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CONCEPTS AND
The global reach of human activities affects all natural ecosystems, so that the environment is best viewed as
a social--ecological system. Consequently, a more integrative approach to environmental science, one that
bridges the biophysical and social domains, is sorely needed. Although models and frameworks for
social--ecological systems exist, few are explicitly designed to guide a long-term interdisciplinary research
program. Here, we present an iterative framework, "Press-Pulse Dynamics!! (PPD), that integrates the bio
physical and social sciences through an understanding of how human behaviors affect "press!! and "pulse!!
dynamics and ecosystem processes. Such dynamics and processes, in turn, influence ecosystem services -
thereby altering human behaviors and initiating feedbacks that impact the original dynamics and processes.
We believe that research guided by the PPD framework will lead to a more thorough understanding of
social--ecological systems and generate the knowledge needed to address pervasive environmental problems.
O
ver the past 50 years, ecosystems have been altered by to move toward a more sustainable world, in which the
humans more than at any other time in recorded his health of ecosystems and human well-being are improved
tory (Vitousek et al. 1997; Chapin et al. 2010), and those and ecosystem services are distributed more equitably for
changes have resulted in reciprocal effects on human well current and future generations. As ecological research
being (MA 2005). Although health and wealth have, on expands from site-based science to regional and global
average, improved, in part as a consequence of these scales (Peters et al. 2008), the conceptual scope of ecology
ecosystem changes, the social and geographic distribution must also expand to embrace not only other scientific dis
of benefits to human populations remains uneven. ciplines, but also the pervasive human dimensions of envi
Furthermore, such improvements are often limited by the ronmental structure and change. Every ecosystem on Earth
inability of ecosystem services to keep pace with human is influenced by human actions (Vitousek et al. 1997;
demand and unequal opportunity for different people to Palmer et al. 2005), and the consensus view now holds
access these services (MA 2005). Learning how to manage that, for many of today's most pressing issues, the environ
feedbacks between ecosystems and humans is vital if we are ment is best understood and studied as a social-ecological
system (Uu et al. 2007).
As recognition of the importance of social-ecological
In a nutshell: science increases, new interdisciplinary linkages are
• T here is growing recognition that the environment must be evolving. Global research programs, such as the
viewed and studied as a social-ecological system International Geosphere-Biosphere Programme and the
• Various conceptual models have been proposed to character
International Human Dimensions Programme on Global
ize social-ecological systems, but new thinking is needed to
guide long-term research that links humans with their envi Environmental Change (Steffen et al. 2004; Carpenter
ronment and Folke 2006), have driven important advances.
• We describe a new model for integrated social-ecological Collaborations between physical scientists and biologists
research, the key components of which include environmen have occurred with the advent of regional- and global
tal and social sciences, press and pulse interactions, and
scale science, whereas in applied sciences, such as agron
ecosystem services
• Application of this approach will bridge the social and nat
omy and fisheries, collaborations between ecologists and
ural sciences and build a knowledge base that can be used to social scientists are more recent. For example, studies on
help solve current and future environmental challenges how ecosystem services benefit society formed the core of
the Millennium Ecosystem Assessment (MA 2005), the
first interdisciplinary global assessment of Earth's ecosys
IDepartrnent of Biology, University of New Mexico, AlbU£juerque, tems conducted at the behest of world leaders. Many early
NM '(scoUins@seviUeta.unm.edu); refer to WebPanel 1 for the re advances in social-ecological research were driven by
maining authors' affiliations, in their entirety coalitions of ecologists and economists (Goulder and
Kennedy 1997; Berkes and Folke 1998; Berkes et al. 2003) therefore a compelling need for a comprehensive concep
seeking to understand how institutions and economies tual framework that is based on highly relevant discipli
solve common-property resource problems (Ostrom 1990; nary research, but at the same time facilitates linkages
Dietz et al. 2003), and more recently by studies of resiliency among disciplines over the time frames and spatial scales at
in regional social-ecological systems (Gunderson and which social-ecological systems operate and interact.
Holling 2002; Walker and Salt 2006). Uu et al. (2007) illus Understanding change is a fundamental challenge for
trated the diversity of approaches that have been applied to environmental science. Social-ecological systems can
site-based social-ecological research, while emphasizing the transform incrementally and at times predictably. Some of
enormous gaps in interdisciplinary science that remain, and the most important routine changes (eg post-fire succes
the need for new theory that will better integrate concep sion or housing prices) are reasonably well understood and
tual and empirical research across disciplines. are incorporated into management practices, yet these
Although research in "pure" social and biophysical sci changes are best understood primarily within the biophys
ences must continue, new emphasis and approaches are ical or social-system contexts. Other changes are large in
also needed to understand the dynamic processes that are magnitude, are spatially extensive, and alter social-eco
unique to social-ecological systems. Ecosystems are com logical systems for long time periods; examples include
posed of numerous species - across the trophic spectrum - the loss of keystone species, land-use change drivers (such
that interact at varying rates and at multiple scales, from as zoning practices and homestead policy), or the
which the patterns and dynamics that we observe emerge increased demand for biofuels. Although large changes
(Levin 1999). Social systems also self-organize and exhibit may account for most of the cumulative dynamics
scale dependencies, but humans within such systems pos observed, they are infrequent - or pulsed - in nature. As a
sess capabilities that qualitatively change these dynamics consequence, observations of these pulsed events are few,
in important ways (Gibson et al. 2000; Westley et al. 2002). individual cases may be unique, and our ability to general
For example, people make forward-looking decisions (ie ize or predict their impacts on social-ecological systems
they act on expectations of the future), generate and remains severely limited. Understanding the drivers and
respond to abstract perceptions that shape their worlds and interactions berween sudden events ("pulse" dynamics)
their expectations, create feedbacks that act on various and extensive, pervasive, and subtle change ("press"
time scales over multiple spatial extents, and develop tech dynamics) is therefore one of the most important chal
nologies with far-reaching consequences (Westley et al. lenges for social-ecological science.
2002). These consequences create complex dynamics and We propose that press-pulse dynamics and ecosystem
often unexpected outcomes, which may have long-term services can form the critical linkage between social and
effects on social-ecological systems (Uu et al. 2007). biophysical domains and serve as the foundation for long
We are now beginning to see some of the emerging term, integrated, social-ecological research across scales.
trends, dynamics, feedbacks, and surprises that are impor Figure 1 presents the basic components of a framework,
tant for human well-being, but we are a long way from known as the "Press-Pulse Dynamics" (PPD) framework,
understanding or being able to manage them. A combina to accomplish this goaL The PPD framework contains
tion of theory development and multiple research four core components: (1) press and pulse events, (2) a
approaches (place-based, cross-scale, long-term, and com biophysical template, (3) ecosystem services, and (4) a
parative) that harmonize diverse disciplinary perspectives social template. The biophysical and social domains
is needed to develop understanding and build the capacity (areas of study) represent traditional disciplinary research
to sustainably manage social-ecological systems. Here, we paradigms that define processes within each domain. The
propose a new mechanistic framework to guide this biophysical template (eg geology, hydrology) constrains
research, which integrates the internal and interactive fundamental and well-documented relationships between
dynamics of social and natural systems. biotic structure and ecosystem functioning, whereas the
social template (eg legal regulation, social networks)
• "Press-Pulse Dynamics" and ecosystem services: encloses a range of possible human outcomes and behav
an integrated, long-term, social-ecological iors, and the dynamics between them.
research framework In the PPD framework, unlike in other models, the
dynamics of biophysical systems are driven by press and
As noted above, scientists have called repeatedly for pulse events (Smith et al. 2009). Pulse events, such as
greater integration between the social and biophysical floods (both natural and human regulated), are relatively
domains (eg Robertson et al. 2004; Palmer et al. 2005; discrete and rapidly alter species abundances and ecosys
Pickett et al. 2005; Farber er al. 2006; Haberl et al. 2006; Uu tem functioning. Most ecosystems have a characteristic
et al. 2007). Typically, these calls are accompanied by illus natural disturbance regime that includes the size, fre
trative case studies and provide general rationales for why quency, and intensity of pulse disturbances. The natural
such research is needed, yet rarely do they propose useful disturbance regime in most ecosystems has been altered by
roadmaps for implementing truly iritegrated, hypothesis human activities, including those related to species extinc
driven research in social-ecological systems. There is tions, as well as land-use change and management deci-
vices. Though much attention has been dynamics negatively affect most ecosystem services; H4 - changes in vital ecosystem
given to the pattern, if not the process, services alter human outcomes; H5 - changes in human outcomes, such as quality of life
of interaction between the social and or perceptions, affect human behavior; H6 - predictable and unpredictable human
the biophysical systems that represent behavioral responses influence the frequency, magnitude, or form of press and pulse
the extreme example of a human-dom disturbance regimes across ecosystems.
inated world - ie urban and wildland
areas - the PPD framework provides the means for a more making, which affects the biophysical template via
nuanced understanding of social-ecological systems across changes in (1) the intensity of press events and (2) the fre
a continuum of developed to undeveloped lands. This has quency, intensity, and scale of pulse events. Collectively,
important implications for social-ecological science, given altered press and pulse events have quantifiable implica
that the environmental changes of greatest consequence tions for and impacts on ecosystem services, and changes
that are expected in the coming decades will derive from in these services feed back to alter human behaviors and
human migration and population growth on rural and outcomes (Figure l).
quasi-rural lands (Brown et al. 2005). The connectivity Because they represent both quantifiable and qualitative
between places and people across this continuum demands benefits that humans derive from ecosystems, ecosystem
that scientists and managers, for example, understand services form the bridge between the biophysical and
water as a natural hydrologic system that supports human social templates. Ecosystem services can be classified as
life - or fails to do so, depending on how the system is provisioning, regulating, cultural, and supporting (MA
altered and managed. Only with such an integrative under 2005). Provisioning ecosystem services that have markets
standing will it be possible to address (and even resolve) (eg food, fiber, biofuel) have been studied extenSively
the tradeoffs and social equities of differing needs, responsi from the standpoint of enhancing supplies. The same is
bilities, and activities required to sustain humans in their true of certain cultural ecosystem services, notably recre
broader environment. ational ones. But the regulating ecosystem services that
Together, the biophysical and social templates accom maintain essential balance in terrestrial ecosystems - as
modate core disciplinary research activities that feed well as the supporting ecosystem services that enable
information into a larger research framework. In essence, ecosystems to supply other types of services that humans
the model assumes a continuous cycle of human decision experience directly - are much less obvious to people,
The northern Negev Desert is a large and rela Cultural: biodiversity, rare species,
open space, recreation, aesthetics
tively sparsely populated region of an otherwise very
Supporting: primary production,
densely populated country. As such, in national-scale nutrient cycling
land-use planning, it has a central role in future devel
opment, even though its status as a semiarid demo
graphic periphery has made it a relatively unpopular
destination for potential residents (Teschner et 01. Figure 3. Example of the PPD framework for social--ecological research in the Negev
20I 0). Land-use managers have responded by
increasing the region's attractiveness to current and
Desert. Hypotheses are shown in the text box below the figure. Hi - changes in
potential residents through investment in economic landscape characteristics influence ecosystem processes (eg water, soil) and landscape
opportunities (agriculture, industry, tourism) and structure (eg patchiness); H2 - human residential patches (eg farms and neighborhoods)
development of recreational areas (forests, parks, and
reservoirs). Because the region is a transition zone
affect aboveground water flows at the landscape scale; H3 - changes in resource flows
between the arid desert in the south and the influence species diversity; H4 - changes in species diversity affect land-use decision
Mediterranean climate zone in the north, forestry is making; H5 - ecosystem services play a role in determining open space preservation
also promoted as an anti-desertification strategy
(Orlovsky 2008). An additional factor in land-use
policy and biodiversity preservation policy in Israel; H6 - different residential community
decision making is the status of the indigenous types (eg city, town, farm) create unique disturbance regimes on the landscape.
Bedouin population and its contentious relationship
with the state on issues of settlement and grazing/cultivation rights (Yahel 2006; Abu-Saad 2008).
The fundamental relationships we are conceptualizing and analyzing via the PPD model are large-scale (kilometers) and small-scale (meters) land-use
changes, their impact on ecosystem structure and function, the resultant changes in ecosystem service provision, and the responses by the public and policy
makers (and so on in this cyclic relationship).The predominant changes are afforestation with high- and low-density plantings, increased land cultivation, and
expanding residential settlement. Unplanned cultivation and residential development also have important ecological and social implications. The most impor
tant pulses in this semiarid ecosystem are floods and droughts, soil erosion (accompanying floods), dust deposition, and human landscape modifications. The
presses are primarily increased human activities, such as recreational use, landscape conversion by settlements, agriculture, forestry, and grazing (Figure 4). The
landscape is viewed at various scales as a mosaic of patches with distinctive structures that control the flow of materials and energy across the landscape
(Shachak et al. 1998), and changes in disturbance regimes alter the mosaic - and thus the distribution - of materials, energy, and ecosystem services (Figure 3).
Importantly, most of the shifts of ecosystem services in the northern Negev are considered by decision makers as desirable in terms of human quality of
life. Afforestation efforts lead to increased water infiltration and carbon sequestration (Grunzweig et 01. 2003; Rotenberg and Yakir 20I 0), decreased erosion
and airborne dust concentrations, as well as the creation of a network of recreational areas popular with local residents (Ginsberg 2000). On the other hand,
the impact on biodiversity is mixed; plant diversity may increase (Boeken and Shachak 1994), for example, but abundance of local specialist species may
decrease (Shochat et 01.200 I; Hawlena et 01. 20I O). The aesthetic impact is Widely debated, as are the political-demographic implications vis-a-vis the Negev
Bedouin. Policies to increase residential opportunities are politically popular, though residential development, depending on the type, leads to potentially detri
mental impacts on ecosystem function and on biodiversity (Orenstein et 01. 2009).
The PPD framework is assisting researchers and policy makers to conceptualize these multiple, concurrent, and often conflicting impacts and generate
hypotheses regarding how changes in land-use policies will affect different ecosystem service flows. One such study is looking at the ecological implications at
the local and regional scale of low-density residential settlement in the Negev. This
study considers the political-demographic drivers of such human settlement pat
terns (Orenstein and Hamburg 2009), the ecological implications, and the popular
response to perceived changes in the provision of ecosystem services (Orenstein et
01. 2009). Such relationships are investigated at the regional scale (eg northern
Negev) with regard to impact on desert aesthetics and landscape fragmentation, and
at the local scale (eg the wadi, an Arabic term for a dry riverbed or intermittent
stream) on water flow, and rodent, insect, and shrub diversity.
research paradigm has evolved, yet the purpose and gen using social constructs to understand biophysical variations
eral utility of these frameworks vary widely, suggesting over the long term (Walker et al. 2009). Social data are
that they serve multiple goals. Several conceptual frame increasingly spatially explicit (Irwin and Geoghegan
works provide evidence for why such research is needed 2001), which permits novel hypothesis testing and analysis
on topics such as environmental degradation, conserva that is spatially relevant, as well as multi-scaled. Moreover,
tion planning, and sustainability (eg Kremen and Ostfeld ecological research is now commonly conducted at socially
2005; Haberl et al. 2006), but they offer limited informa relevant scales. Eventually, the use of spatial data may lead
tion on how to conduct an integrated research agenda. to unifying theories that view phenomena as integrated
Indeed, some of these models are highly linear and pro social-ecological systems and, with the inclusion of long
vide no clear mechanism for understanding key feedbacks term data and analyses, this will move theory from the
between social and biophysical systems (Kremen and realm of correlations and associations to a deeper probing
Ostfeld 2005; Theobold et al. 2005). Other frameworks of both mechanism and pattern.
describe the necessary components of interactive Biophysical and social scientists examine how systems are
social-ecological systems (Grimm et al. 2000), or focus on organized and the roles played by internal versus external
only a subset of potential interactions, such as economics influences (Pickett et al. 2005). Moving environmental sci
and biodiversity (Fisher et al. 2009; Ohl et al. 2007). ence to a new level of research collaboration, synthesis, and
Often such models lack temporal dynamics or specifics on integration requires a shift from viewing humans as external
how other components of social-ecological systems drivers of natural systems to viewing them as affected agents
should or could be integrated. acting within social-ecological systems (Grimm et al. 2000)
A popular research framework in European social-eco - agents that depend on ecosystem services across a range of
logical research is the "Driving force-Pressure-State scales and feedback cycles. As the human population con
Impact-Response" model (eg Ohl et a l. 2007). This general tinues to grow, with attendant land-use, technological, and
model has similarities to the PPD framework, including key economic changes, it will place additional demands on vital
feedbacks, but it lacks an explicit focus on ecosystem ser ecosystem services (MA 2005). These demands will require
vices. The same is true of Redman et al.'s (2004) social-eco integrated, long-term research that spans multiple disci
logical model, which highlights the areas where social and plines and that will ultimately provide solutions for the
ecological systems intersect without ecosystem services and environment and society. The PPD framework provides an
press-pulse constructs (Ohl and Swinton 2010). By con explicit roadmap to guide this research.
trast, quantifiable ecosystem services explicitly link social
and biophysical systems in the Millennium Ecosystem • Acknowledgements
Assessment (MA 2005) as well as in Daily et al. (2009). But
one key difference is that, like Redman et al. (2004), the All authors contributed equally to the development of
PPD framework emphasizes that human behavior is partly this conceptual framework. Support was provided by
influenced by factors external to ecological feedbacks. National Science Foundation award DEB-0435546 to the
Another major difference is the generalizable set of Long Term Ecological Research Network.
hypotheses within the PPD framework that provide guid
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Scott L Collinsl*, Stephen R Carpenter2, Scott M Swinton3, Daniel E Orenstein4, Daniel L Childers5,
Ted L Gragson6, Nancy B Grimm7, J Morgan Groves, Sharon L Harlan9, Jason P KayelO, Alan K Knappll,
Gary P Kofinas12, John J Magnuson2, William H McDowell13, John M Melackl4, Laura A OgdenlS,
G Philip Robertsonl6, Melinda D Smith17, and Ali C WhitmerlS
IDepartment of Biology, University of New Mexico, Albuquerque, NM *(scollins@sevilleta.unm.edu); 2Center for Limnology,
University of Wisconsin, Madison, WI; 3Department of Agricultural, Food and Resource Economics, Michigan State University, East
Lansing, MI; 4Center for Urban and Regional Studies, Technion - Israel Institute of Technology, Haifa, Israel; 5School of
Sustainability, Arizona State University, Tempe, AZ; 6Department of Anthropology, University of Georgia, Athens, GA; 7School of
Life Sciences, Arizona State University, Tempe, AZ; 8US Forest Service, Northern Research Station, Baltimore, MD; 9School of
Human Evolution and Social Change, Arizona State University, Tempe, AZ; IODepartment of Crop and Soil Science, Pennsylvania
State University, University Park, PA; I IDepartment of Biology and Graduate Degree Program in Ecology, Colorado State University,
Ft Collins, CO; 12Department of Resources Management, University of Alaska, Fairbanks, AK; I3Department of Natural Resources
and the Environment, University of New Hampshire, Durham, NH; 14Bren School of Environmental Science and Management,
University of California, Santa Barbara, Santa Barbara, CA; 15Department of Global and Sociocultural Studies, Florida International
University, Miami, FL; 16Kellogg Biological Station, Michigan State University, Hickory Corners, MI; 17Department of Ecology and
Evolutionary Biology, Yale University, New Haven, CT; 18Georgetown University, Washington, DC