs as an antibiotic
to protect against potential pathogenic
microbes, but microbes also produce
AMPs to defend their environmental
niche. In higher eukaryotic organisms,
AMPs can also be referred to as ‘host
defense peptides’, emphasizing their
additional immunomodulatory activities.
These activities are diverse, specific to
the type of AMP, and include a variety of
cytokine and growth factor-like effects
that are relevant to normal immune
homeostasis. In some instances, the
inappropriate expression of AMPs can
also induce autoimmune diseases, thus
further highlighting the importance of
understanding these molecules and
their complex activities. This Primer
will provide an update of our current
understanding of AMPs.
History of AMPs
AMPs are evolutionarily conserved
molecules found in organisms ranging
from prokaryotes to humans. Lysozyme,
the first reported human antimicrobial
protein, was identified in 1922 from
nasal mucus by Alexander Fleming. This
observation was overshadowed when
in 1928 Fleming discovered penicillin,
and in the 1940s he, along with others,
brought the therapeutic use of penicillin
to fruition, for which he was awarded
a share of the 1945 Nobel Prize for
Medicine. Thus, the beginning of the
‘Golden Age of antibiotics’ in the 1940s
led to a loss of interest in the therapeutic
potential of natural AMPs, such as
lysozyme. In the 1960s, the rise of
multidrug-resistant microbial pathogens
awakened the interest in AMPs as host
defense molecules.
AMPs have been identified from all
species of life that have been studied