FEDERAL UNIVERSITY OF TECHNOLOGY,

P.M.B. 1526, OWERRI, IMO STATE

TERM PAPER

ON

THEORIES AND HYPOTHESIS OF HOLLING

BY

EKE, BETHEL A.

20124760508

COURSE CODE: CST 736

COURSE TITLE: INSECT ECOLOGY

SUBMITTED TO DR. F.O. OJIAKO

JANUARY, 2015

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 TABLE OF CONTENT

CHAPTER ONE

Introduction ………………………………………………………1

Works ………………………………………………………………3

CHAPTER TWO

Theories of Holling…………… …………………………………5

REFERENCES ……………………………………………………20

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 CHAPTER ONE

INTRODUCTION

Crawford Stanley (Buzz) Holling, OC (born December 6, 1930) is a

Canadian ecologist, and Emeritus Eminent Scholar and Professor in
Ecological Sciences at the University of Florida. Holling is one of the
conceptual founders of ecological economics (Evans, 2008).

Biography

Crawford Stanley Holling was born in 1930 in the United States to

Canadian parents. He grew up in Northern Ontario, which was
where he first became interested in nature. As a teenager he was a
member of the Royal Ontario Museum's Toronto Junior Field
Naturalists (Bremner and Taplin, 2004).

Holling received his B.A. and M.Sc. at the University of Toronto in

1952 and his Ph.D. at the University of British Columbia in 1957.
He worked for several years in the Canadian Department of Forestry
in Sault Ste. Marie, Ontario.

After working for Forestry Canada, Buzz Holling was, at various

times, Professor and Director of the Institute of Animal Resource
Ecology, University of British Columbia, Director of the
International Institute for Applied Systems Analysis in Vienna, and
Eminent Scholar, Arthur R. Marshall Jr. Chair in Ecological
Sciences in the Department of Zoology at the University of Florida.

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He retired from the University of Florida in 1999, but remains on
the faculty as an Emeritus Eminent Scholar. He currently lives in
Nanaimo, British Columbia, Canada (Brooks et al, 2004).

He has been awarded two major awards from the Ecological Society
of America, the Mercer Award given to a young scientist in
recognition of an outstanding paper in ecology in 1966, and the
Eminent Ecologist Award for "outstanding contributions to the
science of Ecology" in 1999. He also received the Kenneth Boulding
Memorial Prize, in 2000, the Volvo Environment Prize in 2008, an
Honorary Doctor of Science from the University of Guelph in 1998,
and an Honorary Doctor of Science from the Simon Fraser
University in 2011. He is a Fellow of the Royal Society of Canada, a
foreign Fellow of the Royal Swedish Academy of Sciences, and has
been awarded the Austrian Cross of Honour for Science and Art. In
2009, he was made an Officer of the Order of Canada "for his
pioneering contributions to the field of ecology, notably for his work
on ecosystem dynamics, resilience theory and ecological economics"
(Carreiro and Zipper, 2011).

He was founding editor-in-chief of the open access on-line journal

Conservation Ecology, now renamed Ecology and Society. He was
also the founder of the Resilience Alliance, an international science
network (Karkkainen, 2005).

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Works

Throughout his research, C. S. Holling has blended systems theory

and ecology with simulation modeling and policy analysis to develop
integrative theories of change that have practical utility. He has
introduced important ideas in the application of ecology and
evolution, including resilience, adaptive management, the adaptive
cycle, and panarchy (Jones et al, 1994).

His early work included major contributions to population and

behavioural ecology. Later, he was among the first ecologists to
recognize the importance of nonlinear dynamics. This early work on
predation led to a series of papers, including his 1959 Citation
Classic paper in the Canadian Entomologist, in which he developed
the notion of functional response (the relationship between prey
density and the rate at which prey is eaten), an idea that continues
to be a linchpin of modern population ecology (Fischer et al, 2008).

His 1973 paper on the resilience of ecological systems had a

substantial impact within ecology and other natural and social
sciences. He has also contributed important ideas to ecological
management, including Adaptive management and the Adaptive
Cycle. More recently his work on the cross-scale structure and
dynamics of ecosystems has been highly influential. This work
resulted in the 2002 book Panarchy: understanding
transformations in human and natural systems (Skillen and
Maurer, 2008).

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His work is frequently cited in the fields of ecology, environmental
management, ecological economics and the human dimensions of
global change.

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 CHAPTER TWO

THEORIES OF HOLLING

SYSTEMS THEORY

Systems theory is the interdisciplinary study of systems in general,

with the goal of discovering patterns and elucidating principles that
can be discerned from, and applied to, all types of systems at all
nesting levels in all fields of research. Systems theory can
reasonably be considered a specialization of systems thinking or as
the goal output of systems science and systems engineering, with
an emphasis on generality useful across a broad range of systems
(versus the particular models of individual fields).

A central topic of systems theory is self-regulating systems, i.e.

systems self-correcting through feedback. Self-regulating systems
are found in nature, including the physiological systems of our
body, in local and global ecosystems, and in climate—and in human
learning processes (from the individual on up through international
organizations like the UN) (Chipman et al, 1998).

ECOLOGICAL RESILIENCE

In ecology, resilience is the capacity of an ecosystem to respond to a

perturbation or disturbance by resisting damage and recovering
quickly. Such perturbations and disturbances can include
stochastic events such as fires, flooding, windstorms, insect
population explosions, and human activities such as deforestation,
fracking of the ground for oil extraction, pesticide sprayed in soil,
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and the introduction of exotic plant or animal species. Disturbances
of sufficient magnitude or duration can profoundly affect an
ecosystem and may force an ecosystem to reach a threshold beyond
which a different regime of processes and structures predominates.
Human activities that adversely affect ecosystem resilience such as
reduction of biodiversity, exploitation of natural resources,
pollution, land-use, and anthropogenic climate change are
increasingly causing regime shifts in ecosystems, often to less
desirable and degraded conditions (Brunckhorst, 2002).
Interdisciplinary discourse on resilience now includes consideration
of the interactions of humans and ecosystems via socio-ecological
systems, and the need for shift from the maximum sustainable yield
paradigm to environmental resource management which aims to
build ecological resilience through "resilience analysis, adaptive
resource management, and adaptive governance".

The concept of resilience in ecological systems was first introduced

by the Canadian ecologist C.S. Holling in order to describe the
persistence of natural systems in the face of changes in ecosystem
variables due to natural or anthropogenic causes. Resilience has
been defined in two ways in ecological literature:

 as the time required for an ecosystem to return to an

equilibrium or steady-state following a perturbation (which is
also defined as stability by some authors). This definition of
resilience is used in other fields such as physics and

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 engineering, and hence has been termed ‘engineering
resilience’ by Holling.
 as "the capacity of a system to absorb disturbance and
reorganize while undergoing change so as to still retain
essentially the same function, structure, identity, and
feedbacks" (Holling et al, 2002).

The second definition has been termed ‘ecological resilience’, and it

presumes the existence of multiple stable states or regimes.

Some shallow temperate lakes can exist within either clear water
regime, which provides many ecosystem services, or a turbid water
regime, which provides reduced ecosystem services and can
produce toxic algae blooms. The regime or state is dependent upon
lake phosphorus cycles, and either regime can be resilient
dependent upon the lake's ecology and management.

Mulga woodlands of Australia can exist in a grass-rich regime that

supports sheep herding, or a shrub-dominated regime of no value
for sheep grazing. Regime shifts are driven by the interaction of fire,
herbivory, and variable rainfall. Either state can be resilient
dependent upon management (Brooks et al, 2004).

Theory

Ecologists Brian Walker, C S Holling and others describe four

critical aspects of resilience: latitude, resistance, precariousness,
and panarchy.

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The first three can apply both to a whole system or the sub-systems
that make it up.

1. Latitude: the maximum amount a system can be changed

before losing its ability to recover (before crossing a threshold
which, if breached, makes recovery difficult or impossible).
2. Resistance: the ease or difficulty of changing the system; how
“resistant” it is to being changed.
3. Precariousness: how close the current state of the system is to
a limit or “threshold.”
4. Panarchy: the degree to which a certain hierarchical level of an
ecosystem is influenced by other levels. For example,
organisms living in communities that are in isolation from one
another may be organized differently from the same type of
organism living in a large continuous population, thus the
community-level structure is influenced by population-level
interactions (Evans, 2008).

Closely linked to resilience is adaptive capacity, which is the

property of an ecosystem that describes change in stability
landscapes and resilience. Adaptive capacity in socio-ecological
systems refers to the ability of humans to deal with change in their
environment by observation, learning and altering their interactions
(Scheffer et al, 2009).

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PANARCHY

Panarchy is a conceptual term first coined by the Belgian

philosopher, economist, and botanist Paul Emile de Puydt in 1860,
referring to a specific form of governance (-archy) that would
encompass (pan-) all others. The Oxford English Dictionary lists the
noun as "chiefly poetic" with the meaning "a universal realm," citing
an 1848 attestation by Philip James Bailey, "the starry panarchy of
space". The adjective panarchic "all-ruling" has earlier attestations.
In the twentieth century the term was re-coined separately by
scholars in international relations to describe the notion of global
governance and then by systems theorists to describe non-
hierarchical organizing theories (Bremner and Taplin, 2004).

The concept of panarchy provides a framework that characterizes

complex systems of people and nature as dynamically organized
and structured within and across scales of space and time. It has
been more than a decade since the introduction of panarchy.

Humans build mental models of complex systems to make their

structures and dynamics tractable for scientific inquiry.
Multidimensional, nonlinear processes and structures characterize
complex systems, including ecological, social, or coupled social–
ecological systems. Nevertheless, these systems are amenable to
simplification. Panarchy is a conceptual model that describes the
ways in which complex systems of people and nature are
dynamically organized and structured across scales of space and
time. Panarchy uses a systems approach to understand ecosystem

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dynamics and emphasizes hierarchical structuring (Bremner and
Taplin, 2004).

However, panarchy is different from typically envisioned hierarchies

in that control is not just exerted by larger-scale, top-down
processes, but can also come from small scale or bottom-up
processes.

Additionally, the dynamics of renewal and collapse within-scale

domains, that is, adaptive cycles differ from the more static view of
traditional hierarchy theory. Because of the potential for cycling
within adaptive cycles to affect both smaller scales and larger
scales, panarchy theory emphasizes cross-scale linkages whereby
processes at one scale affect those at other scales to influence the
overall dynamics of the system.

Much of the treatment of the concept of panarchy currently in the

literature is focused on a metaphorical use of the term, but some
core concepts have been tested and others are testable with current
data. Most of the existing empirical tests revolve around
discontinuities and their detection in vertebrate body mass
distributions. However, discontinuities have been tested for and
documented in other organisms and social systems (Benson and
Garmestani, 2011), and the ideas underlying panarchy related to
discontinuities, have been sustained. In addition, limited modeling
and empirical tests have demonstrated a strong link between
discontinuities and the grouping of variables they identify, and
scale specific structure in the environment (Allen and Holling,

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2008). Relatively long-term data sets are becoming available that
have allowed for novel approaches to detect discontinuities and
scaling in temporal variables.

ADAPTIVE MANAGEMENT

Adaptive management (AM), also known as adaptive resource

management (ARM), is a structured, iterative process of robust
decision making in the face of uncertainty, with an aim to reducing
uncertainty over time via system monitoring. In this way, decision
making simultaneously meets one or more resource management
objectives and, either passively or actively, accrues information
needed to improve future management. Adaptive management is a
tool which should be used not only to change a system, but also to
learn about the system. Because adaptive management is based on
a learning process, it improves long-run management outcomes.
The challenge in using the adaptive management approach lies in
finding the correct balance between gaining knowledge to improve
management in the future and achieving the best short-term
outcome based on current knowledge (Angeler and Johnson, 2012).

Objectives

There are a number of scientific and social processes which are vital
components of adaptive management, including:

1. Management is linked to appropriate temporal and spatial

scales
2. Management retains a focus on statistical power and controls

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 3. Use of computer models to build synthesis and an embodied
ecological consensus
4. Use of embodied ecological consensus to evaluate strategic
alternatives
5. Communication of alternatives to political arena for
negotiation of a selection

The achievement of these objectives requires an open management

process which seeks to include past, present and future
stakeholders. Adaptive management needs to at least maintain
political openness, but usually aims to create it. Adaptive
management must therefore be a scientific and social process. It
must focus on the development of new institutions and institutional
strategies in balance with scientific hypothesis and experimental
frameworks (www.resilliance.org).

Adaptive management can proceed as either passive adaptive

management or active adaptive management, depending on how
learning takes place. Passive adaptive management values learning
only insofar as it improves decision outcomes (i.e. passively), as
measured by the specified utility function. In contrast, active
adaptive management explicitly incorporates learning as part of the
objective function, and hence, decisions which improve learning are
valued over those which do not. In both cases, as new knowledge is
gained, the models are updated and optimal management strategies
are derived accordingly. Thus, while learning occurs in both cases,
it is treated differently. Often, deriving actively adaptive policies is

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technically very difficult, which prevents it being more commonly
applied (Somer et al, 1986).

FUNCTIONAL RESPONSE

A functional response in ecology is the intake rate of a consumer as

a function of food density. It is associated with the numerical
response, which is the reproduction rate of a consumer as a
function of food density. Following C. S. Holling, functional
responses are generally classified into three types, which are called
Holling's type I, II, and III.

Type I

Type I functional response assumes a linear increase in intake rate

with food density, either for all food densities, or only for food
densities up to a maximum, beyond which the intake rate is
constant. The linear increase assumes that the time needed by the
consumer to process a food item is negligible, or that consuming
food does not interfere with searching for food. A functional
response of type I is used in the Lotka–Volterra predator–prey
model. It was the first kind of functional response described and is
also the simplest of the three functional responses currently
detailed (Bessey, 2002).

Type II

Type II functional response is characterized by a decelerating intake

rate, which follows from the assumption that the consumer is
limited by its capacity to process food. Type II functional response is
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often modeled by a rectangular hyperbola, for instance as by
Holling's disc equation, which assumes that processing of food and
searching for food are mutually exclusive behaviors. The equation is

where f denotes intake rate and R denotes food (or resource)

density. The rate at which the consumer encounters food items per
unit of food density is called the attack rate, a. The average time
spent on processing a food item is called the handling time, h.
Similar equations are the Monod equation for the growth of
microorganism and the Michaelis–Menten equation for the rate of
enzymatic reactions (Bessey, 2002).

In an example with wolves and caribou, as the number of caribou

increases while holding wolves constant, the number of caribou
kills increases and then levels off. This is because the proportion of
caribou killed per wolf decreases as caribou density increases. The
higher the density of caribou, the smaller the proportion of caribou
killed per wolf. Explained slightly differently, at very high caribou
densities, wolves need very little time to find prey and spend almost
all their time handling prey and very little time searching. Wolves
are then satiated and the total number of caribou kills reaches a
plateau.

Type III

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Type III functional response is similar to type II in that at high
levels of prey density, saturation occurs. But now, at low prey
density levels, the graphical relationship of number of prey
consumed and the density of the prey population is a more than
linearly increasing function of prey consumed by predators. This
accelerating function is large descriptive, and often justified by
learning time, prey switching, or a combination of both phenomena,
but the type III functional response lacks the rigorous theoretical
underpinning of the type II functional response (Bessey, 2002).

Learning time is defined as the natural improvement of a predator’s

searching and attacking efficiency or the natural improvement in
their handling efficiency as prey density increases. Imagine a prey
density so small that the chance of a predator encountering that
prey is extremely low. Because the predator finds prey so
infrequently, it has not had enough experience to develop the best
ways to capture and subdue that species of prey. Holling identified
this mechanism in shrews and deer mice feeding on sawflies. At low
numbers of sawfly cocoons per acre, deer mice especially
experienced exponential growth in terms of the number of cocoons
consumed per individual as the density of cocoons increased. The
characteristic saturation point of the type III functional response
was also observed in the deer mice. At a certain density of cocoons
per acre, the consumption rate of the deer mice reached a
saturation amount as the cocoon density continued to increase
(Bessey, 2002).

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Prey switching involves two or more prey species and one predator
species. When all prey species are at equal prey densities, the
predator will indiscriminately select between prey species. However,
if the density of one of the prey species decreases, then the predator
will start selecting the other, more common prey species with a
higher frequency. Murdoch demonstrated this effect with guppy
preying on tubificids and fruit flies. As fruit fly numbers decreased
guppies switched from feeding on the fruit flies on the water’s
surface to feeding on the more abundant tubificids along the bed
(Bessey, 2002).

ECOLOGICAL ECONOMICS

Ecological economics/eco-economics refers to both a

transdisciplinary and interdisciplinary field of academic research
that aims to address the interdependence and coevolution of
human economies and natural ecosystems over time and space.It is
distinguished from environmental economics, which is the
mainstream economic analysis of the environment, by its treatment
of the economy as a subsystem of the ecosystem and its emphasis
upon preserving natural capital. One survey of German economists
found that ecological and environmental economics are different
schools of economic thought, with ecological economists
emphasizing strong sustainability and rejecting the proposition that
natural capital can be substituted by human-made capital (Farrell
and Turning-Ward, 2004).

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Ecological economics was founded as a modern movement in the
works of and interactions between various European and American
academics (see the section on history and development below). The
related field of green economics is, in general, a more politically
applied form of the subject.

According to ecological economist Malte Faber, ecological economics

is defined by its focus on nature, justice, and time. Issues of
intergenerational equity, irreversibility of environmental change,
uncertainty of long-term outcomes, and sustainable development
guide ecological economic analysis and valuation. Ecological
economists have questioned fundamental mainstream economic
approaches such as cost-benefit analysis, and the separability of
economic values from scientific research, contending that
economics is unavoidably normative rather than positive (i.e.
descriptive). Positional analysis, which attempts to incorporate time
and justice issues, is proposed as an alternative. Ecological
economics shares many of its perspectives with feminist economics,
including the focus on sustainability, nature, justice and care
values (Farrell and Turning-Ward, 2004).

ENVIRONMENTAL RESOURCE MANAGEMENT

Environmental resource management is the management of the

interaction and impact of human societies on the environment. It is
not, as the phrase might suggest, the management of the
environment itself. Environmental resources management aims to
ensure that ecosystem services are protected and maintained for

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future human generations, and also maintain ecosystem integrity
through considering ethical, economic, and scientific (ecological)
variables. Environmental resource management tries to identify
factors affected by conflicts that rise between meeting needs and
protecting resources. It is thus linked to environmental protection
and sustainability.

Environmental resource management is an issue of increasing

concern, as reflected in its prevalence in seminal texts influencing
global socio-political frameworks such as the Brundtland
Commission's Our Common Future, which highlighted the
integrated nature of environment and international development
and the Worldwatch Institute's annual State of the World (book
series) reports (Brooks et al, 2004).

The environment determines nature of every objects around the

sphere. The behaviour, type of religion, culture and economic
practices.

Scope

Improved agricultural practices such as these terraces in northwest

Iowa can serve to preserve soil and improve water quality

Environmental resource management can be viewed from a variety

of perspectives. Environmental resource management involves the
management of all components of the biophysical environment,
both living (biotic) and non-living (abiotic). This is due to the
interconnected and network of relationships amongst all living

18
species and their habitats. The environment also involves the
relationships of the human environment, such as the social,
cultural and economic environment with the biophysical
environment. The essential aspects of environmental resource
management are ethical, economical, social, and technological.
These underlie principles and help make decisions.

The concept of environmental determinism, probabilism and

possibilism are significant in the concept of environmental resource
management.

It should be noted that environmental resource management covers

many areas in the field of science: geography, biology, physics,
chemistry, sociology, psychology, physiology, etc (Brooks et al,
2004).

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