, 1164 (2010);
et al. Stuart H. M. Butchart,
Global Biodiversity: Indicators of Recent Declines
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between ORC binding and nucleosome turnover,
suggesting that turnover facilitates ORC binding.
In contrast, other chromatin features that would
be expected for open or dynamic chromatin, in-
cluding nucleosome density, mononucleosome/
oligonucleosome ratio (a measure of micrococcal
nuclease accessibility), H2Av (an H2A.Z his-
tone variant enriched in active chromatin), and
salt-soluble nucleosomes, show little if any de-
pendence on ORC abundance (Fig. 3, H to P).
Our findings support the hypothesis that repli-
cation origins are determined by chromatin, not
by sequence features (20, 21). The better quan-
titative correspondence of ORC to CATCH-IT
data than to other chromatin measurements implies
that the ORC occupies DNA that is made acces-
sible by nucleosome turnover. In support of this
interpretation, we note that very similar corre-
spondences are seen when CATCH-IT data are
aligned with GAF sites (fig. S9) and that GAF
directs nucleosome turnover in vivo (22, 23).
Our direct strategy for measuring the kinetics
of nucleosome turnover does not rely on trans-
genes or antibodies but rather uses native his-
tones and generic reagents. Thus, CATCH-IT
provides a general tool for studying activities
that influence nucleosome turnover. With use of
CATCH-IT, we found direct evidence that epige-
a process that erases histone modifications (10).
The fact that EZ is responsible for di- and tri-
methylation of H3K27, but the nucleosomes that
it modifies turn over faster than a cell cycle,
argues against proposals that histone modifica-
tions required for cellular memory themselves
transmit epigenetic information (24). Rather, by
simply increasing or decreasing accessibility of
DNA to sequence-specific binding proteins, regu-
lated nucleosome turnover may perpetuate active
or silent gene expression states and facilitate ini-
tiation of replication.
References and Notes
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25. We thank T. Furuyama for suggesting this approach,
members of our lab for helpful discussions, and the
Hutchinson Center Genomics Shared Resource for
microarray processing. This work was supported by NIH
grant 1R21DA025758 to S.H. and NIH Postdoctoral
Fellowship 1F32GM083449 to R.B.D. All data sets can be
found in GEO: GSE19788.
Supporting Online Material
Materials and Methods
Figs. S1 to S9
7 January 2010; accepted 1 April 2010
Global Biodiversity: Indicators of
Stuart H. M. Butchart,1,2* Matt Walpole,1Ben Collen,3Arco van Strien,4
Jörn P. W. Scharlemann,1Rosamunde E. A. Almond,1Jonathan E. M. Baillie,3
Bastian Bomhard,1Claire Brown,1John Bruno,5Kent E. Carpenter,6Geneviève M. Carr,7†
Janice Chanson,8Anna M. Chenery,1Jorge Csirke,9Nick C. Davidson,10Frank Dentener,11
Matt Foster,12Alessandro Galli,13James N. Galloway,14Piero Genovesi,15
Richard D. Gregory,16Marc Hockings,17Valerie Kapos,1,18Jean-Francois Lamarque,19
Fiona Leverington,17Jonathan Loh,20Melodie A. McGeoch,21Louise McRae,3
Anahit Minasyan,22Monica Hernández Morcillo,1Thomasina E. E. Oldfield,23Daniel Pauly,24
Suhel Quader,25Carmen Revenga,26John R. Sauer,27Benjamin Skolnik,28Dian Spear,29
Damon Stanwell-Smith,1Simon N. Stuart,1,12,30,31Andy Symes,2Megan Tierney,1
Tristan D. Tyrrell,1Jean-Christophe Vié,32Reg Watson24
In 2002, world leaders committed, through the Convention on Biological Diversity, to achieve
a significant reduction in the rate of biodiversity loss by 2010. We compiled 31 indicators to report
on progress toward this target. Most indicators of the state of biodiversity (covering species’
population trends, extinction risk, habitat extent and condition, and community composition)
showed declines, with no significant recent reductions in rate, whereas indicators of pressures
on biodiversity (including resource consumption, invasive alien species, nitrogen pollution,
overexploitation, and climate change impacts) showed increases. Despite some local successes
and increasing responses (including extent and biodiversity coverage of protected areas,
sustainable forest management, policy responses to invasive alien species, and biodiversity-related
aid), the rate of biodiversity loss does not appear to be slowing.
the current rate of biodiversity loss” (1), and this
Convention on Biological Diversity (CBD),
“2010 target” has been incorporated into the
United Nations Millennium Development Goals
in recognition of the impact of biodiversity loss
on human well-being (2). The CBD created a
framework of indicators to measure biodiversity
loss at the level of genes, populations, species,
and ecosystems (3, 4). Although a minority have
not been synthesized to provide an integrated
outcome. Despite suggestions that the target is
unlikely to be (6–8), or has not been (4, 9, 10),
met, we test this empirically using abroad suite of
To evaluate achievement of the 2010 target,
of significant inflections in trend for individual
indicators (11) and (ii) calculated aggregated in-
dices relating to the state of biodiversity, pres-
sures upon it, policy and management responses,
and the state of benefits (ecosystem services) that
people derive from biodiversity, using the best
available sources. To calculate aggregate indices,
we first scaled each of 24 indicators (out of 31)
the first year with data from 1970 onward (only
eight indicators had earlier trends) and calculated
annual proportional change from this first year.
Then we used a generalized additive modeling
inflections (12). Although absolute values are
difficult to interpret because they aggregate dif-
ferent elements of biodiversity, this approach
permits a synthetic interpretation of rate changes
across the elements measured: For example, the
aggregated state index should show positive
inflections if biodiversity loss has been signifi-
28 MAY 2010VOL 328
on May 27, 2010
Our analyses suggest that biodiversity has
continued to decline over the past four decades,
with most (8 out of 10) state indicators showing
negative trends (Fig. 1 and Table 1). There have
been declines in population trends of (i) ver-
tebrates (13) and (ii) habitat specialist birds; (iii)
shorebird populations worldwide; extent of (iv)
and (vii) the condition of coral reefs. None show
significant recent reductions in the rate of decline
(Table 1), which is either fluctuating (i), stable(ii),
Two indicators, freshwater quality and trophic in-
tegrity in the marine ecosystem, show stable and
marginally improving trends, respectively, which
are likely explained by geographic biases in data
availability for the former and spatial expansion
of fisheries for the latter (5). Aggregated trends
across state indicators have declined, with no sig-
nificant recent reduction in rate: The most recent
2).Becausetherewerefewer indicatorswith trend
data in the 1970s, we recalculated the index from
loss: The most recent inflection (2004) was neg-
ative. Finally, aggregated species’ extinction risk
(i.e., biodiversity loss at the species level) has ac-
rate of change (16, 17), shows negative trends.
The majority of indicators of pressures on
biodiversity show increasing trends over recent
decades (Fig. 1 and Table 1), with increases in (i)
aggregate human consumption of the planet’s
ecological assets, (ii) deposition of reactive nitro-
gen, (iii) number of alien species in Europe, (iv)
proportion of fish stocks overharvested, and (v)
impact of climate change on European bird pop-
ulation trends (18). In no case was there a signif-
icant reduction in the rate of increase (Table 1),
based on too few data to test significance (ii),
have slowed, and this may explain why the most
recent inflection in aggregated trends (in 2006)
was negative (Fig. 2) (5). Global trends for
habitat fragmentation are unavailable, but it is
ing Atlantic Forest fragments are <0.5 km2in
size (19), and 59% of large river systems are
moderately or strongly fragmented by dams and
1United Nations Environment Programme World Conservation
Monitoring Centre, 219 Huntingdon Road, Cambridge CB3
0DL, UK.2BirdLife International, Wellbrook Court, Cambridge
CB3 0NA, UK.
London, Regent’s Park, London NW1 4RY, UK.
Netherlands, Post Office Box 24500, The Hague, 2490 HA,
Netherlands.5Department of Marine Sciences, University of
North Carolina at Chapel Hill, 340 Chapman Hall, CB 3300,
Chapel Hill, NC 27599, USA.
Conservation of Nature (IUCN) and Conservation International
Global Marine Species Assessment, Biological Sciences, Old
Dominion University, Norfolk, VA 23529, USA.
Nations Environment Programme, Global Environment Mon-
itoring System—Water, c/o National Water Research Institute,
867 Lakeshore Road, Burlington, Ontario L7R 4A6, Canada.
8IUCN Species Survival Commission, Conservation Interna-
tional, Biodiversity Assessment Unit, c/o Center for Applied
Biodiversity Science, Conservation International, 2011 Crystal
Drive, Suite 500, Arlington, VA 22202, USA.9Fisheries and
Aquaculture Management Division, Food and Agriculture
Organization of the United Nations, Viale delle Terme di
Caracalla 00153, Rome, Italy.10Secretariat of the Ramsar
Convention on Wetlands, Rue Mauverney 28, 1196 Gland,
Switzerland.11European Commission Joint Research Centre,
Institute for Environment and Sustainability, TP290, Via
Enrico Fermi 2749, 21027 Ispra (VA), Italy.
Applied Biodiversity Science, Conservation International,
2011 Crystal Drive, Suite 500, Arlington, VA 22202, USA.
13Global Footprint Network, 312 Clay Street, Suite 300,
Oakland, CA 94607–3510, USA.14Environmental Sciences
Department, University of Virginia, Charlottesville, VA
Ricerca Ambientale, Via Curtatone 3, I-00185 Rome, Italy.
16Royal Society for the Protection of Birds, The Lodge, Sandy
SG19 2DL, UK, and European Bird Census Council.
17School of Integrative Systems, University of Queensland,
St. Lucia, Brisbane, Qld 4067, Australia.18Department of
Zoology, University of Cambridge, Downing Street, Cam-
bridge CB2 3EJ, UK.
Research, 3450 Mitchell Lane, Boulder, CO 80301, USA.
20World Wildlife Fund (WWF) International, 1196 Gland,
Invasion Biology and Global Invasive Species Programme,
Post Office Box 216, Steenberg 7947, South Africa.22United
Nations Educational, Scientific, and Cultural Organization,
7 place de Fontenoy, 75352 Paris, France.
International, 219 Huntingdon Road, Cambridge CB3 0DL,
UK.24Sea Around Us Project, Fisheries Centre, University of
British Columbia, 2202 Main Mall, Vancouver, BC V6T1Z4,
Institute of Fundamental Research, GKVK Campus, Bellary
Road, Bangalore 560 065, India.26The Nature Conservancy,
4245 North Fairfax Drive, Arlington, VA 22203, USA.27U.S.
Geological Survey, Patuxent Wildlife Research Center, 12100
Beech Forest Road, Laurel, MD 20708–4039, USA.28Amer-
ican Bird Conservancy, 1731 Connecticut Avenue, N.W., 3rd
Floor, Washington, DC 20009, USA.29Centre for Invasion
Biology, Stellenbosch University, Private Bag X1, Matieland
7602, South Africa.
Department of Biology and Biochemistry, University of Bath,
Bath BA2 7AY, UK.31Al Ain Wildlife Park and Resort, Post
Office Box 45553, Abu Dhabi, United Arab Emirates.32IUCN,
Rue Mauverney 28, 1196 Gland, Switzerland.
*To whom correspondence should be addressed. E-mail:
†Present address: Indian and Northern Affairs Canada, 15
Eddy, Gatineau QC K1A 0H4, Canada.
3Institute of Zoology, Zoological Society of
6International Union for
15Istituto Superiore per la Protezione e la
19National Center for Atmospheric
21South African National Parks, Centre for
25National Centre for Biological Sciences, Tata
30IUCN Species Survival Commission,
Fig. 1. Indicatortrendsfor(A)thestateofbiodiversity,(B)pressuresuponit,(C)responsestoaddressits
loss, and (D) the benefits humans derive from it. Data scaled to 1 in 1970 (or for first year of data if
>1970), modeled (if >13 data points; see Table 1), and plotted on a logarithmic ordinate axis. Shading
shows 95% confidence intervals except where unavailable (i.e., mangrove, seagrass, and forest extent,
LPI, Living Planet Index; RLI, Red List Index; IBA, Important Bird Area; AZE, Alliance for Zero Extinction
site; IAS, invasive alien species.
VOL 32828 MAY 2010
on May 27, 2010
Table 1. Summary of global biodiversity indicator trends.
% Change since
Mean annual % change§
rate of change║
1970s 1990s2000s Since 1970
–31* Living Planet Index (LPI)
(mean population trends of vertebrates)
Wild Bird Index [mean population trends
of habitat specialists in Europe and North
America, disaggregated for terrestrial (t)
and wetland (w) species]
Waterbird Population Status Index
(% shorebird populations increasing,
stable, or decreasing)
Red List Index (RLI) (extinction risk of
mammals, birds, amphibians, and corals)
Marine Trophic Index
(shift in fishing catch from top
predators to lower trophic levels)
Coral reef condition
(live hard coral cover)
Water Quality Index
(physical/chemical quality of freshwater)
Number of state indicators declining
1950–2006 +3.0* +0.1–0.1 +0.1+0.1 +0.1S
1980–20050 +0.1+0.0–0.2 +0S
2/38/9 8/107/10 8/10
(humanity’s aggregate resource-consumption)
Nitrogen deposition rate
(annual reactive N deposited)
No. alien species in Europe
(Mediterranean marine, mammal, and freshwater)
Exploitation of fish stocks
(% overexploited, fully exploited, or depleted)
Climatic Impact Indicator
(degree to which European bird population trends
have responded in the direction
expected from climate change)
Number of pressure indicators increasing
1961–2006 +2.0+1.3+1.3 +2.1 +1.6S
1850–2005† +35+2.0+1.3–0.3 +0.2+0.9D?
1970–2007+76*+2.0 +1.4+1.6 +1.1+1.5S
1974–2006+31* +0.6+0.6 +1.1+1.2 +0.9F
4/44/54/5 5/5 5/5
Extent of Protected Areas (PAs)
Coverage by PAs of Important
Bird Areas and Alliance for Zero Extinction sites
Area of forest under sustainable
management (FSC certified)
International IAS policy adoption
(no. signatories to conventions
with provision for tackling IAS)
National IAS policy adoption
(% countries with relevant legislation)
Official development assistance
(US$ per year provided in support of CBD)
Number of response indicators increasing
1995–2008 +12,000*+100+20 +46D 2006
1964–2009 +10,000*+30 +8.7+12+4.1 +13 D 2004–2009
LPI for utilized vertebrate populations
RLI for species used for food and medicine
RLI for bird species in international trade
Number of benefits indicators declining
*Significant trends (P < 0.05).
date with data if this is post-1970.
comparisons between decades for the same indicator.
significant positive and negative changes), or with too few data points to test significance (?); years indicate periods in which second derivatives differed significantly from zero (P < 0.05).
†Identifies indicators with insufficient data to test significance of post-1970 trends, usually because annual estimates are unavailable.
§Because the indicators measure different parameters, some comparisons of mean annual % change between indicators are less meaningful than
║Rate of change decelerating (D), accelerating (A), stable (S, i.e., no years with significant changes), fluctuating (F, i.e., a sequence of
28 MAY 2010 VOL 328
on May 27, 2010
All indicators of policy and management
responses show increasing trends (Fig. 1 and
Table 1), with increases in (i) extent of protected
subsets of Key Biodiversity Areas (21) [39% of
of the area of 561 Alliance for Zero Extinction
sites (22) by 2009]; (iii) area of sustainably
managed forests [1.6 million km2under Forest
(iv) proportion of eligible countries signing inter-
national agreements relevant to tackling invasive
alien species (IAS) [reaching 82% by 2008 (23)];
(v) proportion of countries with national legisla-
tion to control and/or limit the spread and impact
of IAS [reaching 55% by 2009 (23)]; and (vi)
biodiversity-related aid (reaching US$3.13 billion
slowing (ii, iii, and v), or based on too few data to
test significance (vi) (Table 1). The last three in-
flections in aggregated trends (2002, 2004, and
2008) were all negative (Fig. 2), indicating that
the rate of improvement has slowed. Two other
indicators have only baseline estimates: Manage-
ment effectiveness was “sound” for 22% of PAs
(“basic” for 65% and “clearly inadequate” for
13%), and the proportion of genetic diversity for
200 to 300 important crop species conserved ex
situ in gene banks was estimated to be 70% (24).
Only three indicators address trends in the
benefits humans derive from biodiversity (Fig.
1 and Table 1): (i) population trends of utilized
aggregate species’ extinction risk has increased
Fig. 2. Aggregated indices of (A) the state of bio-
trends, habitat extent and condition, and community
indicators of ecological footprint, nitrogen deposition,
impacts; and (C) responses for biodiversity based on six
indicators of protected area extent and biodiversity cov-
able forest management, and biodiversity-related aid.
intervals derived from 1000 bootstraps. Significant
(filled circles) inflections are indicated.
Table 2. Examples of successes and positive trends relevant to the 2010 target (5).
IndicatorSuccesses and positive trends
Living Planet Index of Palearctic
Waterbird populations in
North America and Europe
Species downlisted on the
IUCN Red List
Increased by 43% since 1970 (e.g., Eurasian beaver and common buzzard)
Increased by 44% since 1980 owing to wetland protection and sustainable management
(but populations remain below historic levels).
Species qualifying for downlisting to lower categories of extinction risk owing to successful
conservation action include 33 birds since 1988 (e.g., Lear’s macaw), 25 mammals since
1996 (e.g., European bison), and 5 amphibians since 1980 (e.g., Mallorcan midwife toad).
Annex 1–listed species’ population trends have improved in EU countries (27) and
extinction risk reduced (RLI increased 0.46% during 1994–2004) owing to designation of
Special Protected Areas and implementation of Species Action Plans under the directive
(e.g., white-tailed eagle).
At least 16 bird species extinctions were prevented by conservation actions during 1994–2004,
e.g., black stilt (28).
Improved by 7.4% since 1970.
Slowed from 2.8 million ha in 2003–2004 to 1.3 million ha in 2007–2008, but it is uncertain
to what extent this was driven by improved enforcement of legislation versus reduced
demand owing to economic slowdown.
87% of countries have now developed NBSAPs and therefore have outlined coherent plans
for tackling biodiversity loss at the national scale.
Nearly 133,000 PAs designated, now covering 25.8 million km2: 12% of the terrestrial
surface (but only 0.5% of oceans and 5.9% of territorial seas), e.g., Juruena National Park,
Brazil, designated in 2006, covering 19,700 km2of Amazon/cerrado habitat.
82% of eligible countries have signed international agreements relevant to preventing the
spread and promoting the control/eradication of IAS. Successful eradications/control of IAS
include pigs on Clipperton Atoll, France (benefiting seabirds and land crabs), cats, goats and
sheep on Natividad, Mexico (benefiting black-vented shearwater), and red fox in southwest
Australia (benefiting western brush wallaby).
Increased to at least US$3.13 billion in 2007.
Wild Bird Index and Red List
Index for species listed on the
European Union Birds Directive
Water Quality Index in Asia
National biodiversity strategies
and action plans (NBSAPs)
Protected areas (PAs)
Invasive alien species (IAS)
policy, eradication, and control
assistance for biodiversity
VOL 32828 MAY 2010
on May 27, 2010
at an accelerating rate (as shown by the RLI) for Download full-text
(ii) mammals, birds, and amphibian species used
for food and medicine (with 23 to 36% of such
that are internationally traded (principally for the
pet trade; 8% threatened). Trends are not yet
available for plants and other important utilized
trend data, show (iv) 21% of domesticated an-
imal breeds are at risk of extinction (and 9% are
already extinct); (v) languages spoken by fewer
than 1000 people (22% of the current 6900 lan-
guages) have lost speakers over the past 40 years
and are in danger of disappearing within this
century (loss of linguistic diversity being a proxy
for loss of indigenous biodiversity knowledge);
and (vi) more than 100 million poor people live
in remote areas within threatened ecoregions and
are therefore likely to be particularly dependent
upon biodiversity and the ecosystem services it
Indicator development has progressed sub-
stantially since the2010 targetwasset.However,
there are considerable gaps and heterogeneity in
geographic, taxonomic, and temporal coverage
of existing indicators, with fewer data for devel-
oping countries, for nonvertebrates, and from
before 1980 and after 2005 (4, 5, 25). Interlink-
ages between indicators and the degree to which
they are representative are incompletely under-
stood. In addition, there are gaps for several key
aspects of state, pressures, responses, and espe-
cially benefits (4, 5, 7, 26).
Despite these challenges, there are sufficient
data on key dimensions of biodiversity to con-
clude that at the global scale it is highly unlikely
that the 2010 target has been met. Neither indi-
vidual nor aggregated indicators of the state of
rates of decline, apart from coral reef condition,
for which there has been no further deceleration
in decline since the mid-1980s. Furthermore, all
specific exceptions with positive trends for par-
ticular populations, taxa, and habitats (Table 2)
suggest that, with political will and adequate re-
sources, biodiversity loss can be reduced or re-
at a decelerating rate (and with little direct infor-
mation on whether such actions are effective).
increasing pressures and slowing responses.
Our results show that, despite a few encour-
aging achievements, efforts to address the loss of
biodiversity into broad-scale land-use planning,
incorporating its economic value adequately into
and implementing policies that tackle biodiversity
in coherent global biodiversity monitoring and in-
dicators is essential to track and improve the ef-
fectiveness of these responses.
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P. Martin, I. May, A. Milam, K. Noonan-Mooney, H. Pavese,
B. Polidoro, C. Pollock, D. Pritchard, J. Schipper,
F. Schutyser, V. Shutte, S. Simons, J. Sˇkorpilová,
A. Stattersfield, P. Voříšek, R. Wright, M. Wackernagel,
and M. Waycott. We acknowledge support from the Global
Environment Facility to the 2010 Biodiversity Indicators
Partnership; Shell Foundation; European Commission; the
Sea Around Us Project (University of British Columbia/Pew
Environment Group) to D.P. and R.W.; World Wildlife
Fund, The Nature Conservancy, and the University of
Queensland to M.H. and F.L.; T. Haas and the New
Hampshire Charitable Foundation to K.E.C.; and the
National Science Foundation (NSF) to J.-F.L. Opinions
and findings expressed here do not necessarily reflect the
views of the NSF or other funding bodies.
Supporting Online Material
Figs. S1 and S2
Tables S1 to S4
Data File 1
26 January 2010; accepted 8 April 2010
Published online 29 April 2010;
Include this information when citing this paper.
Plectasin, a Fungal Defensin,
Targets the Bacterial Cell Wall
Precursor Lipid II
Tanja Schneider,1Thomas Kruse,2Reinhard Wimmer,3Imke Wiedemann,1Vera Sass,1
Ulrike Pag,1Andrea Jansen,1Allan K. Nielsen,4Per H. Mygind,4Dorotea S. Raventós,4
Søren Neve,4Birthe Ravn,4Alexandre M. J. J. Bonvin,5Leonardo De Maria,4
Anders S. Andersen,2,4Lora K. Gammelgaard,4Hans-Georg Sahl,1Hans-Henrik Kristensen4*
Host defense peptides such as defensins are components of innate immunity and have retained
antibiotic activity throughout evolution. Their activity is thought to be due to amphipathic
structures, which enable binding and disruption of microbial cytoplasmic membranes. Contrary to
this, we show that plectasin, a fungal defensin, acts by directly binding the bacterial cell-wall
precursor Lipid II. A wide range of genetic and biochemical approaches identify cell-wall
biosynthesis as the pathway targeted by plectasin. In vitro assays for cell-wall synthesis identified
Lipid II as the specific cellular target. Consistently, binding studies confirmed the formation of an
equimolar stoichiometric complex between Lipid II and plectasin. Furthermore, key residues in
plectasin involved in complex formation were identified using nuclear magnetic resonance
spectroscopy and computational modeling.
lectasin is a 40–amino acid residue fungal
defensin produced by the saprophytic as-
comycete Pseudoplectania nigrella (1).
Plectasin shares primary structural features with
mussels and folds into a cystine-stabilized alpha-
28 MAY 2010 VOL 328
on May 27, 2010