Science - December 5 2014 PDF
Science - December 5 2014 PDF
Science - December 5 2014 PDF
CONTENTS
5 D E C E M B E R 2 0 1 4 V O LU M E 3 4 6 I S S U E 6 2 1 4
NEWS
1161
FEATURE
INSIGHTS
BOOKS ET AL.
REPORT P. 1215
By W. Pizer et al.
1193 RESPONSE
By C. Banks-Leite et al.
1193 ERRATA
IN DEPTH
IN BRIEF
REVIEW
RESEARCH
REPORT P. 1212
dx.doi.org/10.1126/science.1246753
RESEARCH ARTICLES
SCIENCE sciencemag.org
Published by AAAS
1151
CONTENTS
5 D E C E M B E R 2 0 1 4 V O LU M E 3 4 6 I S S U E 6 2 1 4
1238 TRANSCRIPTION
Chromatin decondensation is sufficient
to alter nuclear organization in
embryonic stem cells P. Therizols et al.
1155 EDITORIAL
The measure of research merit
By Marcia McNutt
1198
1200 PLANT GENETICS
1258
1223 PALEOCLIMATE
REPORTS
1205 VALLEYTRONICS
Ultrafast generation of pseudo-magnetic
field for valley excitons in WSe2
monolayers J. Kim et al.
1227 OCEANOGRAPHY
Multidecadal warming of Antarctic
waters S. Schmidtko et al.
PERSPECTIVE P. 1180
ON THE COVER
An electric eel
(Electrophorus
electricus) can generate
up to 600 volts using
modified muscles that
act as batteries. This
electrical weapon
allows the eel to
remotely activate the
muscles in nearby prey to cause either
involuntary movement or involuntary
immobility, depending on the eels needs.
See page 1231 for details and links to
movies that illustrate these findings.
Photo: Kenneth Catania
1234 INFLAMMATION
Neutrophils scan for activated platelets
to initiate inflammation
V. Sreeramkumar et al.
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1153
Stella M. Hurtley(UK), Paula A. Kiberstis, Marc S. Lavine(Canada), Kristen L. Mueller, Ian S. Osborne(UK), Beverly A. Purnell,
L. Bryan Ray, Guy Riddihough, H. Jesse Smith, Jelena Stajic, Peter Stern(UK), Phillip D. Szuromi, Brad Wible, Nicholas S. Wigginton,
Laura M. Zahn ASSOCIATE EDITORS Brent Grocholski, Melissa R. McCartney, Margaret M. Moerchen, Sacha Vignieri ASSOCIATE BOOK
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Maryrose Madrid, John Wood(UK)
News
NEWS MANAGING EDITOR John Travis INTERNATIONAL EDITOR Richard Stone DEPUTY NEWS EDITORS Daniel Clery(UK), Robert Coontz,
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INFORMATION FOR AUTHORS See pages 680 and 681 of the 7 February 2014 issue or access www.sciencemag.org/about/authors
1154
sciencemag.org SCIENCE
Published by AAAS
E D I TOR IAL
Marcia McNutt
Editor-in-Chief
Science Journals
It is time to remedy a
flawed bibliometric-based
assessment for young
scientists.
Marcia McNutt
10.1126/science.aaa3796
SCIENCE sciencemag.org
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1155
NEWS
James Watson, who co-discovered the structure of DNA but has been criticized
for making racist remarks, explaining why he is auctioning his Nobel medal.
IN BRIEF
1158
sciencemag.org SCIENCE
Published by AAAS
ike eight-legged birds of paradise, male jumping spiders flash red, blue, green,
and orange patches as they circle, lunge, and wave their legs in a courtship
dance. Taking advantage of the spiders antics (and YouTube fame), researchers have raised more than $7000 from the public to study how well the spiders
see color. Vision ecologist Daniel Zurek of the University of Pittsburgh in
Pennsylvania and colleagues had found a red filter in the eyes of five Habronattus
species of jumping spiders that greatly expands the diversity of color they see. To
find out how widespread the red-filter adaptation is among the 100 Habronattus
species, the team turned to Experiment.com, a crowdfunding website for scientists. Having reached their goal last week, the team plans to head to Arizona and
southern California next spring to collect 12 to 25 species of the spiders; theyll test
the spiders eyes for color sensitivity and determine whether improved color vision
coincides with flashier displays.
http://scim.ag/EPAozone
A LT O S D E P I P E , V E N E Z U E L A
SCIENCE sciencemag.org
Published by AAAS
1159
BY THE NUMBERS
10%
75
B E T H E S DA , M A RY L A N D
1160
10
million
Years ago that a common ancestor of
humans, chimps, and gorillas evolved
a protein that could efficiently
metabolize alcohol. (Proceedings of
the National Academy of Sciences)
NEWSMAKERS
SUE
E 6214
62
214
14
5 DECEMBER 2014 VOL 346 ISSUE
Published by AAAS
The Planck spacecraft has mapped the polarization of the CMB radiation across the sky.
NEWS
IN DEP TH
BIOMEDICAL RESEARCH
CREDITS: (IMAGE) JASON EDWARDS/NATIONAL GEOGRAPHIC SOCIETY/CORBIS; (DATA) NATIONAL HOSPITAL DISCHARGE SURVEY (2005) AND NATIONAL
INPATIENT SAMPLE, U.S. AGENCY FOR HEALTHCARE RESEARCH AND QUALITY, VIA SUN, BRITISH JOURNAL OF ANAESTHESIA, 2010
million
1.5
million
SCIENCE sciencemag.org
Published by AAAS
1161
NEWS | I N D E P T H
ARCHAEOLOGY
Published by AAAS
CREDITS: (PHOTO) JESSE CASANA, UNIVERSITY OF ARKANSAS; (MAP) ASOR SYRIAN HERITAGE INITIATIVE
a massive expansion
in looting between
August 2013 and April
2014 at Dura-Europos,
a sprawling Roman-era
city on the Euphrates
in Syria. But Bronze Age cities like Ebla in
the westdamaged by a Syrian government
military garrisonand Mari, which is under
the Islamic State groups control, are not immune. There are rumors that armed groups
are undertaking the work, ASORs Branting
says about Mari. Other reports suggest that
the Islamic State group is profiting from the
business, possibly by exacting a tax as well
as by overseeing looting operations. But
Danti adds that most looting appears to be
the work of desperate Syrians attempting to
survive in a devastated economy.
Massive looting, bomb craters, and damage by military garrisons can be spotted
on satellite images, but smaller thefts, kidnapping, and violence against guards and
inspectors at sites are difficult to confirm
Damage assessment
Many of Syrias ancient sites have sustained
damage (red), as shown on a map based on
satellite data. Sites in blue appear undamaged.
Turkey
Syria
Iraq
Lebanon
Israel
Jordan
Saudi Arabia
SCIENCE sciencemag.org
Published by AAAS
1163
EPIDEMIOLOGY
1164
Sierra Leone and down in Liberia in October, he was skeptical, and he decided to find
out firsthand what was happening. He canceled his lectures and contacted the Liberian government. Im not a virologist and
Im not a clinician, but I have considerable
experience investigating messy epidemics
in poor parts of Africa, he says.
A quarter-century ago, he faced a threat
that seemed even more terrifying than Ebola. In August 1981, Rosling was working as
a district medical officer in northern Mozambique when he faced a line of 30 women
and children whose legs had become paralyzed over the past month. I had this big
neurology book, and their disease did not
exist in that book, Rosling says. A South
African submarine had been spotted in a
nearby bay a few weeks earlier. It was fully
possible that it was biological warfare.
When he came home that evening, he
told his wife to take their kids and drive
them to a safer place a few hours away. He
didnt sleep the next 48 hours. When you
face a disease that may be infectious, 98% of
your intellectual capacity becomes blocked.
You become so scared, thinking you will
die, thinking you may be crippled, thinking about your children. It took 2 weeks
Published by AAAS
NEWS | I N D E P T H
A declining epidemic
The number of reported new Ebola cases in
Liberia has dropped to about 10 per day, which
poses new challenges.
Guinea
Sierra
Leone
Ivory
Coast
Liberia
Monrovia
15
620
2150
51250
Total cases
15
620
21100
101500
5014000
SCIENCE sciencemag.org
Published by AAAS
1165
NEWS | I N D E P T H
BRAZIL
razilian
neuroscientist
Miguel
Nicolelis has won global attention for
experiments with brain-controlled
prosthesesand for masterminding
a controversial World Cup kickoff
by a paralyzed man in a robotic
exoskeleton. But back home, controversy
over the high-profile researchers recent
windfall has reached a boiling point. Last
week, Brazils largest scientific organization wrote a letter on behalf of more than
120 scientific societies questioning the
federal governments decision to invest
roughly U.S. $100 million in the Campus
of the Brain, the neuroscience research and
education initiative in rural northern Brazil spearheaded by Nicolelis.
According to Nicolelis, the funding provided by Brazils Ministry of Education
will cover operational costs over the next
4 years for the campuss health and education efforts and for its several dozen neuroscience research laboratories. The money
is not being taken out of anyone elses pot,
he stresses. These funds do not come from
annual federal research moneythis is
new money, a new appropriation.
Yet the Brazilian Society for the Advancement of Science (SBPC) argues that the nations entire scientific community should
have been given a chance to compete for
the funding. Allocating substantial resources for an ad-hoc initiative without, to
1166
sciencemag.org SCIENCE
Published by AAAS
FEATURES
UNKNOWN
SIGNIFICANCE
By Jennifer Couzin-Frankel
when they have no family history of the disease, researchers reported Thursday.
Hmmm.
Science has been my professional home
for the past 13 years, and in that time Ive
written extensively about genetic testing
and spoken with dozens of experts at the
fields cutting edge. Ive chronicled the scientific advances, the ethical quandaries,
the lives testing saves, the angst it ignites. I
had never turned the lens on my own DNA.
Abruptly, there was no escaping it. My
SCIENCE sciencemag.org
Published by AAAS
1167
NEWS | F E AT U R E S
1168
Published by AAAS
20
20
40
40
6%
2%
29%
19%
92%
71%
69%
60%
People need to
become more
comfortable with
uncertainty.
SCIENCE sciencemag.org
Published by AAAS
1169
NEWS | F E AT U R E S
Multigene panels for cancer risk are proliferating and evolving, including this one of 21 genes associated with breast, ovarian, and other cancers, shared with the author prior to her own testing.
HIGH-RISK GENES
GENE
BRCA1
BRCA2
CDH1
EPCAM
MLH1
MSH2
MSH6
PMS2
PTEN
STK11
TP53
CHEK2
PALB2
NEWER GENES
BARD1
BRIP1
FANCC
Female breast
NBN
RAD51C
RAD51D
XRCC2
Outside the United States, panels of cancer genes are largely restricted to research
settings, and investigators often dont share
information about mutations that carry
a modest or unknown risk. Theres much
debate over what to tell volunteers. We
struggle with it, says Hans Ehrencrona, a
clinical geneticist at Lund University Hospital in Sweden. Where to draw that line, no
1170
Published by AAAS
INSIGHTS
B O OKS et al .
Tiny Creatures:
The World of Microbes
Nicola Davies,
illustrated by Emily
Sutton
Candlewick, 2014.
36 pp.
Buried Sunlight
Molly Bang and
Penny Chisholm,
illustrated by
Molly Bang
The Blue Sky Press,
2014. 44 pp.
1172
Parrots over
Puerto Rico
Reviewed by Yevgeniya Nusinovich, Solomon
Nusinovich Ucko, Hadassah Nusinovich Ucko
Published by AAAS
Parrots over
Puerto Rico
Susan L. Roth and
Cindy Trumbore,
illustrated by
Susan L. Roth
Buried Sunlight
Reviewed by Brent Grocholski
SCIENCE sciencemag.org
Published by AAAS
1173
INSIGHTS | B O O K S
Millbrook Press,
2014. 48 pp.
Mission: Mars
Pascal Lee,
illustrated by
Ryan Hobson
Scholastic, 2013.
48 pp.
1174
The Case of
the Vanishing
Honeybees:
A Scientific Mystery
Published by AAAS
Plastic, Ahoy!
Investigating the
Great Pacific
Garbage Patch
Patricia Newman,
photography by
Annie Crawley
Plastic, Ahoy!
Investigating
the Great Pacific
Garbage Patch
Reviewed by Marc Lavine
When we think of large-scale waste disposal, we often think of giant, stinky landfill sites where garbage is buried. However,
the largest collection of garbage isnt on
land but rather in the Pacific Ocean. Known
as both the Pacific trash vortex and the
Great Pacific Garbage Patch, it is located
in a part of the ocean called the North Pacific Central Gyre and spans waters from
the west coast of North America to Japan.
Here, the convergence of four currents
creates a pocket of relatively calm water,
which causes large quantities of small,
primarily plastic, debris to accumulate. In
this book, author Patricia Newman follows
three young scientistsMiriam Goldstein,
Chelsea Rochman, and Darcy Taniguchi
on an ocean voyage to explore the garbage
patch as part of the Scripps Environmental
Accumulation of Plastic Expedition (SEAPLEX). Despite its size, which both is hard
to quantify and can vary through the year,
the patch cannot be seen by satellites because most of the garbage is in the form of
small particles and a large fraction may lie
on the sea floor. Scientists hoping to study
it must take to the open ocean, which also
gives them the opportunity to study the impact of the patch on the local marine life
and food chain. Newman narrates the book
from the perspective of the researchers and
crew and their daily activities, including
anecdotes about the voyage itself, as well as
the exploration of the garbage patch. She
uses these adventures to show the reader
what it means to be a scientist and to use
scientific methods.
Mission: Mars
Reviewed by Maria Cruz
SCIENCE sciencemag.org
Published by AAAS
1175
INSIGHTS | B O O K S
Plant a Pocket of
Prairie
Reviewed by Barbara Jasny
1176
The Tale of
the Dueling
Neurosurgeons
Reviewed by Valerie Thompson
In this book, New Yorker staff writer Elizabeth Kolbert presents compelling evidence
that we are in the midst of a human-caused
mass extinction. According to Kolbert, this
sixth extinction is of a similar magnitude
to previous mass extinction events, including the Cretaceous event that famously
spelled the end for the dinosaurs. The author is at her best when recounting her
personal encounters with species or ecosystems in peril. Her account of the demise of
North American bats from an introduced
sciencemag.org SCIENCE
Published by AAAS
Junk Drawer
Physics
Extreme Medicine:
How Exploration
Transformed
Medicine in the
Twentieth Century
Shocked: Adventures
in Bringing Back the
Recently Dead
Reviewed by Guy Riddihough
SCIENCE sciencemag.org
Plant a Pocket
of Prairie
Phyllis Root,
illustrated
by Betsy Bowen
University of Minnesota
Press, 2014. 40 pp.
The Sixth
Extinction:
An Unnatural
History
Elizabeth Kolbert
Henry Holt,
2014. 330 pp.
Extreme Medicine:
How Exploration
Transformed Medicine
in the Twentieth
Century
Kevin Fong
Penguin Press,
2014. 304 pp.
Shocked: Adventures
in Bringing Back
the Recently Dead
David Casarett
Current, 2014.
270 pp.
Published by AAAS
1177
INSIGHTS | B O O K S
The Usborne
Big Book of Stars
and Planets
Reviewed by Maria Cruz
Life on Earth is ubiquitous, multidimensional, and complex, and what better medium
to convey this than a pop-up book? This book
devotes a full-page spread to six of the earths
biological regions, taking the reader from
grassland savannahs to tangled rainforests,
through chilly polar regions and dry deserts,
and to fresh and saltwater environments. The
book opens with some basic concepts and
random facts about seasonality, biomes, and
the evolution of the elephants trunk. Opening the next page makes a rainforest spring
upcomplete with tapirs, macaws, and jaguars. The desert pages are covered with flaps
that show that arid biomes are not as devoid
of life as they may first appear (although why
there is a jerboa in the dung beetles burrow is not immediately clear). Subsequently,
salmon leap out of a river against a backdrop of coniferous forests, and polar animals
plunge off icebergs and bask in their blubber.
Theres even a colorful coral reef and its inhabitants. Here, flaps lift to reveal what animals can be found in the depths of the seas,
and on other pages, the flaps show those that
take over the habitats at night. The book
is also peppered with intriguing facts: For
Lift-the-Flap
Questions and
Answers About
Your Body
Reviewed by Jennifer Sills
Published by AAAS
is aimed at 10- to 14-year-olds, but the delight that comes from putting one over on
our own brains knows no ageall should
find some fun surprises in this book.
What Makes
You YOU?
Gill Arbuthnott,
illustrated by
Marc Mones
A & C Black,
2013. 64 pp.
Lift-the-Flap
Questions and
Answers About
Your Body
Katie Daynes,
illustrated by
Marie-Eve Tremblay
Reviewed by
Margaret Moerchen
Usborne,
2013. 28 pp.
Eye Benders:
The Science
of Seeing
and Believing
Cant believe your eyes? It
could be your brain thats
lying to you. In this book,
journalist Clive Gifford
leads a tour of optical illusions that make you look twice and think
one more time. Gifford begins by laying out
the mechanics of how our eyes and nervous
system collect and process information. Using colorful and clever images that elicit
specific illusions, he goes on to explain
the shortcuts the brain usually takes that
lead it down the wrong path. We learn that
optical illusions exploit everything from
the brains propensity to fill in the gaps
when it receives inadequate sensory information, to its methods for estimating size
and distance, to the curious way it recognizes edges. At times, the brains strength at
a given skillsuch as facial recognitionis
what brings about its momentary lapses.
We see faces on Mars where only a broad
mountain exists, for example, thanks to a
phenomenon called pareidolia. On another
page, two upside-down photos of human
faces appear to be identical, but on upending the book, its impossible not to see that
the features in one photo have been rightside up all along, a phenomenon known as
the Thatcherized illusion. The slim volume
The Usborne
Big Book of Stars
and Planets
Emily Bone,
illustrated by
Fabiano Fiorin
Usborne,
2013. 14 pp.
SCIENCE sciencemag.org
Weve Got
Your Number
Mukul Patel,
illustrated by
Supriya Sahai
Kingfisher,
2013. 96 pp.
Published by AAAS
1179
PERSPECTIVES
OCEANOGRAPHY
obody lives permanently in Antarctica. At first glance, studies of Antarctic climate might thus seem like
a curiosity without obvious societal
implications. Yet, if the entire West
Antarctic Ice Sheet were to melt,
global sea level would rise by 4.8 m, with
major effects on coastal populations. Two
studies published earlier this year offered
convincing evidence that the West Antarctic
Ice Sheet is indeed melting irrevocably (1, 2).
What are the processes behind this melting?
On page 1227 of this issue, Schmidtko et al.
(3) show that the water flowing under the ice
shelves has warmed in recent decades and
that upwelling winds at the continental shelf
break may contribute to this process.
Because sea ice floats on water, melting or
freezing of sea ice has no major effect on sea
level. Ice sheets are a different story, however, because they sit on land and thus keep
water frozen and out of the ocean. Ice shelves
are the oceanic extensions of ice sheets (see
the photo). Like sea ice, they float. However,
they can be 200 to 1000 m thick (4) and can
serve as buttresses, retaining the ice shelves
on land (5, 6).
Ice shelves may disintegrate by calving
icebergs off the leading edge or by melting through warming from above or below.
Scripps Institution of Oceanography, University of California San
Diego, La Jolla, CA 92093-0230, USA. E-mail [email protected]
1180
Calved ice
Sea ice
Circumpolar deep water
Basal cavity
Melting
Continental shelf
Shelf break
Land
Melting from below. The Antarctic ice shelves are the ocean extensions of the Antarctic ice sheets. Ice shelves
buttress the ice sheets onto the land. Comparatively warm water that circulates under the ice shelves can lead to basal
melting of the underside of the ice shelves, ultimately breaking up the ice shelves. Schmidtko et al. (3) show that wind
patterns at the continental-shelf break influence the properties of water on the continental shelf and in the basal cavity.
sciencemag.org SCIENCE
Published by AAAS
By Sarah Gille
INSIGHTS
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
DEVELOPMENTAL BIOLOGY
nsects use the innate immunity signaling pathways Toll and IMD to fight infections by foreign pathogens, bacteria, or
fungi (1). These pathways induce the expression of powerful antimicrobial- antifungal peptides (AMPs) that are released
from the fat body cells. On page 1199 of this
issue, Meyer et al. (2) report that fruit flies
make use of components of both pathways
to build another pathway that removes its
SCIENCE sciencemag.org
Published by AAAS
1181
INSIGHTS | P E R S P E C T I V E S
Pro-Spz
Spz
TRRs
Tube
PGRP-LCs
MyD88
IMD
dFADD
DREDD
Pelle
Loser cell is
engulfed by
winner cell
dTAK1-dTAB2
P P P
Cactus
KennyIRD5
Dif/Dl
P
Relish
rpr
Apoptosis
hid
and rpr, and death of the loser cells. They are subsequently engulfed by the neighboring
winner cells, thereby restoring a healthy tissue. PGRP-LC, peptidoglycan recognition
protein LC; IMD, immune deficiency; TAK1, transforming growth factor betaactivated
kinase 1; DREDD, death-related ced-3/Nedd2-like caspase; FADD, FAS-associated death
domain; TAB2, TAKI-associated binding protein 2; IRD5, immune-response deficient 5.
sciencemag.org SCIENCE
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CHEMICAL KINETICS
Energy, E
SCIENCE sciencemag.org
Published by AAAS
1183
INSIGHTS | P E R S P E C T I V E S
1184
GEOLOGY
sciencemag.org SCIENCE
Published by AAAS
Depth: 150 km
those of the deep mantle, and magmas from beneath ridges can erupt
at lower temperatures than magmas
extracted from the asthenosphere be2
30
30
neath the middle of large plates.
1
Whereas Earths interior has traditionally
been viewed as a series of
0
0
0
uniform layers, seismic tomography
has enabled mapping of complex
1
30
30
three-dimensional structures. French
et al. (3) have resolved horizontally
2
elongated, lowwave speed struc60
60
tures at depths of 200 to 350 km ex3
tending from the East Pacific Rise to
4
the middle of the Pacific plate; they
found
similar features beneath other
2
B
Depth: 2700 km
oceans. These lowwave speed structures occur below the average depth
60
60
of the asthenosphere and represent
1
the interaction of broad, passive up30
30
wellings with the base of the asthenosphere. Most hot spots occur above
these lowwave speed regions (see
0
0
0
the figure, panel A), and the correlation between hot spots and the edges
30
30
of the lowshear velocity large struc1
tures at the core (see the figure, panel
B)a recent observation used to sup60
60
port the existence of deep mantle
plumesis no better than the correla2
tion with the lowwave speed regions
beneath the asthenosphere (15).
Top or bottom? Seismic anomalies in the asthenosphere (A) and at the base of the mantle, just above the core (B); the anomalies
Plate tectonics and midplate volare relative to a one-dimensional reference model. The correlation between hot spots (triangles) and the edges of the slow (red)
canoes are the natural results of
anomalies in the lower mantle has been used to argue that hot spots originate from plumes at these depths. However, hot spots
processes in the asthenosphere and
correlate as well if not better with slowwave speed anomalies in the asthenosphere, the region where the lavas originate.
lithosphere of the cooling Earth.
Given that the hottest part of the mantle lies
of lowwave speed, low-viscosity anisotropic
sphere may be as much as 200C warmer
directly beneath the lithosphere (2, 4, 13),
material that is consistent with the observed
than beneath mid-ocean ridges (4) and even
there is no need for heat from the core to be
properties of the asthenosphere (12). Interachotter than the core-mantle boundary, when
carried up in narrow plumes (4). As a result,
tion of seismic waves with this material percorrected for adiabatic cooling (the coolthe asthenosphere is by far the largest, most
turbs the material and further reduces the
ing that results from transporting material
accessible, and most plausible source for hot
seismic wave speed (12). Density measurefrom high to low pressure). This is imporspot magmas. Although many authors have
ments of volatile-containing alkali basalts
tant because the asthenosphere is at or near
ruled out this obvious source on the basis of
show that these melts are neutrally buoyant
its melting point, and a large temperature
geochemical assumptions, it is not ruled out
at the pressure and temperature conditions
increase would thus produce a substantial
by any data or theory.
of the asthenosphere under continents (10),
amount of melt. Most researchers have asadding to the evidence that the asthenosumed that the hottest material should be
RE FE RE N CES
sphere is a zone of partial melt. Because both
below ridges, leading them to invoke plumes
1. J. Barrell, J. Geol. 22, 28 (1914).
seismic and mechanical properties are used
to explain 200C lateral temperature varia2. N. Zhang et al., J. Geophys. Res. 115, B06401 (2010).
3. S. French et al., Science 342, 227 (2013).
to describe the asthenosphere, the bottom
tions. Yet in the mantle, where a substantial
4. D. L. Anderson, J. H. Natland, Proc. Natl. Acad. Sci. U.S.A.
boundary remains poorly defined.
fraction of the heat is generated by the decay
111, E4298 (2014).
There is also evidence that the asthenoof radiogenic elements, such lateral varia5. B. H. Hager, R. J. OConnell, Tectonophysics 50, 111 (1978).
6. T. Hoink et al., Geophys. J. Int. 191, 30 (2012).
sphere is hotter than the mantle below it
tions in temperature can arise without the
7. J. W. Crowley, R. J. OConnell, Geophys. J. Int. 188, 61
(2, 4, 13). Average temperature profiles from
need to deliver heat from the deep mantle
(2012).
8.
H. Samuel, S. D. King, Nat. Geosci. 7, 602 (2014).
three-dimensional global mantle convection
via a plume (4).
9. D. L. Anderson, J. Petrol. 52, 1547 (2011).
simulations show a local thermal maximum
Further information on mantle tempera10. C. Crpisson et al., Earth Planet. Sci. Lett. 393, 105 (2014).
beneath the lithosphere because of the inture comes from absolute seismic wave
11. R. Dasgupta et al., Nature 493, 211 (2013).
12. L. Li, D. J. Weidner, Phys. Earth Planet. Inter. 222, 1 (2013).
sulating properties of the tectonic plates (4,
speeds, which can be coupled with a ther13. W. B. Moore, J. Geophys. Res. 113, B11407 (2008).
13). This increase in temperature can reduce
modynamic description of mantle minerals
14. J. Ritsema et al., Earth Planet. Sci. Lett. 277, 244 (2009).
the mechanical strength, lower the seismic
to estimate temperature. This approach con15. B. R Julian et al., paper to be presented at the American
Geophysical Union fall meeting, 15 to 19 December 2014;
wave speed, and eliminate the need to bring
firms that the asthenosphere is anomalously
see https://agu.confex.com/agu/fm14/
heat directly from Earths core to the asthehot, whereas the deep mantle is anomalously
webprogrampreliminary/Paper20489.html.
nosphere to explain volcanic activity at hot
cold (14). Thus, the highest temperatures
10.1126/science.1261831
spots. The hottest regions in the asthenoin the asthenosphere can be greater than
60
60
SCIENCE sciencemag.org
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1185
INSIGHTS | P E R S P E C T I V E S
SURFACE SCIENCE
1186
Published by AAAS
IILLUSTRATION: V. ALTOUNIAN/SCIENCE
REF ERENC ES
HUMAN GENOMICS
SCIENCE sciencemag.org
Published by AAAS
1187
INSIGHTS | P E R S P E C T I V E S
1188
sciencemag.org SCIENCE
Published by AAAS
ILLUSTRATION: V. ALTOUNIAN/SCIENCE
ENVIRONMENTAL ECONOMICS
15
10
0
Costs
2013 SCC
2008 SCC
Climate-only beneft
estimates based on diferent
government SCC values
Benefits of regulations vary. Estimated costs and
climate change benefits of emission reductions in 2020
from proposed U.S. power plant regulations using 2008
(3) and 2013 (2) government SCC estimates. Estimates
from table 18 in (2) using a 3% discount rate averaged
over state and regional approaches. SCC estimate from
table V-3 in (3), rising 2.4% per year to $8.67 in 2020,
multiplied by avoided emissions estimates averaged
over state and regional approaches from table 10 in (2),
and inflation adjusted using the implicit GDP price index
from the U.S. Bureau of Economic Analysis.
SCIENCE sciencemag.org
an
SCC for government CBA requires specific choices, beginning with the selection
of which IAMs to include. Models vary in
terms of breadth of use, degree of public access and available peer review, and incorporation of latest scientific results. New IAMs
may emerge. How should a government select among models? Should selection evolve
over time? Should models be weighted? If
so, how?
Next, one must choose what, if any, assumptions to harmonize across models.
Such assumptions may be important for
consistency between the SCC and other
elements of a government CBA, to reflect
important uncertainties, or to address possibly outdated assumptions.
This harmonization requires more tough
choices. For example, the SCC will measure
incremental policy benefits relative to a
baseline or range of baselines, which must
be explicitly selected. One must decide
whether emissions are forecast on the basis of an ambitious climate policy (such as
the scenario in which polluters are already
forced to pay the estimated SCC), a scenario
where only policies already on the books
remain in place, or something in between.
There are also credible differences on
analytic and ethical grounds regarding the
appropriate discount rate. Previous government guidance for CBA suggested discount
rates of 3 and 7% for most projects, with
possibly lower rates for phenomena (like
climate change) with important intergenerational effects (7). Such differences have
enormous implications; federal SCC estimates tripled as the discount rate changes
from 5 to 3% (1). For practical CBA, it is important to have distinct SCC estimates for
different discount rates that can be paired
with cost estimates based on a particular
discount rate(s).
Each IAM will have its own internal discount rate determined by model parameters and socioeconomic forecasts. Low
discount rates typically follow from low
economic growth (8), and economic growth
is tied to climate impacts. Given this connection, how problematic is it to impose a
discount rate in the SCC that is different
from the rate used within the IAM itself ?
5 DECEMBER 2014 VOL 346 ISSUE 6214
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1189
INSIGHTS | P E R S P E C T I V E S
U.S. government practice is vague regarding when and how a process of reviewing
and updating the SCC might occur, which
makes it difficult for stakeholders and re1
1190
sciencemag.org SCIENCE
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ES SAY
SCIENCE & SCILIFELAB PRIZE ESSAY
mTORC1
(inactive)
GATOR2
GATOR2
Growth factors
energy oxygen
GATOR1
Amino acids
GATOR1
GAP activity
v-ATPase
GDP
RagA
Ragulator
RagC
GTP
FLCN-FNIP
GEF activity
GTP
Rheb
v-ATPase
GTP
RagA
Ragulator
RagC
GDP
mTORC1
(active)
FLCN-FNIP
GTP
Rheb
GAP activity
Lysosomal lumen
The pathway of amino acid sensing by mTORC1. (Left) During amino acid deprivation, Ragulator is found as an inactive complex with the v-ATPase. GATOR1 GAP activity toward
RagA keeps this GTPase in an inactive GDP-bound state that cannot recruit mTORC1. (Right) After amino acid stimulation, GATOR1 is inhibited by GATOR2, and Ragulator and the
v-ATPase undergo a conformational change releasing the guanine nucleotide exchange factor (GEF) activity of Ragulator toward RagA. At the same time, the FLCN-FNIP complex
activates RagC, by promoting its GTP hydrolysis. The now-active Rag heterodimer (RagA-GTP, RagC-GDP) recruits mTORC1 to the lysosomal surface, where it interacts with and is
activated by the small GTPase Rheb, which is itself controlled by growth factor, energy, and oxygen levels (1).
SCIENCE sciencemag.org
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1191
Category Winners
Genomics and Proteomics: Dan Dominissini for his essay, Roadmap to the
epitranscriptome. Dr. Dominissini received his Bachelor of Medical Science degree from Tel-Aviv University, Israel, in 2007. He went on to study
RNA posttranscriptional modifications for his Ph.D., focusing on adenosine
deamination and methylation, with Gideon Rechavi at Tel-Aviv University. He
is currently a Human Frontier Science Program postdoctoral fellow in the
laboratory of Chuan He at the University of Chicago, where he develops novel
chemistries for the study of nucleic acid modifications.
Translational Medicine: Simon C. Johnson for his essay, A target for pharmacological intervention in an untreatable human disease. Dr. Johnson is an
American Federation for Aging Research Fellow at the Albert Einstein College
of Medicine in New York. He earned his Bachelor of Science degree at Oregon
State University and received his Ph.D. from the University of Washington.
He was a 2009 Howard Hughes Medical Institute EXROP scholar and was
previously supported by the Nathan Shock Center Genetic Approaches to
Aging predoctoral and Mechanisms of Cardiovascular Diseases postdoctoral
competitive training grants. His work is centered on characterizing the role of
genetic variation in insulin/IGF-1/mTOR signaling genes on human longevity.
Environment: Chelsea Wood for the essay, Environmental change and the
ecology of infectious disease. Dr. Wood received her Bachelor of Arts degree
from Dartmouth College and her Ph.D. from Stanford University. She did
postdoctoral research in Pieter Johnsons lab at the University of Colorado
at Boulder, and is currently a Fellow in the Michigan Society of Fellows and an
Assistant Professor in the Department of Ecology and Evolutionary Biology at
the University of Michigan. She is interested in how parasites and pathogens
respond to human impacts on the environment.
For the full text of all winning essays and further information, see http://scim.ag/SciLifeLab.
1192
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
ACK N OW LE D G M E N TS
I am grateful to my Ph.D. adviser D. M. Sabatini for his mentorship. I would also like to thank all the members of the Sabatini
lab past and present and especially L. Chantranupong, Y.
Sancak, L. Schweitzer, and R. Zoncu.
10.1126/science.aaa1808
sciencemag.org SCIENCE
Published by AAAS
PHOTO CREDITS: (TOP) SCRIPPS RESEARCH INSTITUTE; (SECOND AND LAST) PERSONAL PHOTOS; (THIRD) CITY HEADSHOTS
INSIGHTS | E S S AY
LET TERS
Edited by Jennifer Sills
Conserving Brazils
Atlantic forests
Response
BUCKLEY AND PEGAS claim that it would
SCIENCE sciencemag.org
ERRATA
Erratum for the Report: Washing away your
sins: Threatened morality and physical
cleansing by C.-B. Zhong and K. Liljenquist,
Science 346, aaa2510 (2014). Published online
21 November 2014; 10.1126/science.aaa2510
5 DECEMBER 2014 VOL 346 ISSUE 6214
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1193
INSIGHTS | L E T T E R S
n his Editorial Chinas private universities (24 October, p. 401), H. Jin expresses hope that the Chinese government
will soon treat public and private universities equally, giving private institutions the funding and opportunities they
need to flourish. Readers analyzed the trade-offs involved in such a decision in the online comments section. Excerpts
from their comments are below. Read the full comments, and add your own, at http://comments.sciencemag.org/
content/10.1126/science.1262035.
Science &
Diplomacy
provides an open
access forum for
rigorous thought,
analysis, and
insight to serve
stakeholders who
develop, implement,
or teach all aspects
of science and
diplomacy. Learn
more about the
latest ideas in
science diplomacy and receive regular updates by following
@SciDip on Twitter, liking the quarterlys page on Facebook
(www.facebook.com/sciencediplomacy), and registering for
free at www.sciencediplomacy.org/user/register.
www.ScienceDiplomacy.org
RESEARCH
IN S CIENCE JOUR NAL S
ANIMAL PHYSIOLOGY
PHOTOS: (TOP TO BOTTOM) TENBOER ET AL.; WORLD TRAVEL COLLECTION/ALAMY; SREERAMKUMAR ET AL.
INFLAMMATION
A two-cell collaboration
for inflammation
Immune cells called neutrophils
are first responders to infection. Neutrophils move within
and through blood vessels to
get to sites of infection quickly.
Intravascular
neutrophil (green)
and platelet (red)
aggregates
CHEMICAL KINETICS
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1195
OCEANOGRAPHY
RESEARCH | I N S C I E N C E J OU R N A L S
HUMAN CAPITAL
An ounce of prevention
is better than violence
Children and adolescents spend
a large fraction of their lives in
schools. Many discussions about
improving adult behavior focus
on improving cognitive skills.
Nevertheless, retrospective analyses also highlight the importance
of noncognitive skills. Heller
reports on the positive effects of
a Chicago program that offered
disadvantaged youths 25 hours
per week of summer employment. Youths enrolled in this
program committed fewer violent
crimes for at least a year after the
summer job ended. GJC
Science, this issue p. 1219
VALLEYTRONICS
TRANSCRIPTION
IN OTHER JOURNALS
A shipworm
removed from its
wooden burrow
PLANT GENETICS
Dominance cascades in
self-incompatibility
Plants often cannot use their own
pollen to set seed. This is known
as self-incompatibility. Although
some of the underlying genetics
and mechanisms of self-incompatibility are understood, the
evolution and maintenance of the
system have remained mysterious. Durand et al. identified a
collection of small RNAs and
their respective matching targets
within a self-incompatibility locus
in Arabidopsis halleri. A subset
of these alleles functioned in a
dominant manner, which helps to
explain how self-incompatibility is
maintained. LMZ
Science, this issue p. 1200
BIOCHEMISTRY
Cancer by activating
a binding partner
Some cancers are associated
with mutations that increase
the activity of members of the
EGFR family of tyrosine receptor
kinases, such as HER1. Although
HER3 has little kinase activity, it
can contain cancer-associated
mutations. HER3 binds to other
EGFR family members that do
have kinase activity. Littlefield
et al. crystallized the kinase
domain of HER1 bound to the
kinase domain of normal HER3
or HER3 with cancer-associated
mutations. Cancer-associated
mutations in HER3 increased
its binding to and allosteric
activation of HER1. NRG
Sci. Signal. 7, ra114 (2014)
MICROBIOLOGY
Aiding shipworms
appetite for destruction
STEM CELLS
An inhibitor to expand
mouse stem cells
Among their many side effects,
treatments such as chemotherapy and radiation deplete
stem cells. Therapies to boost
their numbers may positively
affect patient outcomes. One
possible therapeutic target is
the protein phosphatase SHIP1.
Hematopoietic stem cells
expand in mice lacking SHIP1;
sciencemag.org SCIENCE
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PHOTO: R. M. OCONNOR ET AL., PNAS 111, 47 (25 NOVEMBER 2014) 2014 NATIONAL ACADEMY OF SCIENCES
SUPERCONDUCTIVITY
Beefing up thin-film
superconductivity
Conventional superconductors,
such as lead, owe their exotic
properties to the interaction of
electrons and lattice vibrations.
In contrast, in the more recently
discovered iron-based superconductors (IBSs), magnetic
interactions are thought to play
a major role. Lee et al. show
that in thin films of the IBS FeSe
deposited on SrTiO3, a combination of the two mechanisms may
be at play. Using angle-resolved
photoemission spectroscopy, they
discovered a vibrational mode of
the SrTiO3 substrate that interacts
with the electrons in FeSe. The
interaction roughly doubles the
temperature at which the film
becomes superconducting, when
compared with bulk IBSs with
similar electronic structures. JS
Nature 515, 245 (2014).
MARINE BIOLOGY
PHOTOS: (TOP TO BOTTOM) GUOQIANG WAN; LIU YILIN XINHUA NEWS AGENCY/NEWSCOM
NANOMATERIALS
Hair cells (green) and supporting cells
(cyan) in a neonatal mouse cochlea
HEARING LOSS
TROPICAL PALEOECOLOGY
NATURAL HAZARDS
SCIENCE sciencemag.org
Published by AAAS
1197
RESEARCH
Lunar magnetism
persisted via dynamo
Today the Moon has no magnetic
field, but this was not always the
case. Remnant magnetization
in lunar rock and crust samples
indicates that substantial fields
existed billions of years ago.
Weiss and Tikoo review how
modern magnetic studies have
established that these fields were
powered by a magnetic dynamo
that lasted from 4.2 to 3.56 billion
years ago. However, the possible
mechanics behind the dynamo,
such as mantle precession or
core crystallization, remain under
investigation. To find out how
and when the dynamo came
and went now requires improvements in magnetohydrodynamic
models and more accurate
paleointensity measurements,
possibly even those that show
the field direction. MMM
Science, this issue p. 1198
CELL COMPETITION
1197-B
HEAVY FERMIONS
SURFACE STRUCTURE
Stabilization of the
surfaces of magnetite
Accurate structures of iron oxide
surfaces are important for understanding their role in catalysis,
and, for oxides such as magnetite, applications in magnetism
and spin physics. The accepted
low-energy electron diffraction
(LEED) structure for the surface
of magnetite, in which the bulk
surface termination undergoes
an undulating distortion, has a
relatively poor agreement with
experiment. Bliem et al. show
PALEOCLIMATE
STRUCTURAL BIOLOGY
Watching a protein
molecule in motion
X-ray crystallography has
yielded beautiful high-resolution
GEOLOGY
sciencemag.org SCIENCE
Published by AAAS
RESEARCH
REVIEW SUMMARY
LUNAR GEOLOGY
The interior structure of the Moon and the lunar dynamo. New magnetic measurements of lunar rocks have demonstrated that the ancient Moon generated a dynamo
magnetic field in its advecting liquid metallic core (innermost red shell). This dynamo may
have been driven by convection, possibly powered by crystallization of the core (innermost red sphere) and/or stirring from the solid mantle (thick green shell). The magnetic
field was recorded as magnetization by rocks on the lunar surface. [Image created by
Hernn Caellas]
1198
sciencemag.org SCIENCE
Published by AAAS
R ES E A RC H
REVIEW
LUNAR GEOLOGY
1
Department of Earth, Atmospheric, and Planetary Sciences,
Massachusetts Institute of Technology, 77 Massachusetts
Avenue, Cambridge, MA 02139, USA. 2Department of Earth
and Planetary Science, University of California, Berkeley, CA
94720, USA. 3Berkeley Geochronology Center, 2455 Ridge
Road, Berkeley, CA 94709, USA.
SCIENCE sciencemag.org
Fig. 1. Apollo-era paleointensity measurements of the lunar magnetic field and their perceived
difficulties. Each point represents the inferred field intensity for a single lunar rock. Points with arrows
represent upper limits. Both the intensity values and their ages are uncertain. The apparent rapid rise in
paleointensities between 4.0 and 3.9 Ga has been taken as evidence that the origin of the lunar dynamo,
and perhaps of the core itself, were delayed by 500 My (3) (blue), apparently at odds with a giant impact
origin of the Moon. Energy flux scaling (Eq. 2) predicts maximum paleointensities of only less than ~12 mT
(green), apparently well below many measured values. Early thermal evolution models (25, 27) predicted
that a core dynamo powered by purely thermal convection is unlikely to have persisted beyond ~4.1 Ga,
before virtually all measured paleointensities (red). The weakest surface fields today are <0.2 nT (29).
Paleointensities are obtained from the IRM normalization method as compiled by (4, 17).
5 DECEMBER 2014 VOL 346 ISSUE 6214
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R ES E A RC H | R E V IE W
(for which C/MR2 = 0.4, where C is the polar moment of inertia and M and R are the lunar mass
and radius). However, the subsequent discovery
that the mare plains are basaltic by Surveyor 5
(9) and that the lunar highlands are anorthositic
by Apollo 11 (10) confirmed that, beginning with
an early post-accretional large-scale magma ocean,
the Moon repeatedly experienced igneous processing over >2 billion years. Furthermore, improved
spacecraft tracking in the mid-1970s established
that C/MR2 ~ 0.392 T 0.003 (11), consistent with
(although not requiring) the existence of a small
metallic core (1 to 4% of the lunar mass) (12, 13).
Finally, the discovery of natural remanent magnetization (NRM) in the lunar crust and Apollo
samples from spacecraft and laboratory measurements hinted at the possibility of an ancient but
now extinct lunar dynamo.
Paleomagnetic measurement methodology
NRM is a vector quantity that reflects the macroscopic, semipermanent alignment of electron
spins within ferromagnetic minerals. The major
ferromagnetic minerals in unbrecciated lunar
rocks are the metals kamacite (a-Fe1xNix for
x < ~0.05) and martensite (a2-Fe1xNix for ~0.05 <
x < ~0.25) (4, 14, 15). Igneous rocks become magnetized when they cool in the presence of an
ambient magnetic field below their Curie temperature (780C for kamacite), a form of NRM
known as thermoremanent magnetization (TRM).
For weak planetary fields, the TRM intensity,
M, is given by
M = cTRM Bpaleo
(1)
3
Rc
aG 1=3
2m0 cf r1=2
4pR2c qa 1=3
R
3CP
2
RE S EAR CH | R E V I E W
rounded white clast at the location shown in (B) (JSC photo 02973-x2). The
rounded clast is revealed to be itself a breccia composed of shocked fragments
of plagioclase (gray) and pyroxene and olivine (brown, green, and purple grains).
The rock is also crosscut by fine-scale (<0.1 mm thick) young, glassy melt veins
(dark, sinuous, branching channels) that postdate assembly of the breccia (one
example identified with white arrows). (D) Summary of radiometric and exposure ages (top) and inferred geologic events (bottom) for 67915.The Pu/Xe and
oldest 40Ar/39Ar ages indicate that the oldest materials crystallized at 4.3 Ga.
40
Ar/39Ar age spectra from a diversity of clastsincluding granulitic breccias,
pristine troctolites, and sodic ferrogabbrosexhibit a range of apparent ages
from 4.3 to <0.26 Ga, thought to correspond to multiple thermal events recorded
by clasts with different Ar diffusivities. 81Kr/Kr, 38Ar, and 21Ne cosmic ray exposure ages are thought to date the impact that formed Cone Crater and
excavated 67915. Ages and paleomagnetic data are presented in table S7.
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1246753-4
basalts and breccias and 3 to 5 orders of magnitude larger than the NRM of pristine feldspathic rocks, which are thought to make up the
majority of the lunar crust (35).
By the end of the Apollo era, it was certain
that there were magnetic fields on the ancient
Moon, but the intensity, timing, and, most important, physical origin of these fields were
unclear. Whether the magnetization in lunar
materials was the product of a core dynamo
Orbiter Laser Altimeter (83), showing centers of selected major impact basins
(circles) and the antipodes of the eight youngest impact basins (stars). Basin
centers are identified by the following: B, Bailly; C, Crisium; M, Moscoviense; MR,
Mendel-Rydberg; He, Hertzsprung; Hu, Humboldtianum; I, Imbrium; O, Orientale;
N, Nectaris; Sc, Schrdinger; Se, Serenitatis; Sm, Smythii; SPA, South PoleAitken; and SR, Sikorsky-Rittenhouse. Basin antipodes are identified with the
previous abbreviations appended by A. [Modified from (72)]
sciencemag.org SCIENCE
RE S EAR CH | R E V I E W
SCIENCE sciencemag.org
mantle boundary typically find that a pure thermal convection dynamo can persist at most until
sometime between ~4.3 and 4.1 Ga (25, 26, 44).
Enrichment of the lower lunar mantle by 40
parts per million (ppm) H2O [below that recently inferred from mare basalts (45)] may lower
the mantle viscosity by ~2 orders of magnitude,
thereby enabling a thermal convection dynamo
to persist until perhaps ~3.4 Ga (46). Additionally, it is possible that at the end of magma ocean
crystallization, the core was surrounded by a
radiogenic element-rich cumulate layer. Thermal
models (26) have estimated that radioactive heating from such a layer could delay initiation of a
thermal convection dynamo (47) until sometime
between 4.2 and 4.0 Ga and postpone the dynamo cessation until perhaps ~3.5 Ga. The generally short lifetime predicted for a purely thermal
convection dynamo has motivated alternative
hypotheses that the lunar dynamo was powered
either by thermochemical convection from core
crystallization or mechanical stirring by differential rotation between the mantle and core.
Mechanical dynamos have yet to be definitively identified to operate in any planetary body
today. At least two forms of these exotic mechanisms have been described for the Moon. The
first proposes that large meteoroid impacts, by
driving transient large-amplitude mantle librations or even temporarily unlocking the mantle
from synchronous rotation, powered dynamos
lasting for up to several thousand years after
each impact event (48, 49). However, because
this mechanism requires highly energetic impact
(26)], or 400C (dark red curve); and (iv) inner core crystallization (light red and
dark red curves) [model H50E100V5e19 of (54) and DTCMB = 400 K, 4 wt % S
model of (56), respectively]. Dashed curves depict estimated power from mantle
precession for the three orbital models considered by (23): Nominal (green),
Model 2 (light blue), and Model 4 (dark blue). Gray boxes indicate the estimated
range of critical core heat flux values required for field generation in thermal
convection dynamos (3 to 10 mW m2). Dynamos driven by precession and
compositional convection may operate subadiabatically. (Inset) Magnified
model results for the time spanning 3.9 to 2.4 Ga. Because the models shown
here assumed modestly differing properties of the lunar interior (e.g., core radius
and sulfur abundance), their results should only be compared qualitatively.
1246753-5
R ES E A RC H | R E V IE W
events, it seems only likely to have occurred before the final basin-forming impact that formed
the crater Orientale at 3.72 Ga (50).
A mechanical dynamo could also be powered
by mantle precession (40, 51). Although initially
locked, the lunar core and mantle began to precess around different axes when the Moons orbit
migrated to beyond >26 to 29 Earth radii (52).
The power associated with this differential motion should have been extremely large compared
to that driving thermal convection (Fig. 4) but
declined over time as the angle between the core
and mantle spin axes decreased. The lifetime
of the precession dynamo is highly uncertain
because it could operate subadiabatically and
because the lunar orbital evolution is poorly
constrained. Nevertheless, for a typical choice
of lunar physical parameters (23), it is estimated
that such a dynamo could be sustained until the
Earth-Moon distance reached ~43 to 48 Earth
radii (i.e., until ~3.4 to 2.0 Ga, depending on
which lunar orbital evolution model is assumed).
Considering a broader range of acceptable lunar
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RE S EAR CH | R E V I E W
anorthositic breccia 60015 (14, 18) (Fig. 6). Finally, a recent paleomagnetic study of a <7-Ma
impact melt glass demonstrated that lunar surface fields likely remained below ~7 mT into the
recent past (62, 63) (Fig. 6).
The 4-mT upper limit inferred for the Moon
at 3.19 Ga is nevertheless at the upper end of
the range predicted by the energy flux scaling
(Eq. 2). In particular, for typically assumed lunar parameters, essentially all published thermal and evolution models of the lunar dynamo
(23, 26, 44, 47, 48, 56) predict surface fields
>0.1 mT for more than ~90% of the time period
in which the dynamo is active in the models.
Such a minimum field is comparable to estimates of the strongest lunar crustal surface fields
(64) and below even the weakest known dynamo surface field in the solar system today
(65). Therefore, the current paleointensity constraints do not at all exclude a lunar dynamo
after 3.56 Ga. Determining when the dynamo
truly ceased requires a sample with extraordinary magnetic recording capabilities. Tikoo et al.
(66) very recently suggested that the melt-glass
matrix of regolith breccia 15498 (which contains
a population of high-fidelity single-domain ferromagnetic grains) was magnetized in a ~2-mT
core dynamo field. The age of the TRM is uncertain but must be less than the 3.3 Ga age of
basalt clasts in the breccia, whereas trapped Ar
data suggest a lithification age of perhaps only
~1.3 Ga (67). If confirmed, this result would extend the lifetime of the lunar dynamo by at least
0.2 to 2 billion years, suggesting that the dynamo
operated in two different intensity regimes: a
pre-3.3-Ga high-field epoch and a post-3.3-Ga
weak-field epoch. Depending on the age of the
NRM in 15498, precession and core crystallization are the only mechanisms currently thought
5 DECEMBER 2014 VOL 346 ISSUE 6214
1246753-7
R ES E A RC H | R E V IE W
Fig. 6. Paleointensity measurements of the lunar magnetic field using modern methods and estimated lifetimes of various lunar dynamos. Each point represents measurements of a single Apollo
sample. Circles represent actual paleointensities, downward arrows represent upper limits on paleointensity, and right arrows represent upper limits on age. Green and blue points were measured using the
IRM and Thellier-Thellier methods, respectively, and the upper limits were derived from the ARM method.
Note the datum at <7 Ma at the extreme right of the figure.The shaded green box encompasses the mean
paleointensity value for the period 3.56 to 4.25 Ga (central green line) and its estimated 2 SD uncertainty
(upper and lower lines) (see table S6). The paleointensity value for 76535 (leftmost green point) is
currently not well constrained due to spurious demagnetization effects. Vertical dashed lines show
estimated maximum lifetimes of various proposed lunar dynamo mechanisms: purely thermal convection
in a dry mantle (with and without an early thermal blanket surrounding the core); impact-driven mantle
rotational changes, purely thermal convection in a wet mantle; precession; and thermochemical core
convection driven by core crystallization. The horizontal line shows the maximum lunar surface field as
estimated from Eq. 2. The lifetimes of the precession and thermochemical dynamos are highly uncertain.
Paleointensity data set is from (14, 18, 50, 5961, 63, 66) and is also listed in table S6.
of another geologic association with the southern farside anomalies: They lie on the northern
edge of the vast South Pole-Aitken (SPA) basin,
the largest known impact crater on the Moon.
This discovery has led to two explanations
for these anomalies as alternatives to the impactplasma fields hypothesis. Both propose that rather than reflecting locally high Bpaleo, they are a
manifestation of materials with locally high cTRM
that were magnetized in a dynamo field. The first
of these proposals (71) identifies the anomalies
with the inferred locations of <4.0 Ga basaltic
dikes located within the SPA (Fig. 3). The assumed
great vertical thickness (30 km) of these dikes
enables them to produce the strong SPA anomalies despite the weak NRM of typical mare basalts.
The second proposal (72) emphasizes that
the anomalies are in the expected locations of
ejecta and impact melt derived from the SPA
impactor (Fig. 3). The key advantage of this
proposal is that typical chondritic impactors,
which are far more Fe-rich, have cTRM ~100
and 104 times that of mare basalts and pristine
feldspathic rocks, respectively. Therefore, deposition of relatively thin layers (perhaps just tens
to thousands of meters thick) of such material
should have fundamentally enhanced the magnetic properties on the surface. SPA ejecta, as well
as that from more recent impacts, could also account for most of the other isolated crustal anomalies. However, thus far no evidence has yet been
identified indicating that at least the surface layer
(top ~30 cm) of the southern farside anomaly
region is anomalously iron metal-rich (73).
Both of these new proposals require only dynamo fields rather than plasma fields to explain
the magnetic anomalies. Yet more evidence for
a dynamo comes from the recent identification
of many anomalies within the interiors of most
Nectarian (i.e., ~3.85 to 3.92 Ga) basins and some
pre-Nectarian (i.e., more than ~3.92 Ga) basins
(Fig. 3) (7476). These impacts likely demagnetized any existing NRM, producing impact melt
sheets that should have cooled from the kamacite Curie point to ambient lunar surface temperatures far more slowly (given their >1 km
thicknesses, over a period of >104 years) than
any impact-generated field. Therefore, these
central basin anomalies likely require a steady
field like that expected from a core dynamo
during the Nectarian and pre-Nectarian periods.
However, once again, the inferred TRM intensities (~1 A m1) exceed what is achievable with
known endogenous lunar materials and may reflect the addition of Fe-rich impactor materials.
It has been suggested (29) that the apparent
lack of anomalies in the centers of Imbrian basins (<1 nT at the surface, suggesting that the
central impact melt layer has an NRM less than
~103 A m1) may indicate that the dynamo was
no longer active by the time of their formation
(i.e., after ~3.85 Ga). However, this proposal is
not consistent with the strong evidence for a
core dynamo from paleomagnetic analyses of
3.56 to 3.85 Ga (Imbrian-aged) Apollo 11 and 17
basalts (see above). In fact, the sample and orbital data sets are consistent if the composition
sciencemag.org SCIENCE
RE S EAR CH | R E V I E W
1246753-9
R ES E A RC H | R E V IE W
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.
36.
37.
38.
39.
1246753-10
40.
41.
42.
43.
44.
45.
46.
47.
48.
49.
50.
51.
52.
53.
54.
55.
56.
57.
58.
59.
60.
61.
62.
63.
64.
65.
66.
67.
68.
69.
70.
71.
72.
73.
74.
75.
76.
77.
78.
79.
80.
81.
82.
83.
SUPPLEMENTARY MATERIALS
www.sciencemag.org/content/346/6214/1246753/suppl/DC1
Tables S1 to S7
10.1126/science.1246753
sciencemag.org SCIENCE
and Minute-induced cell competition revealed requirements for two related but
distinct cohorts of components from the
IMD and Toll pathways. Both signaling
cohorts required the extracellular ligand
Sptzle and noncanonical Toll-related receptors (TRRs) and led to elimination of
the less-fit loser cells by inducing NFBdependent activation of proapoptotic
genes. However, our analysis uncovered in-
Hi-Myc
RpL14-/+ WT
TRRs 3,9
Competition
WT
Hi-Myc
TRRs 2,3,8,9
Sptzle
Dredd Ect4
Cactus
dl,Dif
Rel
DEATH
1
dl,Dif
rpr
Rel
hid
SCIENCE sciencemag.org
Published by AAAS
1199
R ES E A RC H
RESEARCH ARTICLE
CELL COMPETITION
SCIENCE sciencemag.org
Myc-regulated cell competition has been documented in Drosophila wing imaginal discs and
among mouse epiblast cells, where fluctuations
in Myc expression precede the transition of cells
to a more limited developmental potential and
correlate with survival (9, 11, 14, 19).
The Toll signaling pathway was first identified
as a developmental regulatory module that patterns
the embryonic dorsoventral axis in Drosophila
but was subsequently recognized to also function
in host defense against infection. Toll-like receptors and NFkB transcription factors are key signaling and transcriptional mediators of the
ancient and broadly conserved innate immune
recognition system activated in response to nonself (e.g., microbial infection) or altered-self cells
(e.g., viruses, cancer) (2024). The early appearance of these factors in metazoan evolution and
their conserved use in altered-self recognition in
a process akin to cell competition led us to probe
whether they mediate the response to mutant
or otherwise compromised cells in developing
tissues. We report here that components of the
innate immune system function to eliminate cells
recognized as unfit during cell competition.
Mutations in the Toll and IMD pathways
prevent Myc-induced cell competition
To test for involvement of innate immune factors
in cell competition, we used a simple genetic assay in wing discs for Myc-induced competition,
wherein cells constitutively express a Myc transgenic FRT cassette (>Myc>), increasing the expression of Myc ~1.5-fold above the endogenous
level (11, 13). Removal of this cassette via Flp/
FRT-mediated recombination generates clones
of WT cells and concomitantly allows expression
of the Gal4 transcriptional activator and the
Gal4-regulated upstream activating sequence
green fluorescent protein (UAS-GFP) (Fig. 1A).
1258236-1
R ES E A RC H | R E S EA R C H A R T I C LE
Toll pathway
1500
ns
MyD88
pll7
Dif1
dl1/dl4
spzrm7
ird5
RelE20
key1
tak12
BG4/Fadd
Dredd B118/Y
67 59 42 53 70 108 35 48 34 21 36 98
500
Ctls
5000
Losers
40
ns
***
3000
2000
***
Cells/clone
*** ***
4000
*** ***
30
***
20
Controls
41
30 56
45
56
WT
Toll-3MiMIC
Toll-8R23B
Toll-9KO
43 47
Toll-9KO
Toll-8R23B
Toll-3MiMIC
Toll-9KO
WT
Toll-8R23B
68 39 56 81 72
Toll-3MiMIC
WT
90
WT Ctl
10
1000
0
135 135 96 79
WT
ns
Ect44273
ns
ns
***
***
1000
WT
*** ***
*** ***
Ect44273
IMD pathway
imd shadok
PGRP-LC 5
WT Ctl
1258236-2
WT loser
Toll receptor in innate immunity and in embryonic dorsoventral patterning (30, 3335).
However, RNA interference (RNAi)mediated
knockdown of Toll itself (Toll-1) had little effect
on competition (fig. S2C). The Drosophila genome
encodes nine Toll-related receptors (TRRs), but
most are poorly characterized. Notably, loss of
Toll-2 (18wheeler), Toll-3 (Mst-Prox), Toll-8 (tollo),
and Toll-9 blocked elimination of the loser cells
and significantly increased clone size (Fig. 1, E
and F, and fig. S2A) but did not affect growth
of control clones (Fig. 1E and fig. S2A). RNAimediated knockdown specifically in the loser
population indicated that the receptors were
required within the loser cells (fig. S2A). By contrast, RNAi directed against Toll-4, -5, and -7 had
little effect on competition (fig. S2C).
Losers
Losers
the nucleus and activate AMP genes. In the Toll pathway, binding of Spz to the
Toll receptor recruits a complex consisting of Myd88, Tube/IRAK-4, and Pelle/
IRAK-1, which phosphorylates Cactus/IkB and targets it for degradation. This
releases Dorsal or Dif for activation of distinct AMP genes in the nucleus. Components outlined in red denote those also required for Myc-induced cell competition. P, phosphorylation. (E and F). Null mutations in Toll-3, Toll-8, or Toll-9
suppress elimination of loser cells, increasing loser clone size (E) and cell number
per clone (F), but do not alter control clones. Error bars in this figure are SEM. All
P values are relative to WT loser clones except in (E) at leftthey are relative to
WT control clones. ***P < 0.001. ns, not significant (Mann-Whitney test).
sciencemag.org SCIENCE
RE S E ARCH | R E S E A R C H A R T I C L E
DNA
TUNEL
GFP
15
10
***
5 ***
WT ctl
***
144 103
Ctls
***
2000
1000
51
WT ctl
18 51 75 56 27 42 14
ns
42
Rel RNAi
Dredd B118/Y
Losers
1000
RelE20/Dredd B118
Ctls
Rel RNAi
+ dcr-2
42 35 81 63 32
WT
Rel RNAi
ns
Dredd B118/+
1000
***
WT
***
***
2000
80
Losers
3000
RelE20/+
***
TUNEL
***
ns
WT
3000
4000
2000
TUNEL
3000
ns ***
WT
Rel RNAi
5000
TUNEL
Dredd B118/Y
TUNEL
98
tolloR23B
DNA
TUNEL
GFP
20
81
59
UAS-Dredd
+Rel RNAi
DNA
TUNEL
GFP
25
WT
DNA
TUNEL
GFP
RelE20
Losers in WT
TUNEL(+) cells/clone
WT controls
UAS-Dredd
Losers
Fig. 2. Dredd-mediated Rel activation eliminates loser cells. (A) Rel and Tollo are required for death
of loser cells. TUNEL assays showing cell death in control clones, WT loser clones, and loser clones in
RelE20 and tolloR23B mutant backgrounds 24 hours after clone induction. Many WT loser cells are TUNELpositive, but their death is suppressed in the mutant backgrounds. (B) Quantification of data from (A).
(C) Expression of Rel-RNAi (or Rel-RNAi and Dicer to enhance RNAi efficiency) within loser cells
suppresses their loss but does not affect control clones. (D) Dredd and Rel are critical for elimination of
WT loser cells. Loss of Dredd suppresses the loss of loser clones but does not affect the growth of
control clones. Whereas DreddB118/+ has little effect on loser death, in combination with RelE20/+ it
strongly suppresses death. (E) Expression of Dredd in the absence of competition reduces clone size,
but this is blocked by coexpression of Rel RNAi. The number of clones scored for each genotype is
indicated. All P values are relative to WT loser clone size except in (C): ***P < 0.001. ns, not significant
(Mann-Whitney test).
SCIENCE sciencemag.org
1258236-3
R ES E A RC H | R E S EA R C H A R T I C LE
20
15
**
DNA
hid-lacZ
GFP
**
10
ns
1000
ns
DNA
-gal
% C3-positive cells
ncCM
cCM
***
800
600
400
200
0
GFP Rel68
+/+
+/+
Rel
+
DNA
hid-lacZ
GFP, Rel68
% nuclear Rel-GFP
80
70
**
+/+
60
DNA
-gal
hid PlacZ/+
50
40
***
ns
cCM
ncCM
+PGN
-PGN
30
GFP, Rel68
1258236-4
Rel68
hid lacZ/+
WT GFP
+
dsRNA: -
GFP, Rel68
hid
rpr
in the hid locus (hid-lacZ, red). DNA is stained with Hoechst (blue). hid-lacZ is
not induced in control clones (C and C). Rel68-expressing clones activate expression of hid-lacZ (D and D), grow poorly, and are small compared with controls (E and G) but are rescued by the mutation caused by the hidPlacZ
insertion (F and G). Clones in (D) are shown at higher magnification to facilitate
resolution of hid-lacZ expression. (G) Average clone size of control clones (GFP),
Rel68-expressing clones in a +/+ background, and Rel68-expressing clones in the
hidPlacZ/+ mutant background. Error bars denote SEM. (H) Expression of UASRel68 in WT wing discs induces hid mRNA but not rpr mRNA in quantitative RTPCR experiments on RNA isolated from wing discs. Error bars indicate SD.
***P < 0.001, **P < 0.01, *P < 0.05. ns, not significant (Mann-Whitney test).
sciencemag.org SCIENCE
RE S E ARCH | R E S E A R C H A R T I C L E
RNAi
nc MRpL14-/+ controls
MRpL14-/+ losers
ns
57
85
106
Losers
MRpL14-/+ losers+cactus
4000
3000
2000
1000
DNA
TUNEL
GFP
TUNEL
TUNEL
TUNEL
DNA
TUNEL
GFP
SCIENCE sciencemag.org
TUNEL
20
15
UAS-GFP UAS-dl
14
DNA
TUNEL
GFP
RNAi
5000
DNA
TUNEL
38
hid
103
rpr
H99/+
20
OE
MRpL14-/+
salE>gRpL14
**
p35
**
40
GFP
/+ clones
RpL14
20
nc ctl
OE
nc
Rel
No competition:
GFP+ clones present
Dif
dl
% larvae with M
20
***
60
**
40
80
MRpl14 -/+
+ cactus
40
60
MRpl14 -/+
*** ***
MRpl14 -/+
***
60
***
dl1/DfTW
Competition:
GFP+ clones eliminated
80
***
80
+cact
MRpL14-/+
salE>Gal4
UAS-GFP
UAS-X
100
MRpl14 -/+
salE>RpL14>Gal4
UAS-GFP
% larvae with M
RpL14-/+
M RpL14-/+
Conclusions
Altogether, our results demonstrate that the conceptual resemblance between cell competition
and innate immunity is matched with genetic
and mechanistic similarities. Thus, cells within
developing tissues that are recognized as mutant or compromised are competitively eliminated
via a TRR- and NFkB-dependent signaling mechanism. Although similar core signaling components are activated in both processes, cell
+RpL14
clones
***
12
hid
rpr
10
8
6
4
**
2
0
UAS-dl
UAS-Dif
Left to right: control noncompetitive (nc) MRpL14/+ wing disc rescued by gRpL14
expression; control nc act>Gal4, UAS-GFP clones generated in unrescued
MRpL14/+wing disc; MRpL14/+ loser clone generated in MRpL14/+ salE>gRpL14
disc; and MRpL14/+ loser clone expressing UAS-cactus generated in MRpL14/+
salE>gRpL14 disc. (E) Competitive elimination of MRpL14/+ loser cells requires cell
death induced by Reaper (Rpr). Elimination of MRpL14/+ loser clones is prevented
by expression of the apoptosis inhibitor p35; by one copy of the H99 deficiency
that removes hid, rpr, and other proapoptotic genes; and by RNAi against rpr, but
not RNAi against hid. Number of larvae scored per genotype is indicated. (F)
dorsal-expressing clones are smaller than WT sibling clones. Mitotic recombination generated GFP-positive clones that express UAS-dorsal and WT GFP-negative
sibling clones. The graph shows the mean clone size of the indicated number of
clones. (G) Expression of UAS-dorsal and UAS-Dif induces expression of rpr, but
not hid, in quantitative RT-PCR experiments on RNA isolated from wing discs.
Error bars in (B) and (E) to (G) denote SD; error bars in (C) indicate SEM. ***P <
0.001, **P < 0.01, *P < 0.05 (Students t test).
5 DECEMBER 2014 VOL 346 ISSUE 6214
1258236-5
R ES E A RC H | R E S EA R C H A R T I C LE
Bioinformatics
Rel binding sites were identified in the hid/W
locus using the Web-based program Genome
Surveyor (44).
Cell culture and generation of
conditioned medium
Drosophila S2 cells were maintained at 25C in
Schneiders Drosophila Medium supplemented
Table 1. Suppression of MRpL14/+ loser clone elimination by Toll and IMD pathway genes. P values
are compared with that of MRpL14/+ (t test).
Genotype
M
UAS-RpL14
UAS-p35
UAS-cact
UAS-cact
UAS-dl
UAS-Dif
UAS-pelle
UASToll-3
UASToll-9
UAS-spz
UAS-imd
UAS-Rel
Fig. 5. A model of the
role of TRR-NFkB
modules in cell competition. A model for
TRR-NFkB function in
cell competition,
incorporating data from
both competitive contexts. See text for full
description. We propose
that in both Minute- and
Myc-induced competitive
contexts, loser cells are
eliminated through a
signaling pathway mediated by Spz, TRRs, and
NFkB function. Contextand/or geneticdependent interactions
that cause Rel activation
trigger expression of Hid,
whereas those activating
Dif and Dorsal trigger
expression of Rpr.
Apoptosis is the ultimate
fate of both loser populations. Red denotes genes
required for Myc-induced
competition, blue indicates those required for
competition in M/+
mosaics, and orange
denotes shared
requirements.
SD
15.5
3.7
RpL14/+
MRpL14/+ + overexpression
80.5
43.8
49.2
MRpL14/+ + RNAi
6.6
41.8
37.0
24.3
39.3
30.1
28.8
27.0
29.6
P value
2.5
2.3
1.4
4.30 105
4.40 107
0.03
4.7
7.1
11.9
1.4
13.3
8.0
8.2
4.2
6.8
0.03
0.00
0.03
0.01
0.03
0.03
0.04
0.01
0.02
Competition
WT
RpL14-/+
Competition
Hi-Myc
WT
TRRs 3,9
WT
Hi-Myc
TRRs 2,3,8,9
Sptzle
Dredd Ect4
Cactus
dl,Dif
Rel
DEATH
dl,Dif
rpr
Rel
hid
sciencemag.org SCIENCE
RE S E ARCH | R E S E A R C H A R T I C L E
24 hours T 125 mM CuSO4 treatment and centrifuged at 1200 revolutions per minute for 3 min.
125 mM CuSO4 was added to the conditioned
medium without CuSO4 to control for inducer
effects (12). The CM was then used for treatments
in experiments. Activity of the CM was verified by
C3 activity measurements as described (12). For
Rel-GFP assays, S2 cells were transfected with a
pMT-Rel-GFP transgene (gift from E. Foley) as follows: WT S2 cells were seeded in a 12-well plate with
one coverslip per well at 8 105 cells per ml and
cultured at 25C overnight. Transfection was done
using Cellfectin II Reagent as directed (Invitrogen).
Subcellular location of pMT-Rel-GFP in WT-S2
cells or Myc-S2 cells was scored after treatment T
50 mg/ml LPS as a source of PGN (Sigma) + 125 mM
CuSO4, cCM, or ncCM. At least 200 cells per condition were scored per experiment. The ratio of cells
positive for nuclear localization of GFP-Relish to
total GFP-positive cells was calculated.
Drosophila stocks and care
The following mutant strains were used in this study:
RelE20 (gift from D. Hultmark); imdshadok and imd1
(gifts from D. Ferrandon); key1, tak11, tak12, and Dif1,
(gifts from B. Lemaitre); FADDf06954 (BG4), DreddB118,
ird5KG08072, pll2, pll7, tube2, spz2, dl4, dl1, Df(2L)
TW119, Df(2L)J4, cact1, cact4, hidW05014 (Plac-Z),
Ect44273, and RelE38 (from Bloomington Drosophila
Stock Center, http://flystocks.bio.indiana.edu); caspP1
(gift from J. Chung); Tollo145 (gift of J. Kim);
TolloR23B and TolloR5A (gifts from P. Simpson);
PGRP-LCTotemD5 and PGRP-LCE12 (gifts from
B. Matthey-Prevot); and spzrm7 (gift from C. Hashimoto).
For Myc competition, control or loser clones were
generated in wild type, mutants, or with UAStransgenes using these strains: yw hsflp;;act>y>Gal4,
UAS-GFP (controls), ywhsflp;;tub>Myc>Gal4,
UAS-GFP, ywhsflp;UAS-GFP tub>Myc>Gal4
(11), and ywhsflp;;UAS-Myc (7). ywhsflp;;FRT82B
ubi-GFP, ywhsflp;;FRT82B RelE20/TM6B was
used for fig. S3. FRT82B RelE20 was generated
by recombination and screened for presence
of the mutation by PCR using primers from
(45). For MRpL14/+ competition, UAS-transgenes
were driven in a ywhsflp UAS-GFP::CD8;;
MRpL14(w+)/+salE>gRpL14(w+)>Gal4/TM6B strain.
UAS-RNAi lines were from the Vienna Drosophila
Resource Center (http://stockcenter.vdrc.at) or the
Transgenic RNAi Project (TRiP) collection (BDSC)
and are listed in table S3. Other strains include
UAS-Toll-8DLRR (gift from J.-L. Imler), UAS-dicer2
(BDSC), yw;;UAS-cactus-HA/TM3 and yw;;UAScactus,
yw UAS-GFP hsFLP; tub-Gal80 FRT40, Ubi-GFP
FRT40; tubGal4, UASdorsal, w P(w+, Dros-LacZ),
yw P(w+,Dipt-LacZ), and UAS-Dredd (gifts from
B. Lemaitre), yw;;CecA-lacZ (gift from S. Stven),
UAS-Rel68 (gift from N. Silverman), and UAScaspar (gift from J. Chung). Null mutants were used
when possible. A complete list of mutant strains
is included in table S1. Flies were raised at 25C
on cornmeal-molasses food supplemented with
penicillin and streptomycin, as indicated below.
dsRNA treatment of S2 cells
The sequences for dsRNA against Relish and Spz
were obtained from the Drosophila RNAi ScreenSCIENCE sciencemag.org
discs. In cases where the induced transgene rescues the clones, GFP is readily visible through the
larval cuticle. Each transgene was tested in biological triplicate, and the average number of larvae with surviving clones was compared to the
WT control. yw, yw;;UASRpL14(w+)/TM6B, and
yw;Sp/CyO;UASP35 (w+) strains were used as
controls. Nonparametric t tests were used to test
for statistical significance.
Imaging, image analysis,
and quantifications
Cell clones were imaged in larvae with a Zeiss
MZFLIII and in fixed wing discs with a Zeiss
Axiophot, Leica SP5, or Leica LSM710 confocal
microscope. Clone area was measured (in square
pixels) using ImageJ or Photoshop software. Cells
per clone were counted in the GFP and/or DNA
channels. Only clones in the wing pouch and proximal hinge area were measured; in the MRpL14/+
assays, this corresponded to the salE expression domain. Significance was determined using
Students t and Mann-Whitney tests.
Immunohistochemistry
Wing disc cells were fixed in 4% paraformaldehyde/phosphate-buffered saline (PBS) for 20 min
at room temperature and washed with PBS 0.01%
Tween-20. Hoechst 33258 or 4,6-diamidino-2phenylindole was used to stain DNA. The primary antibodies used include mouse anti-Relish
(DHSB), rabbit anti-b-galactosidase (Cappel), and
rabbit anticleaved Caspase-3 (Cell Signaling Technology). Secondary antibodies were Alexa-conjugated
anti-mouse and anti-rat and Cy3-conjugated
anti-rabbit (Molecular Probes). For cell culture,
cells seeded on glass coverslips were fixed in 4%
paraformaldehyde/PBS for 30 min. Coverslips
were washed with PBS once, permeabilized in
0.5% TX-100/PBS for 5 min, then blocked for
30 min in 3% BSA, 0.1% Tween-20/PBS. After
blocking, each coverslip was flipped onto 50 ml of
primary antibody in blocking buffer for 1 hour at
room temperature. Coverslips were washed 3
with PBS, and primary antibodies were incubated
in AlexaFluor 546 goat anti-rabbit or goat anti-rat
immunoglobulin G (Molecular Probes) for 45 min
at room temperature. Coverslips were washed 3
for 5 min each with PBS, counterstained for DNA
with Hoechst 33258 for 4 min, washed 3 for 5 min
each, then mounted on slides. TUNEL (terminal deoxynucleotidyl transferasemediated deoxyuridine triphosphate nick end labeling) assays
were carried out on fixed discs using either
TUNEL TMR (Roche Diagnostics) or ApopTag
Red (Chemicon).
RNA isolation and qRT-PCR
Total RNA was isolated from wing disc using the
Nucleo SpinII RNA isolation kit (Machery Nagel)
or Trizol (Invitrogen). RNA quantity was measured
using a NanoDrop ND-1000 UV-Vis Spectrophotometer (NanoDrop Technologies, Wilmington,
DE). For qRT-PCR of Rel-expressing cells, yw;;
tub>CD2>Gal4 flies were crossed either with yw;;
UAS-GFP or yw;;UAS-Rel68 and heat-shocked for
30 min at 37C. At 24 hours after clone induction,
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1258236-8
34.
35.
36.
37.
38.
39.
40.
41.
42.
43.
44.
45.
46.
www.sciencemag.org/content/346/6214/1258236/suppl/DC1
Figs. S1 to S8
Tables S1 and S2
2 July 2014; accepted 31 October 2014
10.1126/science.1258236
sciencemag.org SCIENCE
R ES E A RC H
RESEARCH ARTICLE
PLANT GENETICS
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species, and the >50 S alleles observed in outcrossing Arabidopsis species (17) are predicted
to form up to 1225 distinct heterozygous genotypes. In Brassica, class II alleles similarly show a
linear dominance hierarchy that cannot be explained by the action of Smi alone (12, 18). Under
the mechanistic model involving sRNAs and
their targets within the SCR gene, two hypotheses
could explain an allelic series of n distinct S alleles
in a linear dominance hierarchy. First, the mostdominant allele might produce n 1 distinct sRNAs,
each specifically targeting a given more-recessive
S allele, whereas the next-most-dominant allele
might produce sRNAs targeting all S alleles more
recessive than itself, and so on (with the bottom
recessive producing no sRNAs but having targets
for all more-dominant alleles). This model predicts that the top dominant allele produces the
largest number of sRNAs and that the bottom
recessive allele carries the fewest but is targeted
by the largest number of sequences; we refer to
this as model 1 (Fig. 1A). Alternatively, the most
dominant allele might produce a single sRNA,
as in Brassica (12), and all subsequent morerecessive S alleles might carry a specific target
that interacts with the most-dominant allele in
a manner that results in nonexpression (Fig. 1A);
the next allele in the hierarchy (that is recessive
to the top dominant, but dominant to other classes) might produce a different sRNA, whose target sequence is present in all alleles recessive to
itself, and so on. This model (model 2 in Fig. 1A)
predicts that only a single sRNA is produced in
all but the most-recessive allele, where the mostrecessive allele carries the largest number of
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predicted to be the genetic element causing dominance of Ah20 over Ah01 (Figs. 2A and 3B). We
generated Ah20mirS3 A. thaliana transgenic lines,
which we crossed with the AhSCR01 line to obtain
hybrid plants containing the two transgenes in
hemizygous state (i.e., as biologically relevant for
S alleles) (table S1) (19). Pollen from these hybrid
plants could germinate on AhSRK01 stigmas, which
showed that the presence of Ah20mirS3 was
sufficient to abolish the SI response conferred
by AhSCR01 alone (Moods median test P value =
2.2 104 and 3.2 105 on two replicate Ah20mirS3
insertion lines) (Fig. 4A). Finally, we produced a
mutated AhSCR01 line (AhSCR01*) in which we
introduced a set of four point mutations specifically in the Ah20mirS3 predicted target (located
in the intron of AhSCR01) (Figs. 3B and 4B), which
we crossed with lines containing Ah20mirS3. As
expected, pollen from the AhSCR01* Ah20mirS3
hybrid plants was insensitive to the effect of
Ah20mirS3 and was rejected on AhSRK01 stigmas.
Hence, our data demonstrate that our predicted
target site for AhSCR01 was necessary for the regulatory effect of Ah20mirS3 (Fig. 4B). Note that
sRNA target sites within introns of genes have already been observed and validated experimentally,
in a different context (32). Overall, our validation
procedure provides direct experimental support
to identify sRNA genes and their target sites, although a complete experiment involving all 21
predicted sRNA-target interactions would arguably be required to fully validate the proposed
regulatory mechanism between the six S alleles.
Power and accuracy of the
molecular model
We evaluated power and accuracy of our model
of dominance-recessivity interactions by comparing molecular predictions with the phenotypic observations from controlled crosses. With
two exceptions, most (93.1%) of the predictions
connect a sRNA produced by a dominant allele
to a target present in a more-recessive allele above
the proportion expected by chance alone (about
63 to 79%, obtained by 100,000 random permutations) (fig. S6), which suggests that our molecular prediction of target sites is reliable, with a low
false-positive rate. Overall, 13 of the 14 dominancerecessivity relations observed in our tests (93%)
were predicted by at least one sRNA targeting
prediction (Fig. 2A), which suggests that sRNAs
can explain the majority of the interactions we
observe at the phenotypic level. Thus, a simple
molecular model explaining the control of dominance through factors closely linked to the S locus
can predict observed dominance phenotypes.
1202
(dominant alleles on top, recessive alleles at the bottom). Red boxes on the
target side correspond to the two exons on SCR, with arrows giving the direction
of transcription. For clarity, length variation of the SCR intron is not represented.
A line joins a precursor with a target whenever the precursor produces a sRNA
that is predicted to use that target (based on an alignment threshold of 18). Black
and red lines represent targeting relations that are and are not consistent with
the phenotypic network, respectively. The star symbol indicates the sRNA-target
interaction that we functionally validate in Fig. 4 (Ah20mirS3 on AhSCR01).
sciencemag.org SCIENCE
RE S E ARCH | R E S E A R C H A R T I C L E
Fig. 4. In planta functional validation that Ah20 controls phenotypic expression of SCR01
through specific targeting by mirS3. (A) Ah20mirS3 is sufficient to abolish the SI response of AhSCR01.
(B) The mutated target makes AhSCR01* insensitive to the effect of Ah20mirS3. Pollen compatibility was
estimated by the number of germinated pollen grains per stigma. Incompatible crosses (SI) are colored in
yellow-orange, compatible controls (SC) in green, and tests of the mir-target interactions in blue (see table
S1). Mir-target interactions were tested on hybrids produced from two Ah20mirS3-independent insertion
lines, named lines 6 and 12. Thick bars represent the median, boxes the interquartile range, whiskers
extend out 1.5 times the interquartile range, and individual points are outliers.The number of replicates (n)
is indicated above each cross. (A) (Top) The most abundant targeting sRNA produced by the Ah20mirS3
precursor is aligned with the SCR01 predicted target; in (B), it is aligned with the mutated target. Mismatches are represented by x, and G:U wobbles by o. Mutated bases are underlined.
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Fig. 5. Repertoire of sRNA genes and their targets along the phylogeny of S alleles. The
Bayesian phylogeny is based on full A. halleri and A. lyrata SRK amino acid sequences. Node support
was assessed by posterior probabilities (PP), and nodes with PP > 0.95 are represented by white
circles. Phylogenetic classes are reported (I, II, III, and IV) (15). The scenario of gain and loss for
sRNA precursors and their targets is represented on the branch where each event is inferred. Solid
and open squares indicate precursors that are inferred to have been present and absent from the
ancestral repertoire, respectively. For each precursor, asterisks for BFs indicate the level of
confidence on the presence or absence from the ancestral repertoire. *BF > 2: some evidence,
**BF > 5: strong evidence, and ***BF > 10: very strong evidence (38).
sciencemag.org SCIENCE
R ES E A RC H
REPORTS
S.B., and V.C. wrote the paper. V.C. and S.B. coordinated the project.
S-locus sequences have been deposited on the European Molecular
Biology Organization public database, sRNA sequences on the GEO
public database (see table S2 for accession numbers), and data
matrices and phylogenetic trees on the TreeBASE database (http://
purl.org/phylo/treebase/phylows/study/TB2:S16394). BAC clones
produced in this project will be distributed through http://cnrgv.
toulouse.inra.fr/Library/Arabidopsis upon signature of a material
transfer agreement for in-house academic research.
VALLEYTRONICS
Ultrafast generation of
pseudo-magnetic field for valley
excitons in WSe2 monolayers
Jonghwan Kim,1* Xiaoping Hong,1* Chenhao Jin,1 Su-Fei Shi,1,2 Chih-Yuan S. Chang,3
Ming-Hui Chiu,4 Lain-Jong Li,3,4 Feng Wang1,2,5
The valley pseudospin is a degree of freedom that emerges in atomically thin
two-dimensional transition metal dichalcogenides (MX2). The capability to manipulate it, in
analogy to the control of spin in spintronics, can open up exciting opportunities. Here,
we demonstrate that an ultrafast and ultrahigh valley pseudo-magnetic field can be
generated by using circularly polarized femtosecond pulses to selectively control the valley
degree of freedom in monolayer MX2. Using ultrafast pump-probe spectroscopy, we
observed a pure and valley-selective optical Stark effect in WSe2 monolayers from the
nonresonant pump, resulting in an energy splitting of more than 10 millielectron volts
between the K and K valley exciton transitions. Our study opens up the possibility to
coherently manipulate the valley polarization for quantum information applications.
Energy (eV)
SCIENCE sciencemag.org
www.sciencemag.org/content/346/6214/1200/suppl/DC1
Materials and Methods
Figs. S1 to S12
Tables S1 and S2
References (3957)
31 July 2014; accepted 17 October 2014
10.1126/science.1259442
PL (a.u.)
SUPPLEMENTARY MATERIALS
Energy (eV)
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degenerate energy levels. However, they have distinct optical selection rules and couple to photons
of opposite helicity (3, 4). The dipole transition
matrix element of the K valley A-exciton is characterized by PKsT K jp^x Tip^y j0, where jK and j0
correspond to the K valley A-exciton and ground
state, respectively; p^x ; p^y are momentum operators;
and s+/s corresponds to left/right circular light.
In MX2 monolayers, PKs has a finite value, but PKs
is approximately zero. Consequently, the A-exciton
at the K valley couples exclusively to left-circularly
polarized light. The A-exciton at the K valley, on
the other hand, couples only to the right-circularly
polarized light. This dipole selection rule for valley
does not affect K valley A-exciton resonance because of the optical selection
rule (right). (B and C) Transient reflection spectra of A-exciton resonance at 77 K.
The color scale, horizontal axis, and vertical axis represent the relative reflectivity change DR/R, the probe photon energy, and the pump-probe time
delay, respectively. For atomically thin WSe2 on sapphire substrate, DR/R is
proportional to absorption change. Nonresonant s+ pump of photon energy
at 1.53 eV leads to (B) a strong transient absorption signal for probes with the
same polarization s+, but (C) produces no transient response for probes with
the opposite polarization s.
ity has opposite signs for 1.65 and 1.7 eV probes. (B) The transient reflection
spectrum at t = 0 (green circles). The transient reflectivity signal changes
sign at the A-exciton resonance energy EA = 1.68 eV, and it shows a
dispersive line shape. This transient reflection spectrum matches well with
the derivative of linear absorption spectrum in monolayer WSe2 and can be
fully accounted for by a 4-meV blueshift of the K valley A-exciton resonance
(magenta line).
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transitions in monolayer WSe2 governs both resonant and nonresonant excitations, which can have
different manifestations in valley physics.
The valley-dependent resonant excitation enables both the generation of a valley-polarized
exciton population through the absorption of
circularly polarized light and the detection of
valley-polarized excitons through polarizationresolved photoluminescence (PL) (46). Displayed
in Fig. 1B are polarization-resolved PL spectra of
our WSe2 samples upon laser excitation at 1.8 eV
with s+ circular polarization, in which the PL intensity with helicity matching the excitation light
(s+; Fig. 1B, red curve) is about four times stronger than that for the opposite helicity (s; Fig. 1B,
blue curve). This behavior is similar to that reported in previous studies (47) and verifies the
valley-selective excitation and emission in our
WSe2 monolayers. The nonzero s polarized PL
is presumably caused by a finite intervalley scattering because the excitation energy is ~120 meV
higher than the A-exciton resonance.
To manipulate the valley polarization, however, nonresonant coupling based on the optical
Stark effect is more advantageous because it
avoids the dissipation and dephasing naturally
accompanying real excitations. The optical Stark
effect is widely used in quantum optics to control
a rich variety of quantum systems, ranging from
the manipulation of cold atoms (21) and individual trapped ions (22) to coherent control of superconducting qubits (23) and electron spins (24). In
a two-level system, the optical Stark effect can be
readily understood by using the dressed-atom
picture (25). Consider the two-level system composed of the ground state and A-exciton state at
the K (or K) valley in a monolayer WSe2. Illustrated in Fig. 2A is the effect of a left circularly
polarized (s+) pump with photon energies below
the exciton resonance (Fig. 2A, red arrow). In the
dressed-atom picture, the dressed-ground state
(with N s+ photons), and the K valley A-exciton
(with N-1 s+ photons) are coupled by the dipole
transition, which leads to wave function hybridization and energy-level repulsion. It effectively
shifts down the ground-state energy and shifts
up the K valley exciton energy. The A-exciton at
the K valley, on the other hand, cannot couple to
the ground state with an extra s+ photon because
of the optical selection rule, and the related states
are not shifted by s+ polarized light. As a result,
the energy degeneracy between K and K valley
states is lifted by the valley-selective optical Stark
effect, which can be characterized by a valley
pseudo-magnetic field.
We measured the valley-selective optical Stark
shift in WSe2 monolayers with nonresonant
circularly polarized excitation using pump-probe
spectroscopy. Femtosecond pump pulses with
tunable photon energies below the A-exciton resonance were generated with an optical parametric amplifier. Optical Stark shifts of the WSe2
exciton transitions induced by the pump pulses
was then probed in transient reflection spectra
over the spectral range of 1.59 to 1.77 eV by using
a laser-generated supercontinuum light. Displayed
in Fig. 2, B and C, are two-dimensional plots of
transient reflection spectra in a monolayer WSe2
with s+ and s polarized probe light, respectively,
upon s+ polarized pump excitation. Here, the
nonresonant pump photons are at 1.53 eV, which
is 150 meV below the exciton resonance and do
not excite any real transitions. The color scale
in Fig. 2, B and C, represents the pump-induced
change of the probe beam reflectivity DR/R, the
horizontal axis shows the probe photon energy,
and the vertical axis shows the pump-probe time
delay. For atomically thin WSe2 layers on a transparent sapphire substrate, the reflection change
DR/R is directly proportional to the change in
the absorption coefficient (26, 27). It is apparent that strong changes in the exciton absorption
are present only for s+ probe pulses (Fig. 2B),
and no pump-induced signals can be detected
above the noise level for s probe pulses (Fig. 2C).
This indicates that the nonresonant s+ pump
Fig. 4. Pump detuning and pump intensity dependence of the valley-selective optical Stark
shift. (A) The transient reflection signal at 1.7 eV (left axis) and the corresponding optical Stark
shift (right axis) as a function of the pump detuning energy (green dots). The dependence can be
nicely described by an inverse proportional relationship (magenta line). (B) The transient reflection
signal at 1.7 eV (left axis) and the corresponding optical Stark shift (right axis) as a function of the
pump laser intensity (green dots), which shows a linear scaling behavior (magenta line). The optical
Stark shift can selectively shift the K valley exciton transition by as much as 10 meV, which
corresponds to a valley pseudo-magnetic field of ~60 T.
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line). Such scaling matches well with the theoretical prediction of optical Stark shift dw0
2SEp2 =W, where S is the optical Stark shift constant related to the transition dipole moment
and Ep is the electric field of the pump pulse
(25, 28). From the experimental data, we can determine an optical Stark shift constant S ~ 45 D2
for A-exciton in monolayer WSe2, which is of
similar magnitude to that for exciton transition
in semiconductor quantum wells (29).
The valley-selective optical Stark shift breaks
the degeneracy of valley exciton transitions in
monolayer WSe2 and defines an effective valley
pseudo-magnetic field. In our experiment, the
photo-induced energy splitting between K and
K exciton transitions can be as large as 10 meV
(Fig. 4 and supplementary text). The corresponding pseudo-magnetic field Beff for valley excitons can be
estimated with Beff
DE
2g m , where mB is the
ex B
1208
SUPPLEMENTARY MATERIALS
www.sciencemag.org/content/346/6214/1205/suppl/DC1
Materials and Methods
Supplementary Text
Figs. S1 and S2
References (3034)
30 June 2014; accepted 5 November 2014
10.1126/science.1258122
HEAVY FERMIONS
RE S EAR CH | R E P O R T S
t m0 ca Ha Hc m0 cc Hc Ha
m0 DcH 2 sinfcosf
In our experimental setup, an SmB6 single crystal is glued to the tip of a thin brass cantilever
(Fig. 1C, inset). The magnetic torque t is measured by tracking the capacitance change between the cantilever and a gold film underneath.
An example torque t versus magnetic field m0H
curve (Fig. 1C), taken at temperature T = 0.3 K
and f ~ 44, is quadratic overall, reflecting the
linear H dependence of the sample magnetization and the paramagnetic nature of SmB6.
Large oscillations and small wiggles start to
appear as the magnetic field goes beyond 5 T;
for m0H > 10 T, the fast oscillation patterns
dominate.
The first question that arises is whether it is
possible for the thin atomic layer of the surface
state to be responsible for the observed magnitude of the magnetic torque and magnetization. The magnitude of the observed magnetic
torque was recorded by the relative capacitance
change, then converted to absolute values using
the calibrated spring constant of the cantilevers
(see supplementary materials). Figure 1D shows
the field dependence of the effective magnetic
moment M m tH in a magnetic field H as high
0
(001) c
(101) plane
(100) a
(011) plane
(011) plane
Sm
B
(101) plane
(001) plane
Fig. 1. Experimental setup. (A) The crystal structure of Kondo insulator SmB6. The B6 cluster and rare
earth element Sm form a simple cubic structure. (B) Photograph of an SmB6 single crystal. Guide lines
are drawn to label the crystalline plane of each surface. (C) The field dependence of the magnetic torque
t in SmB6. The data were taken at temperature T = 300 mK. Inset: sketch of the measurement setup.
The sample stage is rotated to tilt magnetic field H in the crystalline a - c plane. The magnetic field was
applied to the sample with a tilt angle f ~44 relative to the crystalline c axis. The spring constant of the
cantilever was determined using the sample weight. (D) The field dependence of the magnetic moment
M, taken at 0.3 K and under a magnetic field as high as 45 T. The low-field linear MH dependence
reflects the paramagnetism in SmB6. The vertical gray scale bar shows the M value corresponding to
0.5 mB per surface unit cell.
SCIENCE sciencemag.org
A
2pe
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and F0b =cosf225 . We observe F b at frequencies as high as 900 Ta more than 200% increase
from the minimum value of 286 T. Such large
divergence and the close tracking of the inverse
cosine dependence strongly support the 2D nature of the observed b pocket on the (101) surface
plane families.
In contrast, the angular dependence of the
oscillation frequency of the a pocket F a follows
a different pattern. In Fig. 3A, the F a values are
plotted against the tilt angle f of two SmB6 single
crystals. The uncertainty of Fa is determined by
the half width at half-height of the F a peak in the
FFT plot. Similar to F b, the F a pattern has a
fourfold symmetry, but the minima are located
at f = 0, 90, and 180, that is, along the (100)
crystalline axes. Fitting the data to the 2D form
for this family, F0a =cosf0 , F0a =cosf90 ,
and F0a =cosf180 , with F0a = 30.5 T (solid
lines in Fig. 3A) results in reasonable agreement.
This suggests that the observed a pocket arises
from a surface state on the (100) plane families.
Similarly, Fig. 3C shows that the angular dependence of the g pocket Fg follows the functional form of F0g =cosf0 , F0g =cosf90 ,
and F0g =cosf180 with F0g ~385 T, suggesting the two-dimensionality of the g pocket on
the (100) surface plane.
Further, given the small value and large uncertainly of Fa, there is still a chance that an
extremely elongated 3D ellipsoidal Fermi surface
may fit the Fa versus f dependence. Experiments
with cleaner SmB6 crystals may resolve the issue.
The angular dependence of the oscillation frequency suggests that the Fermi surface b is twodimensional and likely arises from the crystalline
(101) plane. In contrast, most of the theoretical
modeling focuses on the surface states in the
Fig. 2. Quantum oscillation pattern observed by torque magnetometry. (A) The oscillatory magnetic torque tosc is plotted as a function of 1/m0H. tosc is calculated by subtracting a polynomial background from the full torque signal. (B) The Fast Fourier Transform (FFT) of the oscillatory torque tosc
shows three major peaks, labeled a, b, and g. The higher harmonics of the b peak are also observed,
together with the b peak from the neighboring surface plane. (C) The temperature dependence of the
FFTamplitude of the three major peaks, normalized by their 0 K limit.The magnetic field tilt angle f ~ 32.
Fitting the oscillatory amplitude to the LK formula (solid lines) yields the effective mass m = 0.119me for
the low-frequency a pocket, m = 0.129me for the high-frequency b pocket, and m = 0.192me for the
higher-frequency g pocket. (D) At 0.3 K and f ~ 30, the oscillation amplitude is tracked as a function of
field H, generally known as the Dingle plot. Solid lines are fits to the Dingle factor.
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F (T)
F (T)
(degree)
F (T)
(degree)
(degree)
Fig. 3. Angular dependence of the oscillating frequencies in SmB6. (A) Oscillation frequency Fa
of the observed Fermi surfaces a is shown as a function of f. The error bar of Fa is defined as the halfwidth at half-height of the peak in the FFT plot. To improve the statistics, two SmB6 samples were
cooled down twice. All the oscillation frequencies are inferred from the FFTspectra at many tilt angles f
from 30 to 180 (figs. S3 to S5) (37).The data are in agreement with a fit to the solid lines (described
by F0 =cosf f0 , where f0 = 0, 90, and 180). The lines correspond to 2D Fermi surfaces on the
surface planes (100), (001), and (1 0 0). (B) Oscillation frequency Fb of the observed Fermi surfaces
b measured in two SmB6 samples. Fb is consistent with the expected 2D Fermi surface sitting at the
four surface planes of the (101) surface family. The solid line is the fitting curve F 0 =cosf f0 , where
f0 = 45, 45, 135, and 225. (C) The angular dependence of the oscillation frequency Fg of the
pocket g. Solid lines are fits to F 0 =cosf f0 with f0 = 0, 90, and 180. (D) Sketch of the Brillouin
zone of SmB6. Band inversion occurs at the X points. The projection of these X points to the (101)
plane shows a possible location of the Fermi surfaces in the (101) surface state.
Table 1. Parameters in the three Fermi pockets in SmB6. The oscillation frequency F and Fermi
wavevector kF are obtained from the dHvA period. The effective mass m, Fermi velocity vF, mean free
path l, and the mobility m are inferred from the T and H dependences of the oscillation amplitude. The
Landau level index plot yields the infinite field limit d.
a
F (T)
Crystalline surface Origin
kF (nm1)
m/me
vF (105ms1)
l (nm)
m (103 cm2/Vs)
kFl
d
SCIENCE sciencemag.org
29.3 T 4
(1 0 0)
0.30 T 0.02
0.119 T 0.007
2.90 T 0.39
22 T 8
1.1 T 0.5
6.6 T 2.9
0.45 T 0.07
b
286 T 5
(1 0 1)
0.941 T 0.008
0.129 T 0.004
8.45 T 0.33
52 T 6
0.84 T 0.10
49 T 6
0.44 T0.06
g
385 T 5
(1 0 0)
1.080 T 0.011
0.192 T 0.005
6.50 T 0.21
25 T 4
0.36 T 0.05
27 T 4
0.32 T 0.07
(p= 2a,0). Based on the time-reversal symmetry, only (0, p/a) has a Dirac point, which is
consistent with the experimental observation.
However, the topological theory does not prohibit pairs of Dirac points at low symmetry points
(37), which offers another possible origin for the
observed pockets.
The other important question is whether
the 2D electronic state on the surface follows
the Dirac dispersion. A general test is to track the
Landau Level index plots to find out the infinite
field limit, i.e., the geometric Berry phase factor.
Using 45 T, the quantum limit is reached for
pocket a, which, in the infinite magnetic field
limit, points to 0.45 T 0.07, very close to 1/2,
the geometric Berry phase contribution, similar to other 2D Dirac electronic systems such as
graphene (18, 19). As the oscillation frequencies
of Fermi pocket b and g are quite close, filtering
is needed to isolate the H dependence of the
oscillation patterns for each Fermi pocket. This
filtering may cause additional uncertainty of
the oscillation phase, and the Zeeman effect and
correlations may lead to some nonlinear effect
of the Landau level index plots (37).
Because Al flux is used in the sample growth,
the observed effective masses, oscillation frequencies, and angular dependence bear some
similarity to those of pure aluminum (Al) (38).
However, the oscillation periods of all the observed Fermi surfaces are symmetric about crystalline symmetry axes in the rotation plane. The
observed fourfold symmetry and the behavior
F a ~ 1=cosf, F b ~1=cosf45 and F g ~ 1=cosf
cannot be explained by a residual Al impurity
[for more details, see (37)]. Therefore, the observed quantum oscillation pattern is an intrinsic property of single-crystalline SmB6.
We note also that our observed quantum oscillation feature is quite robust against oxidization, as the samples were always in atmosphere
for storage. The ordinary surface states known to
occur for vacuum clean surfaces of metal hexaborides such as LaB6 disappear under even modest oxygen exposure (37, 39).
REFERENCES AND NOTES
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www.sciencemag.org/content/346/6214/1208/suppl/DC1
Materials and Methods
Supplementary Text
Figures S1 to S14
Table S1
Equation S1
References (4050)
2 January 2014; accepted 5 November 2014
10.1126/science.1250366
1212
CHEMICAL KINETICS
Predictive a priori
pressure-dependent kinetics
Ahren W. Jasper,1 Kenley M. Pelzer,2 James A. Miller,2 Eugene Kamarchik,1
Lawrence B. Harding,2 Stephen J. Klippenstein2*
The ability to predict the pressure dependence of chemical reaction rates would be a
great boon to kinetic modeling of processes such as combustion and atmospheric
chemistry. This pressure dependence is intimately related to the rate of collision-induced
transitions in energy E and angular momentum J. We present a scheme for predicting this
pressure dependence based on coupling trajectory-based determinations of moments of
the E,J-resolved collisional transfer rates with the two-dimensional master equation. This
completely a priori procedure provides a means for proceeding beyond the empiricism of
prior work. The requisite microcanonical dissociation rates are obtained from ab initio
transition state theory. Predictions for the CH4 = CH3 + H and C2H3 = C2H2 + H reaction
systems are in excellent agreement with experiment.
ressure-dependent reactions are ubiquitous in applications of gas-phase chemical kinetics to practical problems, such
as combustion (<100 bar), atmospheric
chemistry (1 bar), and chemical vapor
deposition (<<1 bar). These reactions can take
various forms involving chemical or thermal activation, single or multiple unimolecular potential
wells, and single or multiple sets of bimolecular
products. Such reactions are enormously complicated problems to treat theoretically, even for
a single-channel, single-well dissociation, which
is the case we focus on here. The accurate firstprinciples prediction of pressure-dependent rate
coefficients would dramatically improve the utility
of theoretical kinetics as a tool for global chemical modeling.
There are two constants of motion in an isolated dissociating molecule: the total internal
(vibrational-rotational) energy, E, and the total
angular momentum, J. A physically correct description of the kinetics requires a quantitative
understanding of the rate of collision-induced
transitions in these two variables, kc(E,J;E',J'),
and the corresponding E- and J-resolved dissociation/isomerization rate constant, k(E,J). The
master equation treats the time dependence of
the state-resolved species populations arising from
collisions and dissociation. This time dependence
can be directly related to pressure-dependent
thermal dissociation rate constants k(T,p) (14).
Although many of the foundational formulations of reaction rate theory considered pressure
dependence, subsequent theoretical developments
have almost exclusively focused on pressureindependent reactions. Such pressure-independent
rate coefficients are generally obtained from a
thermal average of k(E,J) and are independent
of the collisional transition process. Many classes
of pressure-independent reactions can now be
1
Combustion Research Facility, Sandia National Laboratories,
MS 9055, Livermore, CA 94551-0969, USA. 2Chemical
Sciences and Engineering Division, Argonne National
Laboratory, Argonne, IL 60439, USA.
RE S EAR CH | R E P O R T S
amplified, particularly for higher-energy channels and well-skipping reactions that involve
dynamical passage over intermediate wells with
only partial stabilization (15).
The most commonly used model (1, 3) for the
energy transfer process neglects the dependence
of the kinetics on the total angular momentum J
and employs a so-called exponential down form
for the probability of a collision-induced transition
from E' to E (16):
DE
0
1
; E < E0
PE; E exp
a
In Eq. 1, DE = E E', and a is a fitting parameter, which represents the average downwards energy transferred, <DEd>, in a single
deactivating collision. The energy transfer rate
kc(E;E') is then generally represented as the
product of a Lennard-Jones collision rate, ZLJ,
with P(E;E'). Within this one-dimensional master equation (1DME) model, k(T,p) may be obtained via empirical estimates for the parameter a.
An obvious shortcoming of this simple approach is its neglect of J. Explicitly including J
requires the solution of the two-dimensional
(in E and J) master equation (2DME) and some
characterization of the four-dimensional function kc(E,J;E',J'), which we also write as the product of a total collision rate, Z, and a normalized
probability, P(E, J;E',J'), of a collision producing
the state (E, J) from an initial state (E', J'). The
2DME reduces to a 1DME if one assumes the
distributions in E and J are separable and then
makes a microcanonical strong collider in J assumption (i.e., the final rotational state is assumed
to be independent of the initial rotational state
with its probability determined by the J dependence of the microcanonical density of states)
(17, 18). In this 2D/ model, we write P(E,J;E',J')
as P(E;E') (E,J), where (E,J) = r(E,J)/r(E), with
r(E,J) and r(E) as the E,J- and E-resolved density
of states for the molecular complex, respectively.
A priori kinetic predictions were recently obtained within this 2D/ model for CH4 dissociation via classical trajectory calculations of the
parameter a = <DEd> (19). However, detailed
comparisons with experimental data indicate
that such predictions are not highly accurate,
being about a factor of 2 too high, at least once
anharmonic effects on the vibrational state densities are accounted for (19).
The assumption in the 2D/ model that single
collisions produce microcanonical statistical distributions of J is clearly not valid; kinetically
relevant distributions of J typically are much
wider than the average J-transferred in a single
collision (12) (e.g., by a factor of 5 or more; compare fig. S2, a and b). One might also question the
use of classical mechanics in the estimation of the
energy transfer rates. However, the Ehrenfest
theorem states that averaged quantum mechanical variables increasingly obey classical laws as
the classical (high-E) limit is approached. The
sensitivity of kinetic predictions to collisional
transfer rates only at energies near the dissociation threshold then suggests that quantum
effects on these rates should be negligible (20).
SCIENCE sciencemag.org
dJ dE PE; J; E ; J DE m D J n
0
E0
DE m DJ n dJ hJ DE m DJ n E0 , J 3
0
where h(J') are reference (e.g., thermal) J' distributions, and E'0 is the rotationless threshold
Fig. 1. Plot of the pressure-dependent
rate constant for CH3 + H (+He) CH4
(+He) at 600 K for various approximate
models: the simple separable twoparameter 2DME model as a function
of the DJ range parameter g, the
one-parameter 2D/ model, and the
one-parameter 1DME model. Symbols
denote experimental results and error
bars from Brouard et al. (29).
1213
R ES E A RC H | R E PO R TS
energy for dissociation. We have explicitly neglected to characterize the E' dependence in P
because the solution to the 2DME depends only
on E' values near the dissociation threshold,
where the dependence of P on E' is very weak.
These low-order moments may be readily evaluated with a limited number of trajectories (e.g.,
103), which facilitates routine implementation
of the approach and even suggests the feasibility of performing direct trajectory evaluations.
To solve the master equation, we use an ad
hoc analytic representation of the energy transfer function that is simple, yet flexible enough to
reproduce the low-order double moments defined in Eq. 3. Details are given in the supplementary material. Briefly, the functional form
allows for the modeling of weak collisions in J,
the nonseparability of DE and DJ, long tail (nonexponential) collisions in both DE and DJ, and an
explicit dependence on J'. The analytic functional
form has nine parameters that are determined
such that the analytic representation reproduces 135 low-order double moments of the energy
transfer function (Eq. 3) calculated using classical trajectories.
Barker and Weston have recently explored
the mapping of the four-dimensional collisioninduced transition kernel with trajectory simuFig. 3. Plot of the 1DME, 2D/f, and full
2DME predictions for the pressuredependent rate constant for H +
C2H2 C2H3 at room temperature
with He as a bath gas. Symbols denote
experimental results and error bars from
Keil et al. (30) and Payne and Stief (31).
1214
lations in a study of the energy deactivation process for excited pyrazine and ethane (23). These
and earlier trajectory studies cited therein elucidated important details of P, but these details
have not found their way into kinetics calculations. Some prior master equation studies have
implemented the full 2DME (2426), but these
studies used empirical separable models owing
to the absence of data for the collision kernel.
The details of the present theoretical calculations are provided in the supplementary material, where the (i) implementation of the 2DME,
(ii) ab initio transition state theory calculations,
(iii) representation and fitting of the collisional
energy transfer kernel, and (iv) classical trajectory simulations are each discussed at length.
These descriptions include an illustration of the
quality of the present moment-fitting procedure
in fig. S2 and in table S2.
As a demonstration of our approach, we obtained a priori predictions for three representative classes of reaction, focusing on specific
reactions for which the experimental record appears fairly definitive. In particular, we studied
CH3 + H radical-radical recombination, H + C2H2
radical-molecule recombination, and C2H3 radical decomposition. Each of these reactions has
also been the subject of previous theoretical
studies by our team where the pressure dependence was treated empirically via adjustments
of a to reproduce the experimental k(T,p) using
the 2D/ model (18, 27, 28). Although the present
comparisons serve as a useful test of the approach, its true power will be in making predictions of rate coefficients for cases where there
are little or no experimental data. In this regard,
it is important to emphasize that the present approach is readily applicable to essentially arbitrary reactions and bath gases.
The pressure dependence of the CH3 + H recombination rate coefficient was considered at
four temperatures ranging from 300 to 600 K.
The predicted rate coefficients (Fig. 2) are seen to
be in excellent agreement with the experimental
data of Brouard et al. (29), generally within the
error bars. Although there are a few discordant
points at the lowest pressures for the two lowest
temperatures, there are some apparent irregularities in the experimental data in that range.
For example, the value near p = 25 Torr for 300 K
is below that for 400 K, which is difficult to
rationalize theoretically. Aside from that irregularity, theory and experiment generally agree to
within 20%.
The pressure dependence of the room temperature rate coefficient for the H + C2H2 radicalmolecule recombination to form C2H3 has been
studied experimentally over an exceptionally
broad range of pressures (from 1 to 1000 Torr)
by Keil et al. (30) and by Payne and Stief (31).
Meanwhile, the related reverse dissociation of
vinyl was studied by Knyazev and Slagle (32) for
a range of pressures at temperatures near 1000 K.
Taken together, the two sets of data provide a
stringent test for the present a priori approach.
As shown in Figs. 3 and 4, the predictions for
both the low-temperature recombination rate
constants and the high-temperature dissociation
rate constants are in very good agreement with
the experimental data, with maximum deviations
of about 20%.
In summary, the present pressure-dependent
kinetic analysis solves the two-dimensional master equation for the representation of the collisional E,J transition kernel specified by eqs. S1
to S5, with the parameters in the representation
obtained from fits to trajectory-based determinations of its low-order moments (Eq. 2). The
requisite (E,J)-resolved dissociation rate constants are obtained from ab initio transition
state theory calculations. The excellent performance of this fully a priori J-resolved pressuredependent kinetic analysis for these three test
cases strongly suggests that the present representation of the collision kernel (eq. S3) is accurate enough for obtaining kinetic estimates and
that the trajectory-based prediction of the underlying parameters is also quantitatively accurate. Thus, this method should prove highly
effective for predicting the pressure dependence
of experimentally unstudied reactions and will
serve as a great boon to the development of
chemical models for complex gas phase environments. It should also allow for the quantitative
prediction of the bath gas dependence of the rate
sciencemag.org SCIENCE
RE S EAR CH | R E P O R T S
www.sciencemag.org/content/346/6214/1212/suppl/DC1
Materials and Methods
Figs. S1 and S2
Tables S1 and S2
References (3545)
5 September 2014; accepted 28 October 2014
10.1126/science.1260856
SURFACE STRUCTURE
RE FE RENCES AND N OT ES
SCIENCE sciencemag.org
Iron oxides play an increasingly prominent role in heterogeneous catalysis, hydrogen production,
spintronics, and drug delivery. The surface or material interface can be performance-limiting
in these applications, so it is vital to determine accurate atomic-scale structures for iron oxides
and understand why they form. Using a combination of quantitative low-energy electron
diffraction, scanning tunneling microscopy, and density functional theory calculations, we show
that
ordered array of subsurface iron vacancies and interstitials underlies the well-known
p an
p
( 2 2)R45 reconstruction of Fe3O4(001). This hitherto unobserved stabilization mechanism
occurs because the iron oxides prefer to redistribute cations in the lattice in response to oxidizing
or reducing environments. Many other metal oxides also achieve stoichiometry variation in
this way, so such surface structures are likely commonplace.
The surface VO concentration is usually in equilibrium with the bulk, where such defects form
when the oxide is reduced. At the surface, VOs
strongly affect the electronic structure, are active
sites for chemical reactions (4, 5), and play a
central role in surface reconstructions. There
exists another class of metal oxides, however,
in which stoichiometric variation is accommodated primarily via the cations; the iron oxides
are the prototypes. Wstite (FeO), magnetite
(Fe3O4), and hematite (a-Fe2O3) are all based
on a close-packed O2 anion lattice and differ
primarily in the number, oxidation state, and
distribution of the cations among octahedral
and tetrahedral interstitial sites, Feoct and Fetet
(6). Near the phase boundaries, FeO can incorporate up to 17% of Feoct vacancies, whereas the
metastable phase maghemite (g-Fe2O3) is essentially Fe3O4 with 1/6 of the Feoct removed.
We now show that an ordered array of subsurface cation vacancies and interstitials underlies
5 DECEMBER 2014 VOL 346 ISSUE 6214
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R ES E A RC H | R E PO R TS
p p
the well-known ( 2 2)R45 reconstruction of
Fe3O4(001). This stabilization is driven by the
material adopting a stoichiometry compatible
with its environment. The oxides of Co, Mn, and
Ni exhibit similar cation redistributions in reducing and oxidizing environments, so such reconstructions may be a common occurrence.
In the (001) direction, Fe3O4 consists of alternating planes of Fetet and (Feoct)2O4 and thus
appears to be a classic Tasker typep
3polar
psurface
1216
character that is also observed in x-ray photoelectron spectroscopy (XPS) (9); angle-resolved
XPS spectra are provided in fig. S1 (17). The Feoct
and Fetet sublattices remain antiferromagnetically
coupled as in the bulk, yielding a reduced net
magnetic moment in the SCV reconstruction. The
absence of Fe2+ suggests that the contribution
of the orbital moment should be small. The magnetic moments of all atoms in the surface region
are contained in table S1 (17) together with their
optimized DFT+U coordinates. The energetic
preference for the SCV structure is borne out by
atomistic thermodynamics calculations (Fig. 1E)
(18). The distorted bulk truncation is only favored
below an oxygen chemical potential of 3 eV, corresponding to an O2 partial pressure of <1020 mbar
at 900 K. Under such reducing conditions, however, Fe-rich surface phases occur (19). Although
a-Fe2O3 is predicted to take over as the thermo9
dynamically
p p stable phase above 10 mbar, the
( 2 2)R45 reconstruction persists up to
105 mbar. It has been shown that a-Fe2O3 inclusions grow slowly at 106 mbar O2 (20), whereas
conversion of the surface to g-Fe2O3(001) requires
extremely oxidizing conditions (9).
A key success of the SCV structure is that it explains the extraordinary thermal stability of Au,
Ag, and Pd adatoms at the Fe3O4(001) surface,
which resist agglomeration into clusters up to
sciencemag.org SCIENCE
RE S EAR CH | R E P O R T S
surface reveal no clear preference for one specific site. On the SCV structure, the interstitial
Feint blocks the adsorption of an adatom in the
bulk continuation site directly above. For Au, a
Fig. 2. Quantitative low-energy electron diffraction measurements unambiguously confirm
the subsurface cation vacancy termination of
Fe3O4(001). (A) Comparison of selected experimental LEED IV spectra and theoretical curves for
the optimized subsurface vacancy structure. The
final RP for the best-fit structure is 0.125; the selected beams have an individual RP close to this
average. (B and C) Experimental LEED patterns
for electron energies of 149 and 285 eV, respectively. Diffraction spots highlighted with yellow circles
correspond topthe
in (A). The red square
curves
p
indicates the ( 2 2)R45 unit cell.
SCIENCE sciencemag.org
1217
R ES E A RC H | R E PO R TS
www.sciencemag.org/content/346/6214/1215/suppl/DC1
Materials and Methods
Figs. S1 to S3
Tables S1 to S3
References (3335)
Database S1
28 August 2014; accepted 17 October 2014
10.1126/science.1260556
sciencemag.org SCIENCE
RE S EAR CH | R E P O R T S
HUMAN CAPITAL
very day in the United States, acts of violence kill almost 150 people and injure over
6000 more (1, 2). This public health crisis
(3) disproportionately involves youth, who
are twice as likely as adults to be both victims and perpetrators of violence (4). The problem is most concentrated among disadvantaged
minority youth; violent-crime arrest rates for
African-American juveniles are five times as high
as that of their white counterparts (5).
Poverty scholar W. J. Wilson identifies one major
cause of these racial violence disparities among
young people: joblessness (6). His argument adds
to decades of social science investigating how poor
job prospects cause crime, from weakening social
bonds to generating psychic strain to reducing
the perceived cost of punishment (710). Policy
discussions often conclude that public employment and training programs are a solution to
youth violence on the grounds that nothing stops
a bullet like a job (11).
The empirical literature on youth employment
programs, however, suggests that the practical applications of this idea are limited (12). Several major
employment programs fail to reduce delinquency
among youth (13, 14). The two experimentally evaluated interventions that do lower crime, at least
during the program, involve such intensity and expense that their benefits fail to outweigh their costs
(15, 16). Attempts to provide shorter, more scalable
employment services have, if anything, increased
adolescents criminal behavior (17). The theoretical
promise of using employment interventions to fight
youth violence does not appear to translate easily
into empirical reality; only very high levels of investment seem able to reduce crime (supplementary
materials, section 2.1) (18).
But, one widespread type of youth employment program has not yet been rigorously evaluated: summer jobs (19, 20). Despite some
promising observational studies (21), there is little convincing causal evidence on the effects of
1
Department of Criminology, University of Pennsylvania,
Philadelphia, PA, USA. 2University of Chicago Crime Lab,
Chicago, IL, USA.
SCIENCE sciencemag.org
rather than managing or trying to reverse a problem once it occurscan improve outcomes more
effectively, with less intensive treatment (2325)
If this well-established idea that prevention can
be more effective than remediation also applies to
the employment domain, teaching adolescents how
to be successful employees, facilitating connections
to employer networks, and providing work experience before they drop out of school might reduce
crime with less intensive intervention than is required for already disconnected youth. Offering
summer employment at this key point in the life
course could make crime a relatively less attractive option, strengthen social bonds, and develop
soft skills such as self-efficacy and impulse control
(7, 2628). Waiting until after dropout to intervene
may make it more difficult to reduce violence in
particular, given the apparently causal relationship
between school dropout and criminal behavior, especially murder and assault (29). Limiting the intervention to summer avoids direct conflicts
between work and school. Summer jobs also provide wages and structured activity during a highcrime season when youth might otherwise be idle,
both of which may affect crime (30, 31).
However, the theory on summer jobs is not entirely clear-cut. Additional income could be spent
on crime-inducing goods such as drugs and alcohol, and time spent traveling to and from work
might increase exposure to criminal opportunities.
Even the attempt to keep youth busy (the incapacitation effect) may be poorly targeted because
most jobs are during business hours, whereas most
crime occurs during evenings and weekends. More
broadly, a short-term subsidized job might not be
enough to generate behavior changes beyond the
summer itself. The effects of summer jobs are
therefore an empirical question.
Data come from matching study youth to
administrative data sources (supplementary materials, materials and methods). Program participation is from provider-tracked attendance
records. Student-level administrative records from
the Chicago Public Schools capture pre- and postprogram academic outcomes from the 20112012
and 20122013 academic years, respectively. Demographic information on applicants neighborhoods comes from matching the Census tract of
youths home addresses to the 2010 American Community Survey. The main outcome measures are
from individual-level Chicago Police Department
arrest records covering both juveniles and adults
(32). The study uses data through 25 September
2013, which is 16 months after randomization
(13 months after the end of the program).
The analysis categorizes arrests by offense type
(violent, property, drug, and other). It is common
for social interventions to have differential impacts
on these different types of crime (31, 33, 34), likely
because some of the underlying causes differ.
Violent crime stems from conflicts between people, so problematic cognitive and emotional responses to social interactionsincluding hostile
attribution bias, uncontrolled anger, and hot
decision-makingare thought to be proximal
causes of youth violence in particular (35, 36).
Nonviolent crimes, which involve property or
5 DECEMBER 2014 VOL 346 ISSUE 6214
1219
R ES E A RC H | R E PO R TS
1220
Table 1. Mean preprogram characteristics for treatment and control groups. To test baseline
equivalence, each characteristic was regressed on treatment indicator and blocking variables using
heteroskedasticity-robust standard errors; the P value column reports statistical significance of
treatment indicators coefficient. Demographic and schooling data are from Chicago Public Schools
administrative records on pre-program year (20112012 school year). Free/reduced price lunch is
proxy for family poverty. Days absent are reported as a percentage of days enrolled (average missed,
29 days). Arrest and victimization data are from Chicago Police Department administrative records.
Neighborhood characteristics are from 2010 American Community Survey and Chicago Police
Departments community-area crime rates. Percent unemployed is percent of civilian labor force over
age 16 years looking for work but without a job. Table uses nonmissing data only (5 youth missing school
attendance data, 60 missing fall GPA, 1 missing neighborhood violent-crime rate). Gender is not included
in the table because it is captured by the blocking variables; 38% of the sample is male.
Control mean
Treatment mean
(n = 904)
(n = 730)
16.79
10.15
96%
2.9%
92%
16.78
10.12
94%
3.8%
92%
Demographics
Age
Grade
Percent Black
Percent Hispanic
Percent free/reduced price lunch
P value
0.46
0.55
0.84
0.74
0.98
Schooling
Percent days absent
GPA in fall 2011 (4 point scale)
Percent enrolled in summer school 2011
18%
2.37
8.8%
18%
2.32
9.0%
0.99
0.93
1.00
19%
0.13
0.09
0.05
0.15
21%
0.29
22%
0.18
0.09
0.08
0.19
24%
0.33
0.37
0.20
0.71
0.55
0.55
0.36
0.30
19%
33%
35,665
2,128
19%
35%
34,321
2,136
0.94
0.71
0.58
0.95
Neighborhood characteristics
Percent unemployed
Percent below poverty line
Median household income (US$)
Violent crime rate (per 100,000)
.12
Treatment mean
.1
0.123 0.122
0.091
0.051
0.051
0.049
0.054
0.039
Control mean
Property
Drug
Other
.04
Violent
0.0395**
(43%)
.04
0.0124
0.0054
0.0004
.08
Difference (ITT)
95% confidence interval
sciencemag.org SCIENCE
RE S EAR CH | R E P O R T S
to engage with less intensive pro-social programming (40). If so, OSPs SEL curriculumwhich
focuses on emotion and conflict management,
social information processing, and goal setting
could be a key driver of the violence decrease. Because SEL was randomly assigned, the study can
separately identify any additional role of SEL.
ITT violence effects by treatment group are
shown in Table 2. Not only is the difference between groups statistically insignificant, but the
magnitude is also tiny (1% of the control mean).
The nearly identical point estimates suggest that
the statistical similarity across groups is not just
the result of limited power to detect subgroup
differences. Instead, it appears that both groups of
youth experienced very similar drops in violence. The same is true for other crime types (supplementary materials, section 2.6, and table S4).
Although the experimental design can only directly isolate the role of SEL, another potentially
important mechanism is testable indirectly: whether the decrease in violence is a mechanical result of
youth having less time to engage in crime while
working over the summer (an incapacitation effect). If so, and youth returned to their prior behavioral patterns immediately after the program
0.0391*
(0.0205)
Jobs-only
0.0399**
(0.0203)
Jobs + SEL
0.0008
(0.0220)
SCIENCE sciencemag.org
End of program
5 6 7 8 9 10 11 12 13 14 15 16
Months since random assignment
1221
R ES E A RC H | R E PO R TS
hugely costly to improve outcomes for disadvantaged youth; well-targeted, low-cost employment policies can make a substantial difference,
even for a problem as destructive and complex
as youth violence.
RE FERENCES AND NOTES
26.
27.
28.
29.
30.
31.
32.
33.
34.
35.
36.
37.
38.
39.
40.
41.
42.
43.
44.
45.
46.
47.
48.
49.
50.
51.
52.
sciencemag.org SCIENCE
RE S EAR CH | R E P O R T S
Youth and Family Services, and Youth Guidance. I thank Chicago Public
Schools partners J. Foreman, K. Klein, J. Loudon, and S. Norris;
Chief R. Tracy, T. Lavery, and the Chicago Police Department; R. Goerge
and Chapin Hall; R. Ander, S. Coussens, J. Davis, G. Cusick, M. Egan,
H. Pollack (Chicago PI), R. Harris, N. Hess, A. Mtivier, and J. Rountree
for project support; and K. Charles, J. Guryan, C. Loeffler, J. Ludwig,
J. MacDonald, E. Owens, S. Raudenbush, S. Sotelo, and D. Tannenbaum
for helpful comments. Replication data are posted at the University of
Michigans ICPSR data depository (http://doi.org/10.3886/E18627V1);
see supplementary materials section 1.5 for details. All content is
the responsibility of the author and does not represent the official
position or policies of the Chicago Police Department, Chicago Public
PALEOCLIMATE
he future response of African rainfall to increasing greenhouse gas (GHG) concentrations is a critical socio-economic issue, with
implications for water resources, agriculture,
and potential conflict (1), but uncertainties
among model projections remain (24). African
hydroclimate changed substantially during the
last deglaciation, the most recent time period
during which natural global warming was associated with increases in GHG concentrations.
Numerous proxy records from Africa indicate
that dry conditions during the Last Glacial Max1
Climate and Global Dynamics Division, National Center for
Atmospheric Research, Boulder, CO 80307-3000, USA.
2
Department of Earth, Environmental, and Planetary
Sciences, Brown University, Providence, RI 02912, USA.
3
College of Earth, Ocean, and Atmospheric Sciences, Oregon
State University, Corvallis, OR 97331, USA. 4Center for
Climatic Research and Department of Atmospheric and
Oceanic Sciences, University of Wisconsin-Madison, Madison,
WI 53706, USA. 5Laboratory for Climate, Ocean and
Atmosphere Studies, School of Physics, Peking University,
Beijing 100871, P. R. China. 6Department of Geosciences and
Institute of the Environment, University of Arizona, Tucson,
AZ 85721, USA. 7Cooperative Institute for Research in
Environmental Sciences, University of Colorado Boulder,
Boulder, CO 80309, USA. 8Department of Earth and
Environmental Sciences, Columbia University, New York, NY 10027,
USA. 9Department of Earth, Ocean, and Atmospheric Science,
Florida State University, Tallahassee, FL 32306, USA.
SCIENCE sciencemag.org
www.sciencemag.org/content/346/6214/1219/suppl/DC1
Materials and Methods
Supplementary Text
Tables S1 to S8
References (5396)
23 June 2014; accepted 7 November 2014
10.1126/science.1257809
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R ES E A RC H | R E PO R TS
18
17
16
15
14
13
12
11
18
0.05
14
0.06
10
0.07
0.08
0.09
0.1
400
0.5
0.9
200
4
40
45
50
55
60
Humidity Index
0.3
600
NA Precip
(mm/year)
Th
19
230
20
Pa/
21
Terrigenous
AMOC
(Sv)
22
NA TLAI
130
110
1000
90
70
0.8
0.6
0.4
BIT index
1
1200
Dwax
231
SEA Precip
(mm/year)
closely parallels changes in precipitation, starting from low values signifying primarily bare
ground followed by increasing values starting
at ~16 ka (Fig. 1C). To the extent that precipitation and vegetation cover influence availability
of sediment for aeolian transport, the temporal
evolution of these simulated properties can be
used to explain a record of terrigenous dust flux
sourced from subtropical NA (8) (Fig. 1C).
Simulated deglacial changes in SEA hydrology
are similar to those simulated for NA, including a near-synchronous start and subsequent
persistence of the AHP. After peak dry conditions at ~17 ka, TraCE simulates a gradual increase in precipitation starting at ~16.5 ka and
a large, abrupt increase at ~14.7 ka associated
with the start of the AHP (Fig. 1D). These changes
are observed in two SEA proxy records from
Lakes Tanganyika (16) and Challa (14) (Fig. 1D),
which show an abrupt onset of the AHP synchronous within a few hundred years of the
precipitation increase simulated by the model.
Rainfall changes in the Congo basin of western
equatorial Africa, which integrate precipitation
changes north and south of the equator, indicate drying from LGM to 17 ka and then increasing precipitation in both TraCE and proxy data
(25) (Fig. 1E).
We use empirical orthogonal functions (EOFs)
and their associated principal components (PCs)
to extract the dominant modes of precipitation
variability from the TraCE simulation and mois-
0.2
2200
160
2100
150
140
2000
Dwax
800
130
1900
120
22
21
20
19
18
17
16
15
14
13
12
11
Age (ka)
1224
sciencemag.org SCIENCE
RE S EAR CH | R E P O R T S
Fig. 2. EOFs and associated PCs for the deglacial period 20 to 11 ka. EOF1 and PC1 of (A) proxy data
for moisture availability and (B) TraCE annual precipitation (millimeters per year). (C and D) Same as (A) and
(B) except for EOF2 and PC2. Model results and proxy records (table S1) are interpolated to the same
100-year resolution.
SCIENCE sciencemag.org
1225
R ES E A RC H | R E PO R TS
30oN
10.6
7.6
4.7, 5.7
30 N
2.8
15oN
15oN
1.7
2.1
0o
0o
1.7, 3.2
1.3, 0.8
-0.1
15oS
15oS
o
30oS
30 S
o
45 W
45 E
90 E
45oW
135 E
-1
0o
-0.5
-0.1
45oE
90oE
135oE
30 N
Fig. 4. Deglacial annual SST change (C), 11 ka minus 17 ka, as simulated by the model and from proxy records. (A) TraCE, (B) TraCE GHGonly, and (C) TraCE orbital-only. Proxy records plotted in (A) are tabulated
in table S2.
15oN
0o
15oS
o
30 S
45oW
1226
0o
45oE
90oE
135oE
sciencemag.org SCIENCE
RE S EAR CH | R E P O R T S
of this century (3). Although multiple forcings influenced African climate during the deglaciation,
the agreement between proxy records and our
simulated evolution of rainfall indicates that the
processes and sensitivity of GHG-driven rainfall
change are well simulated in our fully coupled
global climate model. This lends confidence to
the CMIP5 climate model ensemble projections
of future precipitation change over tropical Africa
at the continental scale.
SUPPLEMENTARY MATERIALS
www.sciencemag.org/content/346/6214/1223/suppl/DC1
Supplementary Text
Fig. S1 to S8
Tables S1 and S2
References (3372)
4 August 2014; accepted 6 November 2014
10.1126/science.1259531
OCEANOGRAPHY
RE FE RENCES AND N OT ES
SCIENCE sciencemag.org
Multidecadal warming of
Antarctic waters
Sunke Schmidtko,1,2* Karen J. Heywood,1 Andrew F. Thompson,3 Shigeru Aoki4
Decadal trends in the properties of seawater adjacent to Antarctica are poorly known, and the
mechanisms responsible for such changes are uncertain. Antarctic ice sheet mass loss is
largely driven by ice shelf basal melt, which is influenced by ocean-ice interactions and has
been correlated with Antarctic Continental Shelf Bottom Water (ASBW) temperature. We
document the spatial distribution of long-term large-scale trends in temperature, salinity, and
core depth over the Antarctic continental shelf and slope. Warming at the seabed in the
Bellingshausen and Amundsen seas is linked to increased heat content and to a shoaling of
the mid-depth temperature maximum over the continental slope, allowing warmer, saltier water
greater access to the shelf in recent years. Regions of ASBW warming are those exhibiting
increased ice shelf melt.
he Antarctic ice sheet is the largest reservoir of terrestrial ice and is a significant
contributor to sea level rise in a warming
climate (1). Massive ice shelf disintegration
and rapid acceleration of glacial flow have
occurred in recent decades (2) or are potentially
looming (3). These events are generally linked to
enhanced basal melt (4, 5), which reduces buttressing and accelerates glacier flow. An increase
in basal melt may be linked to stronger subice
shelf circulation of Circumpolar Deep Water
(CDW). It is not known whether changes in the
delivery of warm water to the underside of the
ice shelf are caused by increased heat content,
increased volume flux responding to changes
in wind and buoyancy forcing, or some combination of the two (6, 7). Here, we focus on documenting long-term temperature and salinity
changes in ocean properties over the continental
shelf and slopes.
We refer to the water occupying the sea floor
on the Antarctic continental shelf as Antarctic
Continental Shelf Bottom Water (ASBW) and
the temperature minimum layer, representing the remnant of the winter mixed layer, as
Winter Water (WW). Atmospheric processes,
adjacent water masses, ice shelf and continental
1
Centre for Ocean and Atmospheric Sciences, School of
Environmental Sciences, University of East Anglia, Norwich
NR4 7TJ, UK. 2GEOMAR Helmholtz Centre for Ocean
Research Kiel, Dsternbrooker Weg 24, 24105 Kiel, Germany.
3
Environmental Science and Engineering, California Institute
of Technology, Pasadena, CA 91125, USA. 4Institute of
Low Temperature Science, Hokkaido University, Sapporo
060-0819, Japan.
1227
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Fig. 1. Temporal means, linear trends, and multiannual variability in Antarctic Continental Shelf
Bottom Water (ASBW). (A to D) Conservative temperature [(A) and (C)] and absolute salinity [(B) and
(D)] at the seabed for depths shallower than 1500 m for the period 1975 to 2012 are shown in terms of
temporal means [(A) and (B)] and linear trends [(C) and (D)].Trends not significantly different from zero are
hatched. (E and F) Multiannual variability as shown by 5-year median properties since 1975, interquartile
ranges, and median trends for selected areas. Abbreviations for surrounding seas: BS, Bellingshausen Sea;
AS, Amundsen Sea; RS, Ross Sea; CS, Cosmonaut Sea; WS, Weddell Sea (excluding the Antarctic Peninsula).
1228
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RE S EAR CH | R E P O R T S
The trends in Figs. 1 and 2 show a link between changes in the properties of ASBW and
those of CDW over the continental slope. The
CDW thermal structure may be broadly categorized into two regimes, shown schematically
in Fig. 4: CDW (i) sloping upward or (ii) sloping
downward toward the shelf break. The Weddell
and Ross gyres are subject to a large-scale cy-
Fig. 2. Temporal means and linear trends in Circumpolar Deep Water (CDW). (A to F) Conservative temperature [(A) and (D)], absolute salinity [(B) and
(E)], and core depth [(C) and (F)] of CDW between 1975 and 2012 are shown in terms of temporal means [(A) to (C)] and linear trends [(D) to (F)]. Trends not
significantly different from zero are hatched. Fronts (28) related to the ACC are shown as black lines. Only regions where the ocean depth exceeds 1500 m and the
underlying CDW is cooler than 2.8C are shown. (G) Trends and data for selected locations.
34.4
0.5
1
34.2
WW Sabs (g kg1)
WW (C)
1.5
150
100
WW P (dbar)
200
0.5
50
5
10
0
5
(dbar yr
yr )
10
WW Pt
30
10
WW S t (mg kg
30
WW t1 (mC yr1)
60
10
60
Fig. 3. Temporal means and linear trends in Winter Water (WW). (A to F) Conservative temperature [(A) and (D)], absolute salinity [(B) and (E)], and core
depth [(C) and (F)] of WW between 1975 and 2012 are shown in terms of temporal means [(A) to (C)] and linear trends [(D) to (F)]. Trends not significantly
different from zero are hatched. Fronts (28) related to the ACC are shown as black lines. Only regions where the ocean depth exceeds 1500 m and the
underlying CDW is cooler than 2.8C are shown.
SCIENCE sciencemag.org
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R ES E A RC H | R E PO R TS
Fig. 4. Schematic differences of shelf water masses. (A) Schematic of the Amundsen and Bellingshausen
seas with warm water at the bottom of the water column on the shelf. (B) Schematic of the shelf in the Ross
and Weddell seas, with a strong Antarctic Slope Current.
1230
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RE S EAR CH | R E P O R T S
SUPPLEMENTARY MATERIALS
ACKN OWLED GMEN TS
ANIMAL PHYSIOLOGY
to 600 V (1). Early attempts to understand electricity made use of electric eels (2), and more
recently, eels were important for identifying acetylcholine receptors (3) and for providing insights
into the evolution of electric organs (4), but little
is known about how the eels electrical discharge
affects prey. In this study, I designed a set of expe-
www.sciencemag.org/content/346/6214/1227/suppl/DC1
Materials and Methods
Figs. S1 to S11
Table S1
References (29, 30)
15 May 2014; accepted 27 October 2014
10.1126/science.1256117
43 cm/s
20
40
60
80
100
120
140
160
180
200
220
Milliseconds
40ms
160ms
2 cm
0 ms
20 ms
40 ms
60 ms
80 ms
15 cm/s
Shocked
100 ms
120 ms
140 ms
160 ms
180 ms
160ms
Matched
Unshocked
55 cm/s
Fig. 1. Eels discharge and strike. (A) Electric organ discharge corresponding to plates below. Arrow indicates low-amplitude discharge. (B) Video frames
showing that fish movement is arrested by discharge. Red frames indicate electric organ discharge (movie S1). (C) The utility of the discharge illustrated. Shown
are the prey fish at 40 ms (green) and later, the position and velocity of the eel and fish at 160 ms (red fish). Green dotted fish outline shows velocity and location of
uninterrupted escaping fish matched in time, size, and position from 40 ms, suggesting that the eel would have missed without the discharge.
SCIENCE sciencemag.org
1231
R ES E A RC H | R E PO R TS
Force
Transducer
Agar Barrier
Force
Transducer
20 ms
3.7 ms
3 45 6 7 8 9
Fish
Tension
Eel
EOD
Pithed
Fish
Eel
EOD
10 ms
23 4 5 6 7 8 9
100 ms
Fish Tension
Time 2:40
3:14
6:29
9:16
12:04
14:55
17:58
21:03
23:10
EOD
253g
Fish 1
Tension
SHAM
429g
CURARE
Fish 2
Tension
Fig. 2. Paradigm for investigating strong electric organ discharge. (A) An agar barrier separated eels from pithed fish. Eels shocked earthworms
while fish tension was recorded. (B) All eels induced whole-body tension, occurring 2 to 4 ms after strong discharge onset. No tension was developed
from weak discharge. At low frequencies, individual twitches emerged for each discharge (top right) (fig. S2). (C) Two pithed fish (fish 1, 19 g; fish 2, 21 g)
preparation. (D) Effect of curare. Red trace indicates strong electric organ discharge matched in time to unnormalized fish tension (green). Arrows
indicate time of injections (fig. S3). Bar in (D) = 500 ms.
Voltage Tension
90g
200 ms
Doublet
Expanded Doublet
5 ms
Weak EOD
Fish
Twitch
Doublet
Attack
Volley
Agar Barrier
Prey Movement
1
2 3
6 7
1
2
Prey Movement
1
3 4
1
5 6
9
3
10
9
25 ms
10
10
10
Fig. 3. Doublets during hunting. (A) Examples of doublets and corresponding tension responses. (B) Expansion of the first doublet and corresponding
tension trace (off-scale peaks were estimated). (C) Schematic of attack sequence. (D) Example of high-voltage electric organ discharge for an attack
preceded by a doublet. (E) Video frames from volley shown in (D). Numbers correspond to numbers in (D). (F) Timing of the high-voltage discharge for
attack preceded by a triplet. (G) Video frames for volley shown in (F).
1232
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RE S EAR CH | R E P O R T S
Carbon Electrode
Agar Barrier
EOD Recording
PowerLab 8/35
Stimulator
Plastic Bag
Stimulator
Off
Stimulator
(Movie S6 Clip 6)
Plastic Bag
Eel Attack
Eel
Stimulator
Triggered
By Doublet
Stimulator
Plastic Bag
Fish Twitch
Eel Attack
Stimulator
Triggered
By
Experimenter
Eel
Stimulator
Plastic Bag
Fish Twitch
Eel
Stimulator
Triggered
By Doublet
Stimulator
Plastic Bag
Stimulator
Triggered
By Doublet
Freeze-Thawed
Fish
Eel
Stimulator
(Movie S6 Clip 5)
Plastic Bag
Eel
Freeze-Thawed
Fish
No Plastic Bag
Below Plexiglas
Stimulator
Triggered
By
Experimenter
Eel
Stimulator
Fish Twitch
Plexiglas
(Movie S6 Clip 7)
Fig. 4. Paradigm and controls showing eels attack doublet-generated movements. (A) Movement
in electrically isolated pithed fish (below agar) was generated through stimulator. (B) Without fish twitch,
eels did not follow doublets with attack (10 trials each for two eels). (C) When stimulator triggered fish
twitch after doublets, eels attacked (10 trials each for two eels). (D) Without doublets, fish twitches also
elicited attack volleys (10 trials each of two eels). (E) Doublets that triggered stimulator leads in bag did
not elicit attack (10 trials for each of two eels). (F) Likewise, no attack volleys were elicited after stimulation
of a freeze-thawed fish (10 trials each of two eels). (G) Doublets directed at a freeze-thawed fish under
agar without the plastic bag or stimulator did not elicit eel attack volleys or strikes (10 trials each of two
eels). These latter conditions, along with (H) trials with Plexiglas barrier, show that visual cues did not
generate eel attacks. Examples are provided in movie S6 and (10).
SCIENCE sciencemag.org
1233
R ES E A RC H | R E PO R TS
www.sciencemag.org/content/346/6214/1231/suppl/DC1
Materials and Methods
Supplementary Text
Figs. S1 to S4
Movies S1 to S7
4 September 2014; accepted 13 November 2014
10.1126/science.1260807
INFLAMMATION
eutrophils are primary effectors of the immune response against invading pathogens
but are also central mediators of inflammatory injury (1). Both functions rely on
their remarkable ability to migrate within and through blood vessels. The migration of
neutrophils is initiated by tethering and rolling
on inflamed venules, a process mediated by endothelial selectins (2). Selectin- and chemokinetriggered activation of integrins then allows firm
adhesion, after which leukocytes actively crawl
on the endothelium before they extravasate or
return to the circulation (3). A distinct feature
of leukocytes recruited to inflamed vessels is the
RE S EAR CH | R E P O R T S
SCIENCE sciencemag.org
activated platelets might be mediating the interactions with the PSGL-1 clusters. Analyses of
mice deficient in P-selectin (Selp/ mice) indeed
demonstrated patterns of platelet interactions
with the two leukocyte subdomains that were
similar to those found in mice lacking PSGL-1
(Fig. 1B). These results indicated that neutrophils recruited to inflamed vessels extend a PSGL1bearing microdomain into the vessel lumen
that scans for activated platelets present in the
bloodstream through P-selectin.
During the course of our IVM experiments,
we also noticed alterations in the intravascular
behavior of adherent neutrophils in the different
mutant mice. Deficiency in Mac-1 severely compromised neutrophil crawling on the inflamed
vasculature (Fig. 2, A and B), a process previously reported to be mediated by this integrin
(3). Surprisingly, although PSGL-1 was excluded
from the area of contact with the endothelium
(Fig. 1, D and E), neutrophils deficient in this glycoprotein also displayed reductions in crawling displacement and velocity (Fig. 2, A and B),
Fig. 1. Neutrophils recruited to inflamed venules interact with activated platelets via protruding
PSGL-1 clusters. (A) Micrographs of polarized neutrophils interacting with platelets (red; yellow arrowheads) through the leading edge or the CD62L-labeled uropod (blue). (B) Quantification of total or domainspecific platelet interactions in wild-type mice or mice deficient in P-selectin (Selp/), PSGL-1 (Selplg/),
or Mac-1 (Itgam/); n = 5 to 8 mice, 38 to 133 interactions. (C) In vivo receptor distribution on polarized
wild-type neutrophils. (D) Examples of luminal and lateral projections from 3D reconstructions of polarized
Dock2-GFP neutrophils (see also movie S4). (E) Frequency of neutrophils extending PSGL-1 clusters into
the lumen (Lu), laterally (La) or between the cell body and the endothelium (En). n = 6 mice, 251 cells. (F)
3D reconstructions of an inflamed vessel showing the distribution of PSGL-1 clusters (movie S5). (G)
Representative micrographs of neutrophils interacting with nonactivated (arrow) or activated P-selectin+
platelets (arrowhead), and quantification of interactions of each domain with P-selectin+ or JON/A+
platelets. n = 3 to 4 mice, 66 to 116 interactions. Scale bars, 10 mm. Bars show mean T SEM. *P < 0.05; ***P <
0.001, one-way analysis of variance (ANOVA) with Tukeys post-hoc test.
5 DECEMBER 2014 VOL 346 ISSUE 6214
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R ES E A RC H | R E PO R TS
Fig. 3. PSGL-1 at the uropod becomes a preferred docking site for platelets during pathological
inflammation. (A) Survival curves of Balb/c mice treated with lipopolysaccharide (LPS) alone or LPS
plus antiMHC-I (MHC, major histocompatibility complex) to induce ALI. Neutrophils were depleted using
anti-Ly6G, and platelets using antiplatelet serum before the induction of ALI; n = 5 to 20 mice. (B) (Left)
Representative micrographs of inflamed venules during ALI. The asterisks indicate platelets interacting
with the uropod of neutrophils. (Right) Quantification of platelet interactions with the leading edge or
uropod in control (LPS only) and ALI-induced mice. Scale bar, 10 mm. n = 3 to 4 mice, 32 to 73 interactions.
(C) Frequency of interactions with the leading edge or uropod in TNF-atreated or ALI-induced mice, and
distribution of interactions in wild-type mice and mice deficient in PSGL-1 (Selplg/) or Mac-1 (Itgam/);
n = 3 to 5 mice, 23 to 137 interactions. (D) Frequency of interactions with the leading edge or uropod during
sepsis in wild-type mice and mice deficient in PSGL-1 or Mac-1. n = 3 to 4 mice, 32 to 56 interactions. Bars
show mean T SEM. *P < 0.05; ***P < 0.001 as determined by ANOVA with Tukeys multigroup test.
sciencemag.org SCIENCE
RE S EAR CH | R E P O R T S
Fig. 4. PSGL-1mediated
interactions trigger vascular
injury. (A) Survival curves of
Balb/c mice treated with LPS
alone or LPS plus antiMHC-I
to induce ALI. The absence of
Mac-1 or inhibition of PSGL-1
protects from death; n = 5 to
19 mice. (B) Representative
axial slices of the thorax of
Balb/c mice at different times
after induction of ALI. The
white signal in the pulmonary
space identifies edema,
which is quantified in (C); n = 7
to 8 mice per group. (D)
Quantification of hepatic injury
as levels of AST and ALT
transaminases in plasma of
the indicated group of mice
24 hours after treatment with
LPS; n = 7 to 11 mice. (E)
Representative brain sections
of wild-type mice 24 hours
after inducing ischemia,
showing vessels at increasing
magnifications and intravascular neutrophil (Ly6G,
green)platelet (CD41, red)
aggregates. Scale bars, 10 mm. (F) Percentages of infarcted hemispheres 24 hours after arterial occlusion in control wild-type mice, mice deficient in Mac-1 (Itgam/),
and wild-type mice after blocking PSGL-1. Images are representative brain sections stained with TTC, showing the extent of ischemia as white areas with a red
outline; n = 5 to 8 mice. Bars show mean T SEM. *P < 0.05; **P < 0.01; ***P < 0.001, ANOVA with Tukeys multigroup test.
1237
R ES E A RC H | R E PO R TS
19.
20.
21.
22.
TRANSCRIPTION
Chromatin decondensation is
sufficient to alter nuclear
organization in embryonic stem cells
Pierre Therizols, Robert S. Illingworth, Celine Courilleau, Shelagh Boyle,
Andrew J. Wood, Wendy A. Bickmore*
During differentiation, thousands of genes are repositioned toward or away from the
nuclear envelope. These movements correlate with changes in transcription and replication
timing. Using synthetic (TALE) transcription factors, we found that transcriptional
activation of endogenous genes by a viral trans-activator is sufficient to induce gene
repositioning toward the nuclear interior in embryonic stem cells. However, gene relocation
was also induced by recruitment of an acidic peptide that decondenses chromatin without
affecting transcription, indicating that nuclear reorganization is driven by chromatin
remodeling rather than transcription. We identified an epigenetic inheritance of chromatin
decondensation that maintained central nuclear positioning through mitosis even after
the TALE transcription factor was lost. Our results also demonstrate that transcriptional
activation, but not chromatin decondensation, is sufficient to change replication timing.
1238
www.sciencemag.org/content/346/6214/1234/suppl/DC1
Materials and Methods
Figs. S1 to S20
Tables S1 and S2
References (2429)
Movies S1 to S11
27 May 2014; accepted 7 November 2014
10.1126/science.1256478
(FISH) showed that Ptn, Sox6, and Nrp1 loci relocate away from the nuclear periphery and
toward more central nuclear positions, correlated
to their expression changes, during the differentiation of ESCs to EpiSCs or to NPCs (P <
0.01; Fig. 2B and fig. S1).
To directly address the role of transcription in
nuclear reorganization, we ectopically activated
Ptn, Sox6, or Nrp1 in ESCs by means of synthetic
transcription factors composed of TALE (transcription activatorlike effector) DNA binding
domains with specificity for the respective gene
promoters (9) fused to VP64, a tetramer of the
VP16 acidic transcriptional activator (10, 11) (Fig.
1A). When transfected into ESCs, tPtn-VP64
induced expression of its target by a factor of
>30 to 90 (Fig. 1, B and C). Other than Ptn, only
two additional genes (Il33 and Nnmt) were significantly up-regulated, and genes involved in
ESC pluripotency or differentiation were not
significantly changed. This suggests that Ptn
up-regulation is not just an indirect consequence
of differentiation triggered by transfection or
the nonspecific expression of an acidic activator (Fig. 1C and fig. S2). Specific activation of
Nrp1 or Sox6 in cells transfected by tNrp1-VP64
or tSox6-VP64, respectively, also showed no expression signature of differentiation (Fig. 1B).
Control plasmids lacking the VP64 domain
(tPtn-D, tSox6-D, and tNrp1-D) had almost no
effect (Fig. 1, B and C). Moreover, we did not
detect any changes in the expression of genes
neighboring those targeted by the TALEs (Fig. 1C
and fig. S3).
As well as activating Ptn, Nrp1, or Sox6,
FISH showed that tPtn-VP64, tNrp1-VP64, and
tSox6-VP64 caused specific relocalization of the
targeted loci toward the center of ESC nuclei,
relative to control [enhanced green fluorescent
protein (eGFP)] transfection (tPtn-VP64, P =
4.6 109; tNrp1-VP64, P = 5.3 1014; tSox6VP64, P = 6.4 1012) or to constructs lacking
the activation domain (tPtn-D, P = 3 106;
tNrp1-D, P = 8.1 1010; tSox6-D, P = 8 1012)
(Fig. 2, A and B). The extent of this relocalization was similar to that seen upon normal
sciencemag.org SCIENCE
RE S EAR CH | R E P O R T S
0.06), and tSox6-VP64 did not affect Nrp1 positioning (P = 0.08) (fig. S4, A to C). This reinforces the idea that TALE transfection does
not simply induce differentiation. We conclude
SCIENCE sciencemag.org
that forced Ptn, Nrp1, or Sox6 expression induces relocalization of the targeted loci toward
the center of the nucleus, independent of differentiation. This is consistent with a report
target genes (Ptn, Nrp1, Sox6), genes involved in pluripotency (Klf4, Oct4), or
genes involved in differentiation (FoxA2, Nestin, Fgf5) in ESCs transfected with
the different TALE vectors. Expression is shown relative to eGFP transfection.
Expression changes are also shown for the differentiation of ESCs to EpiSCs
or NPCs (n = 3 biological replicates). (C) Heat map showing 27 genes from the
expression microarray selected to represent different differentiation states, as
well as genes surrounding the regions targeted by the TALEs. Log2 transformed
mRNA ratios for ESCs transfected by the different tPtn constructs relative to
eGFP transfection, or for cells differentiated into NPCs relative to levels in
ESCs, are displayed using the color code shown.
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R ES E A RC H | R E PO R TS
was accompanied by relocation of the respective loci toward the nuclear center (tPtn-DEL,
P = 6.22 106; tNrp1-DEL, P = 3.4 1023;
tSox6-DEL, P = 1.8 108; Fig. 2B). The nuclear
repositioning induced by DEL peptide was similar to that induced by VP64 (tPtn-VP64, P = 0.93;
tNrp1-VP64, P = 0.03; tSox6-VP64, P = 0.9). Neither tPtn nor tNrp1 affected Sox6 condensation,
and Nrp1 condensation was not affected by
TALEs targeting Sox6 (fig. S4D and table S1).
We also did not observe any off-target effects
for TALE-DEL proteins on radial positioning
(fig. S4, A to C). Using chromatin immunoprecipitation (ChIP) for RNA polymerase II (RNAPII) (9), we confirmed that the effect of the DEL
peptide was independent of the transcription
sciencemag.org SCIENCE
RE S EAR CH | R E P O R T S
two biological replicates). (C) FISH at Ptn (green) and Sox6 (red) in ESCs
7 days after transfection by eGFP, tPtn-D, tPtn-VP64, or tPtn-DEL. Scale bars,
5 mm. Histograms below quantify the nuclear distribution of Ptn and Sox6
signals (as in Fig. 2B) for ESCs 7 days after transfection by eGFP, tPtn-D,
tPtn-VP64, and tPtn-DEL (n = 100 to 150 nuclei for each condition for each of
at least two biological replicates).
Fig. 4. Synthetic activation of transcription, but not nuclear repositioning, shifts replication timing. Mean (T SEM) log2 ratio of early/late
S phase fraction for Ptn, Nrp1, Sox6, Clec2l, and Mgam in ESCs after
transfection with (A) tPtn-D, tPtn-VP64, or tPtn-DEL; (B) tNrp1-D, tNrp1VP64, or tNrp1-DEL; and (C) tSox6-D, tSox6-VP64, or tSox6-DEL. For
comparison, changes in replication timing during differentiation of ESCs to EpiSCs or NPCs are included in all panels. Transfection with eGFP is also
shown at left of each group in all panels.
SCIENCE sciencemag.org
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R ES E A RC H | R E PO R TS
detectable transcriptional changes. A full dissection of chromatin events mediated by the DEL
peptide, and those present at targeted genes after
loss of the original TALE transcription factor,
would give further insight into the mechanisms
involved.
RE FERENCES AND NOTES
www.sciencemag.org/content/346/6214/1238/suppl/DC1
Materials and Methods
Figs. S1 to S6
Tables S1 to S3
References (1925)
4 August 2014; accepted 29 October 2014
10.1126/science.1259587
STRUCTURAL BIOLOGY
RE S EAR CH | R E P O R T S
by synchrotrons and by XFELs (10, 18). Timeresolved synchrotron studies take advantage of
an x-ray beam with exceptional stability, where
ideally a data set is collected on one large single crystal at essentially constant beam energy,
bandwidth, photon flux, and volume of the crystal exposed to the x-rays. The resulting data consist of sets of consecutive light and dark images
collected at the same orientation from the large
single crystal. This consistency of data acquisition is important, as structure factor changes between the light and dark states are often very
small. By contrast, several inherent pulse-to-pulse
variations make TR-SFX at atomic resolution
challenging: (i) The XFEL photon flux per pulse
can vary by up to an order of magnitude; (ii) the
peak energy and spectral content of the x-ray
beam changes from pulse to pulse; and (iii) the
crystal size is variable, and even if it were constant, the volume of the crystal interacting with
the beam can change. These factors give rise to
large fluctuations in the diffracted intensities.
However, the resulting total error is inversely proportional to the square root of the number of
diffraction patterns (18), and by collecting diffraction patterns from a large number of tiny crystals,
high-quality x-ray data can be obtained (15, 16, 19).
Despite these challenges, TR-SFX offers several advantages over time-resolved Laue crystallography at a synchrotron: (i) Time resolution,
largely set by the duration of the x-ray pulse in
the femtosecond time range, is substantially
higher; (ii) the diffraction-before-destruction
principle overcomes the x-ray damage problem;
(iii) each crystal diffracts only once and crystals
are rapidly exchanged, which provides an easy
way to address irreversible processes; (iv) the
quasi-monochromatic FEL x-ray beam allows
investigation of crystals with large unit cells;
(v) diffraction patterns are less sensitive to crystal
mosaicity than in the Laue method; and (vi) the
small size of the crystals (often <10 mm) allows
more uniform laser initiation of the reaction of
the molecules in the crystal. These advantages
were exploited in the first TR-SFX studies of
the large protein complexes, the photosystem
Iferredoxin complex and photosystem II, as
model systems (20, 21). Structural changes were
SCIENCE sciencemag.org
1243
R ES E A RC H | R E PO R TS
Fig. 2. Stereo view of the light-dark 1.6 difference electron density map at 1-ms time delay,
superimposed on the dark PYP structure (cyan). Contour levels: red/blue 3s/+3s. Chromophore
and some important chromophore pocket residues are shown in yellow and marked in (A). (A) Red
arrow: Plume of structural displacements extends to Met18, close to the N-terminal helix, which may
be strongly displaced at longer times (26, 34). (B) View rotated by ~90. A large part of the molecule
does not display sizable DED features and remains structurally unaltered.
extending to the periphery of the protein (red arrow). The positive (blue) and negative (red) DED
peaks are contiguous and can be interpreted in
terms of atomic models (Fig. 3, B and C). When
the resolution is reduced below 3 , the difference
signal disappears (fig. S7) and clear interpretation
of structural changes in PYP becomes impossible.
We note, however, that the minimum resolution
needed to observe structural changes is likely
system-dependent, and an extension to other
systems must await further work.
RE S EAR CH | R E P O R T S
Fig. 3. Comparison of electron density and DED maps in the chromophore pocket obtained by TR-SFX and the Laue method. The dark state
is shown in yellow in all maps. (A and D) Electron density maps for the PYP
dark state obtained with TR-SFX and Laue, respectively (contour level 1.1s,
1.6 resolution).The PCA chromophore and nearby residues are marked in
(A). Arrow: Double bond in the chromophore about which isomerization
occurs. (B) TR-SFX DED map at 10 ns. Light green structure: ICT interme-
diate. Features marked by dotted arrows belong to additional intermediates not shown. (C) TR-SFX DED map at 1 ms. Pink and red structures:
structures of pR1 and pR2 intermediates, respectively. (E) Laue 32-ns
DED map correlates best to the TR-SFX 10-ns map. (F) Laue 1-ms DED
map. Contour levels of the DED maps: red/white 3s/4s, blue/cyan
+3s/+5s, except for (C) where cyan is +7s. See fig. S8 for stereo versions
of (B) and (C).
1245
R ES E A RC H | R E PO R TS
1246
www.sciencemag.org/content/346/6214/1242/suppl/DC1
Materials and Methods
Supplementary Text
Figs. S1 to S10
Tables S1 to S4
References (3555)
29 July 2014; accepted 7 November 2014
10.1126/science.1259357
sciencemag.org SCIENCE
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Postdoctoral Researcher
Computational Biology
Sarcoma Disease Management Program
Departments of Surgery, Medical Oncology and Pathology
New York, NY
The Sarcoma Disease Management Program and the Computational Biology Center at MSK is searching
for an outstanding scientist with a strong interest in developing and applying computational methods
to cancer biology. Specically we are seeking a candidate with experience in both bioinformatics - in
particular, the bioinformatics of next-generation sequencing and high throughput genomic analysis and systems biology to lead the effort to analyze high throughput and genomic data from an extensive
dataset of patients treated at MSK for soft tissue sarcoma.
The candidate will have the opportunity to interact with expert investigators in computational biology,
sarcoma biology, translational therapeutics, and molecular pathology. The candidate will also analyze high
throughput genomic data and work with computational and laboratory-based investigators to identify the
mutational programs and pathways involved in sarcomagenesis and to identify novel therapeutic targets.
Position requires a PhD or equivalent in biological science, mathematics, physics, bioinformatics, or a related
eld with a strong quantitative background. Prociency with scientic computing software (Matlab, R) and
scripting languages are also required. The ideal candidate will have solid programming skills and extensive
research experience in the biological sciences. For consideration, please send CV, bibliography, brief
statement of research interests, and names of 3 references to [email protected]. Please put
#sarcoma in the subject line of your reply.
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MSK is an equal opportunity and afrmative action employer committed to diversity and inclusion in all aspects
of recruiting and employment.
Become a member of an elite research and development community involved in basic and applied
scientifc research and advanced technological development for tomorrows Navy.
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The Rutgers School of Environmental and Biological Sciences at Rutgers University - New Brunswick announces faculty searches
for three endowed chairs. We seek scholars whose records in teaching, research and service to society are consistent with anticipated
appointment as full Professor with tenure. Please visit the websites provided for further information and instructions for applications
and nominations.
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COLUMBUS, OHIO
Associate Dean
for Research
George C. Pafenbarger
Alumni-Endowed Chair in
Dental Research
The Ohio State University College of Dentistry seeks a
dynamic, experienced candidate for the position of
Associate Dean for Research.
Thisfull-time,tenuredpositionwillprovideleadershipandvisionfor
the research mission of the college, while building its extramural
funding base and expanding its reputation for world-class research.
The National Science Foundation ranks Ohio State 10th in total
research expenditures among public and private universities;
seventh among public universities; and second in industrysponsored research. The College of Dentistry is well funded with a
diverse and dynamic research portfolio that includes basic, clinical,
and translational science programs that are supported by seven
institutes at the NIH and by industry and various foundations.
Research foci include cancer, oral and gut microbiome, taste and
feeding behavior, psychoneuroimmunology, hard tissue and
muscle biology, and dental biomaterials.
The successful candidate will be an established and funded
investigator who can develop a compelling research focus for the
college, lead and manage existing programs, and foster a collegial
work environment. Required qualifcations include an earned
D.D.S., D.M.D., Ph.D., or equivalent degree in a related feld with
experience in academic health sciences.
The Associate Dean for Research also will be eligible for the George
C. Pa0enbarger Alumni-Endowed Chair in Dental Research, which
provides additional compensation for the position and resources to
support scholarship and o1ce operations.
For more information about the college and
this position, visit dent.osu.edu
Initial screening of applications will begin immediately and will
continue until an appointment is made. Application materials
should include: a letter addressing how the candidates experiences
match the position requirements; curriculum vitae; and contact
information for fve references. Confdential inquiries, nominations,
and application materials should be directed to:
Betty Turner Asher or Jan Greenwood
Greenwood/Asher & Associates, Inc.
42 Business Center Drive, Suite 206
Miramar Beach, FL 32550
Phone: 850.650.2277 | Fax: 850.650.2272
[email protected]
[email protected]
The Ohio State University is an equal opportunity employer. All qualifed applicants will
receive consideration for employment without regard to race, color, religion, gender, sexual
orientation or identity, national origin, disability status, and protected veteran status.
http://join.sjtu.edu.cn/
http://www.swjtu.edu.cn/
Institutes of Biomedical
Sciences, Fudan University
Southwest Jiaotong University (SWJTU), founded in 1896, situates itself in Chengdu, the provincial capital of Sichuan.
It is a national key multidisciplinary 211 and 985 Feature Projects university directly under the jurisdiction of the
Ministry of Education, featuring engineering and a comprehensive range of study programs and research disciplines
spreading across more than 20 faculties and institutes/centers. Boasting a complete Bachelor-Master-Doctor education
system with more than 2,500 members of academic staff, our school also owns 2 first-level national key disciplines, 2
supplementary first-level national key disciplines (in their establishment), 15 first-level doctoral programs, 43 first-level
master programs, 75 key undergraduate programs, 10 post-doctoral stations and more than 40 key laboratories at national
and provincial levels.
Our university is currently implementing the strategy of developing and strengthening the university by introducing and
cultivating talents. Therefore, we sincerely look forward to your working application.
More information available at http://www.swjtu.edu.cn/
I. Positions and Requirements
A.High-level Leading Talents
It is required that candidates be listed in national top talents programs such as Program of Global Experts, Top Talents
of National Special Support Program, Chang Jiang Scholars, China National Funds for Distinguished Young
Scientists and National Award for Distinguished Teacher.
Candidates are supposed to be no more than 50 years old. The limitation could be extended in the most-needed areas of
disciplinary development.
Candidates who work in high-level universities/institutes and reach the above requirements are supposed to be no more
than 45 years old.
B. Young Leading Scholars
Candidates are supposed to be listed in or qualified to apply for the following programs:
G J MN P
MS T
M N MJ
Y
G P
T
M N MJ J MN N
J
Y
Y
J M
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M N MJ
G
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MSJ M JP ab NN MJ T
JP P N
Candidates should have good team spirit and leadership, outstanding academic achievements, broad academic vision and
international cooperation experience and have the potential of being a leading academic researcher.
C. Excellent Young Academic Backbones
Candidates under 40 years old are expected to graduate from high-level universities/institutes either in China or other
countries. Those who are professors, associate professors and other equal talents from high-level universities/institutes
overseas could be employed as professors and associate professors as well.
D. Excellent Doctors and Post Doctoral Fellows
Candidates under 35 years old are supposed to be excellent academic researchers from high-level universities either in
China or other countries.
II. Treatments
The candidates will be provided with competitive salaries and welfares that include settling-in allowance, subsidy of
rental residence, start-up funds of scientific research, assistance in establishing scientific platform and research group as
well as international-level training and promotion . As for outstanding returnees, we can offer further or specific
treatments that can be discussed personally.
III . Contact us:
Contacts: Ye ZENG & Yinchuan LI
Telephone number: 86-28-66366202
Email: [email protected]
Address: Human Resources Department of SWJTU, the western park of high-tech zone, Chengdu, Sichuan, P.R.China,
611756
WU
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opportunities in china
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General enquiries should be directed to the SAMS President, Professor Geoffrey Boulton OBE
[email protected]; +44 131 667 2531 and/or Acting Director, Professor Axel Miller
[email protected]; +44 1631 559272.
Further information is available at www.sams.ac.uk, or e-mail [email protected],
or write to: Director/CEO SAMS Group, (D10/14.AM), Human Resources,
SAMS, Scottish Marine Institute, Oban, Argyll PA37 1QA
The closing date for applications is MONDAY 2ND FEBRUARY 2014
SAMS is a registered Scottish charity (SC 009206), has an equal opportunities policy and welcomes applications from all sections of the community.
University of Colorado
Denver Anschutz Medical Campus
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Jefferson Science
Fellowship
The National Academies is pleased to announce a call for
nominations and applications for the 2015 Jefferson Science
Fellows program. Initiated by the Secretary of State in 2003, this
fellowship program engages the American academic science,
technology, engineering and medical communities in the design
and implementation of U.S. foreign policy.
Jefferson Science Fellows (JSF) spend one year at the U.S.
Department of State or the U.S. Agency for International
Development (USAID) for an on-site assignment in Washington,
D.C. that may also involve extended stays at U.S. foreign
embassies and/or missions.
The fellowship is open to tenured, or similarly ranked, academic
scientists, engineers and physicians from U.S. institutions of higher learning. Nominees/applicants must hold U.S. citizenship and
will be required to obtain a security clearance.
The deadline for 2015-2016 program year applications/
nominations is January 12, 2015. To learn more about the
Jefferson Science Fellowship and to apply, visit the website at:
www.national-academies.org/jsf
Open Position in
Microbial Ecology: Microbiome/Infectious Disease
Department of Microbiology & Molecular Genetics
The Department of Microbiology & Molecular Genetics at Michigan State
University seeks candidates at the Assistant or Associate Professor level for a
position in the microbial ecology of infectious disease including interactions
between host, microbiome, pathogen and environment. Of special interest
within the broad area of the microbial ecology of infectious disease are
systems biology and evolution at the molecular level, physiology and metabolic
networks of microbial consortia, interactions of pathogens with mammalian
hosts, insect vectors, and/or microbiota within them, host innate immunity
to microbes, and the application of omics technology to study these areas.
Signifcant opportunities exist for collaboration with faculty associated with
the NSF BEACON Center (Evolution in Action), the NIH Enterics Research
Investigational Network, the NIH International Center of Excellence for
Malaria Research, the RDP, and the Center for Water Sciences. A strong record
of research accomplishment and/or potential for an independent, externally
funded research program with national visibility is expected. Academic rank
will be commensurate with experience. Teaching in graduate, professional, and/
or undergraduate programs is expected. The offer will include a competitive
startup package and laboratory facility.
Applications should be submitted electronically at https://jobs.msu.edu
(posting #0519) with requisite material (letter of interest, curriculum vitae listing
past and current funding if appropriate, a statement of future research goals,
and names of three potential references (not to be contacted until approval is
received from the applicant)) uploaded as a single PDF. Review of applications
will begin immediately and remain open until flled.
Additional information could be obtained via email to the Search Committee
Executive Assistant at [email protected]. http://www.mmg.msu.edu
MSU is an Affrmative-Action, Equal-Opportunity Employer and is
committed to achieving excellence through diversity. The University actively
encourages applications of women, persons of color, veterans, and persons
with disabilities, and we endeavor to facilitate employment assistance to
spouses or partners of candidates for faculty and academic staff positions.
DIRECTOR
Forensic Science Program
Middle Tennessee State University invites qualified
applicants for a tenure-track faculty position serving as
Director of the B.S. in Forensic Science program (rank
open). The Forensic Science degree program is an interdisciplinary program involving the departments of
Biology, Chemistry, and Criminal Justice Administration. Responsibilities for the director will include direction and evaluation of program activities, oversight of
internships, and management of initial program accreditation through FEPAC. The director will teach in undergraduate specialty courses. The director will also be
expected to direct his/her own research program involving undergraduate and graduate students. For more
information about the position requirements and application procedure, visit website: http://www.mtsu.
edu/hrs/.
Stop searching
for a job;
start your career.
ScienceCareers.org
Chair
Department of Computer and Information Sciences
College of Arts and Sciences
The UAB College of Arts and Sciences (CAS) invites applications for the position
of Chair of the Department of Computer and Information Sciences. For additional
information on the department, please visit https://cis.uab.edu/.
Computer and Information Sciences at UAB offers BS, MS, and PhD degrees. The
Department has a strong research focus, excellence in teaching, and a strong commitment
to outreach, innovation, and industrial partnerships. Research grants are growing
signifcantly, and the Department has a leading role in an Interdisciplinary Research
Center in Information Assurance and Computer Forensics. Departmental strengths
include research programs in diverse areas of Computer Science including programming
languages and software engineering, information assurance, natural language processing,
machine learning, data mining, computer graphics, visualization, 3D printing, computer
forensics, and high performance computing. Collaborations with UABs school of
engineering and the medical enterprise are strong and growing, with opportunities for
faculty to participate in interdisciplinary R&D.
The successful candidate will combine excellence in research and management with
a fundamental understanding of the operation and advancement of a research-focused
Computer Science department at a Carnegie Research Extensive University (RU-VH).
The chair is responsible for overall departmental administration and reports to the Dean
of the College of Arts and Sciences. The successful candidate will be expected to provide
strategic direction, vision, and leadership to the department and to participate actively
in development across the UAB campus and in the city of Birmingham. The University
is now embarking on the development of a 5 and 10 year plan to develop research
computing. There is an opportunity for signifcant growth of the department over the
next fve years and the new Chair is expected to play an active role in the development
of research computing infrastructure at UAB.
The Chair is the chief academic, fnancial, and strategic offcer for the Department. He or
she will work to recruit and retain outstanding faculty and students at all levels, contribute
to the intellectual life of the department as an active member of the faculty, and promote
the department within the University and, as importantly, outside the University to secure
support. A PhD in Computer Science or a closely related feld is required. Candidates must
possess a distinguished record of scholarship, grant funding, teaching, and professional
service consistent with appointment at the rank of Full Professor with tenure. Candidates
must exhibit excellent interpersonal, verbal, and written communication skills, and
interest in commercialization and entrepreneurship is welcome.
Applicants should submit (1) a cover letter summarizing qualifcations and explaining
interest in the position, (2) a complete curriculum vitae, (3) a statement of vision and
management style, and (4) contact information for at least fve references. References
will not be contacted without prior permission of the candidate.
Screening will begin on October 1, 2014, and will continue until the position is flled.
Please go to the following web-site to apply and upload application materials.
http://www.uab.edu/cas/application
For more information or to make inquiries about this position, contact:
Yogesh K. Vohra, PhD
Professor/Associate Dean
Chair, CIS Chair Search Committee
E-mail: [email protected]
Tel: (205) 934-6662
UAB is an Equal Opportunity/Affrmative Action Employer committed to fostering
a diverse, equitable and family-friendly environment in which all faculty and staff
can excel and achieve work/life balance irrespective of race, national origin, age,
genetic or family medical history, gender, faith, gender identity and expression as
well as sexual orientation. UAB also encourages applications from individuals with
disabilities and veterans.
A pre-employment background investigation is performed on candidates
selected for employment.
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POSITIONS OPEN
The Department of Biological Sciences (DBS) at the
University of North Carolina at Charlotte is seeking applicants for a tenure-track ASSISTANT/ASSOCIATE
PROFESSOR position. Applicants should possess a
Ph.D. in the biological sciences with expertise in Protein functions in a biological context. Candidates are
expected to document expertise in their specialty by
a record of postdoctoral training, peer-reviewed publications, and plans for supporting an extramurally funded
research program. Higher consideration will be given
to applicants that bring needed expertise in large glycoprotein function (i.e., Chaperones, Receptor and
Adaptor complexes). Previous teaching and mentoring
experience is highly desirable. In addition to contributing to graduate programming in the DBS, this position is part of a long-term effort to establish a resident
BProtein Function & Biotechnology Research[ Group
and will be expected to also make contributions to
the Nanoscale Science Ph.D. Program (jointly with
the Chemistry, Math & Statistics, Physics & Optics
Departments, and academic units in the College of
Engineering).
The successful candidate is expected to contribute
to teaching and mentoring in our undergraduate and
graduate curricula with a focus on Protein function in
a cell biological context. The Department of Biological
Sciences supports B.S., B.A., M.S. (thesis and non-thesis),
and Ph.D. programs with a diverse body of faculty and
students and prides itself with hands-on training of its
students. The Department and College strongly support
and value diversity among their students and faculty.
Candidates must apply online at website: http://
jobs.uncc.edu, position number: 6753. Please provide
complete curriculum vitae, philosophy statements for
research and teaching in an ethnically diverse environment, contact information for three references, and
three representative publications. Screening of applications will begin December 15, 2014, and continue
until the position is filled. The expected start date is
August 15, 2015.
For more information, please refer to the Department
of Biological Sciences website: http://biologicalsciences.
uncc.edu for contact information.
The University of North Carolina at Charlotte is an Equal
Opportunity Employer/Affirmative Action Employer and an
ADVANCE Institution that strives to create an academic climate
in which the dignity of all individuals is respected and maintained.
Therefore, we celebrate diversity that includes, but is not limited to
ability/disability, age, culture, ethnicity, gender, language, race,
religion, sexual orientation, and socio-economic status.
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POSITIONS OPEN
IMETs mission is to study the biology of coastal marine biosystems and ensure their sustainable
use, as well as exploit marine-derived systems to improve human health (www.imet.usmd.edu).
IMET brings together faculty members from three major USM research institutions - the University
of Maryland Baltimore (UMB), the University of Maryland Baltimore County (UMBC), and the
University of Maryland Center for Environmental Science (UMCES) - in an integrated state-of-theart research facility located at Baltimores Inner Harbor.
The Illinois Natural History Survey (INHS) is soliciting applications for a two-year POSTDOCTORAL
RESEARCH ASSOCIATE. Scientists whose research
interests fit within those of the Illinois Natural History
Survey are encouraged to apply. A salary of $42,000 per
year with benefits is provided as well as a $5,000/year
research stipend. Applicants should submit curriculum
vitae and Research Proposal. The Research Proposal is
limited to three pages, not including references, and
should address the research plan and budget proposed
for the two-year postdoctoral position. In advance of
the application deadline, applicants must identify and
contact a research sponsor at the INHS (website: http://
wwx.inhs.illinois.edu/opportunities/postdocsponsors/list/) who is willing to sponsor the Postdoctoral Associate. Preference will be given to applicants
who can develop a strong research plan that merits
additional and continued external funding. Research
plans that build on existing INHS research strengths
by adding new directions or new analytical techniques
are encouraged.
Applicants should have completed a Ph.D. by the
start date of the position (expected before December
31, 2015) and within the last five years. In addition to
the Research Proposal and curriculum vitae, applicants
should arrange for two letters of recommendation to be
sent. Applications must be sent electronically to e-mail:
[email protected] by January 31, 2015.
The USM is an Equal Opportunity, Affrmative Action Employer. The University of Maryland,
Baltimore is an Equal Opportunity/Affrmative Action Employer. Minorities, women, individuals
with disabilities, and protected veterans are encouraged to apply.
www.ScienceCareers.org
MemberCentral.aaas.org
GROWING IANR
Faculty Positions
Institute of Agriculture and Natural Resources
The Institute of Agriculture and Natural Resources (IANR) at the University
of Nebraska-Lincoln (UNL) is committed to world-class excellence in
applications of agricultural and life sciences towards a sustained high
quality of life for the citizens of Nebraska, and for a quickly growing
global population. Early in 2013, refecting this commitment, IANR
launched an initiative to hire new tenure-track faculty members in strategic
impact areas of Science Literacy; Stress Biology of Plants, Animals, and
Agroecosystems; Healthy Humans; Healthy Systems for Agricultural
Production and Natural Resources; and Computational Sciences. This Phase
1 effort was very successful, resulting in the recruitment and hiring of 35
highly skilled tenure-line faculty members (with two searches still active).
We are pleased to announce Phase 2 of this effort with recruitment for
an anticipated 30+ additional tenure-line faculty positions. The focus for
Phase 2 will be to strengthen the six strategic impact areas in Phase 1,
with an additional focus area Drivers of Economic Vitality for Nebraska.
We invite you to view brief explanations of the positions currently being
released and those to be released soon at http://ianr.unl.edu to explore
whether your skills and experience make you a good At for our team.
The positions will be advertised and posted beginning December 1, 2014.
We invite you to visit the UNL employment web site at http://employment.
unl.edu to learn which positions are posted.
The University of Nebraska is committed to a pluralistic campus
community through affrmative action, equal opportunity, work-life
balance, and dual careers.
Cernet
AAAS is here
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Cernet/
Sciencecareers.org/CER
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WORKING LIFE
By Vincent F. Scalfani
Finally free
s a child, I found relief from the stress of stuttering by burying myself in numerous hobbies. I
built rockets and radio-controlled vehicles. I rebuilt engines. I did these things because when
I was building something, I did not have to think about speaking. My hobbies eventually
led me to study chemistry. After finishing a B.S. in chemistry at the State University of New
York, Oswego, I studied polymer chemistry and earned a Ph.D. at Colorado State University.
I enjoyed chemistry research for the same reasons I enjoyed my hobbies as a child: I could
do interesting things, free from the stress of speaking. In the laboratory, I could be alone and build
things, although now I was building molecules instead of model cars.
I noticed early on that when I forgot about stuttering, my speech became much more fluent. The first
few words of a question or first
few minutes of a presentation were
rough, but then I forgot about the
audience, forgot about stuttering,
and forgot about worrying. When
that happened, my speaking improved. I felt free.
I wanted to feel free all the time.
Unfortunately, there were always
new people to meet and presentations to deliver, so there were many
awkward beginnings. I was ashamed
of my stutter and did everything I
could to hide it. Far from forgetting
about it, I developed strategies for
dealing with it. Whenever I felt I was
going to stutter, I would substitute a
new word that was easier to say. I
obsessed over how to deliver a talk
or just hold a conversation. I studied every tiny detail and came up
with alternative explanations ahead of time so that I could
always change terminology or the direction of the conversation. It was a wonderful irony: My incapacity for spontaneous speech led to a highly developed skill for improvisation.
My adaptations were so successful that most of my peers and
professors probably did not even know I stuttered.
Later, I would learn that hiding my stutter only perpetuated my anxiety and frustration with my speech. The skills
I acquired by hiding it, though, never left me.
After graduate school, my interests shifted away from
the laboratory, and I started to investigate alternative career paths. I accepted a position as assistant professor and
science and engineering librarian at the University of Alabama, Tuscaloosa. In my new job, I communicate regularly
and spontaneously with students, faculty, and staff. I knew
that if I wanted to succeed, I needed to change my attitude
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sciencemag.org SCIENCE
Published by AAAS
My incapacity for
spontaneous speech led to
a highly developed skill for
improvisation.