illumination.
Therefore, measurements ANTHROPOLOGY
should be taken before aging (open circle
in bottom right figure panel) and after ag-
ing, including a resting period in the dark
(solid circles in the bottom right figure
panel) to determine the “reversible loss.”
Devices with different starting perfor-
mances vary largely: A low-efficiency de-
vice may be considerably less stable than
a high-efficiency counterpart (see red and
black lines in the bottom right figure panel).
Hence, PSC aging routines need to disclose
all data with absolute rather than relative
(or “normalized”) values and ideally pro-
vide repeated measurements to ensure the
validity of the data. Bare perovskite layers
are very sensitive to extrinsic degradation,
e.g., ambient moisture or oxygen (9). These
can be remedied with encapsulation or,
alternatively, an inert nitrogen atmosphere
resembling “perfect encapsulation” (10, 11),
but these procedures and conditions must
be stated as well. The outlined aging routine
is only a starting point and may well change
as the understanding of aging mechanisms
in PSCs is improved. However, it is along
these parameters that a calibration curve
for accelerated aging can be developed to
start the process of establishing a standard.
The power outputs from PSCs are suffi-
ciently high to make them serious contend-
ers for sustainable energy sources. In the
coming years, PSCs will undergo relentless
scrutiny for one of the most important top-
ics in terms of market viability: long-term
stability. Unfortunately, characterizing PSCs
is challenging because of ion mobility. These
issues already cast serious doubt on the reli-
ability of efficiency measurements from the
early period of perovskite research. A critical
self-reflection on aging standards is needed
to establish a basis for norms and common
procedures with the chief aim of providing
a calibration curve that fully captures the
long-term stability of PSCs. Aging routines
that account for the hour-long settling time
of mobile ions, as outlined here, can then al-
low for the development of accelerated aging
routines to guide industrial development. j
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1. National Renewable Energy Laboratory, Best research-cell
efficiencies; www.nrel.gov/pv/assets/images/efficiency-
chart.png.
2. S. S. Shin et al., Science 356, 167 (2017).
3. M. Saliba et al., Science 354, 206 (2016).
4. M. A. Green, A. Ho-Baillie, ACS Energy Lett. 2, 822 (2017).
5. H. J. Snaith et al., J. Phys. Chem. Lett. 5, 1511 (2014).
6. E. L. Unger et al., Energy Environ. Sci. 7, 3690 (2014).
7. Y. G. Rong, L. F. Liu, A. Y. Mei, X. Li, H. W. Han, Adv. Energy
Mater. 5, 1501066 (2015).
8. K. Domanski et al., Energy Environ. Sci. 10, 604 (2017).
9. J. W. Lee et al., Adv. Energy Mater. 5, 1501310 (2015).
10. F. Matteocci et al., Nano Energy 30, 162 (2016).
11. K. A. Bush et al., Adv. Mater. 28, 3937 (2016).
10.1126/science.aar5684
SCIENCE sciencemag.org
When did modern
humans leave Africa?
A ~180,000-year-old fossil from Israel provides evidence
for early forays of Homo sapiens into western Asia
By Chris Stringer and
Julia Galway-Witham
T
he skeletal features of our species,
Homo sapiens, include a globular
braincase, brow ridges that are divided
into central and side portions, a flat
and retracted midface, a chin on the
lower jaw, and a narrow pelvis. Fossils
showing many of these characteristics have
been excavated from the Ethiopian sites of
Omo Kibish and Herto, dated at ~195,000
and ~160,000 years ago, respectively (1). Pos-
sible more primitive members of the species
are known from Jebel Irhoud (Morocco) and
Florisbad (South Africa), dated at ~315,000
and ~259,000 years ago, respectively (1). Yet,
the oldest known H. sapiens fossils outside of
Africa, from Skhul and Qafzeh in Israel, have
been dated to just 90,000 to 120,000 years
old. On page 456 of this issue, Hershkovitz
et al. (2) provide fossil evidence from Misliya
Cave, Israel, suggesting that our species had
already left Africa by ~180,000 years ago (see
the figure).
Misliya Cave lies on Mount Carmel, Israel,
and has been excavated over the last century.
It shows successive periods of hominin oc-
cupation with Lower and Middle Paleolithic
tools, the use of fire, and the exploitation of
a diverse fauna. Given Misliya’s position at a
crucial crossroads between Africa and Eur-
asia, the latest discoveries have enormous po-
tential for understanding early populations
of H. sapiens in western Asia.
The newly excavated material is repre-
sented by a partial upper jaw (Misliya-1),
which includes some of the bone surround-
ing the tooth sockets, part of the cheekbone,
the roof of the mouth, the bottom of the na-
sal cavity, and the complete upper left denti-
tion. The size and shape of the specimen fall
within the known range of variation of later
H. sapiens fossils. Furthermore, Misliya-1
lacks the unique diagnostic features of Nean-
dertals and earlier hominin species.
Hershkovitz et al. note Misliya-1’s similari-
ties to the later Skhul and Qafzeh fossils, al-
Department of Earth Sciences, Natural History Museum,
London SW7 5BD, UK. Email: c.stringer@nhm.ac.uk;
j.galway-witham@nhm.ac.uk
Published by AAAS
though the size of one of the tooth cusps in
Misliya-1 is reduced relative to the other fos-
sils. Individual traits associated with the mor-
phology exhibited in Misliya-1 are sometimes
seen in other hominin taxa, but the combi-
nation of traits is characteristically that of
H. sapiens.
Stone tools excavated from the same strati-
Downloaded from http://science.sciencemag.org/ on January 25, 2018
graphic layer as that of Misliya-1 show the
use of Levallois technology, a complex tool
preparation method involving a prepared
core. This technology has also been identified
in ~190,000- to 260,000-year-old artifacts
from nearby Tabun Cave (3), but the mate-
“…the latest discoveries
have enormous potential
for understanding
early populations of H.
sapiens in western Asia.”
rial at Misliya represents the earliest known
association of this industry with modern hu-
man fossils in the region. Levallois tools as-
sociated with putative early H. sapiens fossils
have also been found at Jebel Irhoud (Mo-
rocco) (1), suggesting that the emergence of
this tool technology may be linked with the
appearance and dispersal of our species in
both Africa and western Asia.
Hershkovitz et al. dated the fossil and
archaeological specimens from Misliya us-
ing multiple methods. They used uranium-
thorium dating to determine the age of
tooth dentine samples; combined uranium
series and electron spin resonance tech-
niques to directly date enamel samples;
uranium-thorium dating to determine the
age of the sediment adhering to the upper
jaw; and thermoluminescence to measure
the age of the burned tools found close to
the fossil. Collectively, the results of these
dating methods provide an estimated age of
~177,000 to 194,000 years for Misliya-1 and
its associated artifacts.
It remains unclear where and when hu-
mans lived in western Asia in the period prior
to that of the Misliya specimen. Fragmentary
26 JANUARY 2018 • VOL 359 ISSUE 6374 389
INSIGHTS | P E R S P E C T I V E S
fossils are known from the Israeli caves of
Qesem and Zuttiyeh, the former consisting
of isolated teeth dated to ~400,000 years
old, the latter represented by a partial skull,
perhaps of similar antiquity. Both show prim-
itive traits, and some that are found in Ne-
andertals or H. sapiens (4, 5). Archaeological
evidence for occupation of the region spans
the time between these fossils and Misliya,
but imprecise dating makes it impossible to
tell whether human presence was continuous
or episodic.
Paleoclimatic reconstructions using speleo-
thems, deep sea cores, and paleoenvironmen-
tal data suggest that there were several humid
phases between 244,000 and 190,000 years
ago, one or more of which could have facili-
tated the spread of H. sapiens into the region
(6). But there were severe periods of aridity
before and after this time, meaning that the
region was probably more often a “boulevard
of broken dreams” than a stable haven for
early humans (7). Direct local evolutionary
continuity between the population repre-
sented by the Misliya fossil and later Skhul
and Qafzeh peoples thus seems unlikely.
If the Misliya population extended further
into Eurasia, encountering Neandertals, this
may have led to gene flow between these
two lineages of humans. The main phases
of genetic introgression from Neandertals
into H. sapiens are estimated to have oc-
curred between ~50,000 and 60,000 years
ago (8), but genetic analyses of Neandertal
fossils from Denisova Cave (Siberia, Russia)
and Hohlenstein-Stadel (Germany) indicate
at least one earlier phase of introgression,
from H. sapiens into Neandertals. This event
Earliest modern human migrations from Africa
Hershkovitz et al. report an age of ~180,000 years for a modern human fossil from Misliya in Israel,
suggesting that modern humans left Africa much earlier than previously thought.
Denisova
Hohlenstein-Stadel
Jebel Irhoud
Herto
Omo Kibish
Fossil age
(thousand years)
>200
>150
90–130
Florisbad
60–90
390 26 JANUARY 2018 • VOL 359 ISSUE 6374
has been estimated at ~219,000 to 460,000
years ago, suggesting that it predated the
Misliya fossil (9). The H. sapiens fossils from
Misliya, Skhul, and Qafzeh could therefore
represent relatively late excursions of our
species from Africa. During earlier humid
phases, western Asia could have attracted
more primitive members of the H. sapiens
and H. neanderthalensis lineages, provid-
ing repeated opportunities for exchanges of
genes and technologies.
Beyond western Asia, evidence for early dis-
persals of H. sapiens prior to ~120,000 years
ago is weak. Age estimates for fragmentary
Chinese fossils from Zhiren and Daoxian
Caves only range between ~80,000 and
113,000 years ago (10). The Misliya find is im-
portant in establishing an earlier dispersal of
H. sapiens from Africa into western Asia. It
also highlights how little we know about the
early occupants of the region, nearly a cen-
tury after the first discoveries from the caves
of Zuttiyeh, Skhul, Qafzeh, and Tabun. j
RE FERENCES AND NOTES
1. C. B. Stringer, J. Galway-Witham, Nature 546, 212 (2017).
2. I. Hershkovitz et al., Science 359, 456 (2018).
3. N. Mercier, H. Valladas, J. Hum. Evol. 45, 401 (2003).
4. I. Hershkovitz et al., Am. J. Phys. Anthropol. 144, 575 (2011).
5. S. E. Freidline et al., J. Hum. Evol. 62, 225 (2012).
6. P. S. Breeze et al., Quat. Sci. Rev. 144, 155 (2016).
7. J. J. Shea, in Rethinking the Human Revolution, P. Mellars et
al., Eds. (McDonald Institute for Archaeological Research
Monographs, Cambridge, 2007), chap. 19.
8. R. Nielsen et al., Nature 541, 302 (2017).
9. C. Posth et al., Nat. Commun. 8, 16046 (2017).
10. M. Martinón-Torres et al., Curr. Anthropol. 58, S434 (2017).
ACKNOWLEDGME NTS
Authors are supported by the Calleva Foundation and the
Human Origins Research Fund.
10.1126/science.aas8954
Daoxian
Zhiren
Zuttiyeh
Misliya
Tabun (2) Skhul Qafzeh
Tabun (1)
Kebara
H. sapiens lineage
Neandertal 0 20
Qesem
Uncertain km
Published by AAAS
CELL BIOLOGY
Complexity in
targeting
membrane
proteins
Discovery of a new
pathway provides a role
for a conserved
membrane protein complex
Downloaded from http://science.sciencemag.org/ on January 25, 2018
By Michelle Y. Fry and
William M. Clemons Jr.
F
or a cell, making hydrophobic integral
membrane proteins (IMPs, or trans-
membrane proteins) is a complicated,
but critical, process. All proteins are
made by ribosomes in the cytosol, but
IMPs, which account for ~30% of the
proteins encoded in the eukaryotic genome,
must also be properly delivered to and in-
serted into their respective subcellular mem-
branes, a process known as targeting. This
is important because hydrophobic IMPs are
rapidly degraded in the cytoplasm to prevent
aggregation, which can lead to broad disrup-
tions in cellular homeostasis. Because of the
number and diversity of IMPs, identifying
and targeting them relies on pathways that
often overlap in function. The information
for targeting, typically stored in hydropho-
bic a-helical transmembrane domain (TMD)
signals in the IMPs, is recognized by factors
that then ferry the IMP client to the destined
lipid bilayer. On page 470 of this issue, Guna
et al. (1) demonstrate that, for an essential
class of IMPs, the current picture is incom-
plete. They identify a new targeting pathway
for these IMPs, contributing to the elucida-
tion of how the cell synthesizes this impor-
tant class of proteins.
IMP targeting is dominated by the secre-
tory (SEC) pathway, with the signal-recogni-
tion particle (SRP) being the central targeting
factor. For most IMPs, the SRP binds the N-
terminal signal as it emerges from a ribo-
some. The subsequent nascent protein–SRP
complex is delivered to the endoplasmic re-
GRAPHIC: J. YOU/SCIENCE
ticulum (ER) membrane for cotranslational
insertion via the SEC translocon, an ER-
Division of Chemistry and Chemical Engineering, California
Institute of Technology, Pasadena, CA 91125, USA.
Email: clemons@caltech.edu
sciencemag.org SCIENCE
When did modern humans leave Africa?
Chris Stringer and Julia Galway-Witham

Science 359 (6374), 389-390.

DOI: 10.1126/science.aas8954

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