LESSON 2: ENERGY FLOW IN AN ECOSYSTEM AND THE BIOGEOCHEMICAL CYCLES

INTRODUCTION
This lesson discusses the food chain, food web, trophic levels, the biogeochemical or nutrient cycles
including the hydrologic cycle, nitrogen cycle, carbon-oxygen cycle, phosphorous and sulfur cycles.

LEARNING OUTCOMES

At the end of the lesson, you should be able to:

1. illustrate clearly a simple grazing food chain;

2. explain comprehensively the pyramid of energy, pyramid of numbers and pyramid of biomass;
3. discuss systematically the energy pathway from the living to the non-living components through
the various nutrient cycles.

FOOD CHAIN, FOOD WEB AND TROPHIC LEVELS

Observe the illustration below. Explain how the abiotic component of the ecosystem affect the biotic
components.

Figure 1: biotic and abiotic components

Ecosystem
Ecosystem is the community of biotic or living organisms and its environment functioning as an
ecological unit.

Example of an Ecosystem
All the living and nonliving factors inside a pond:
 The water in the pond
 The algae and plants that grow in the water
 The animals and bacteria that live in the water
 The dirt and rocks on the bottom
 The sunlight on the water

Biotic vs. Abiotic Factors

Biotic Factors:
 Living organisms and factors from formerly living organisms
 Include interactions between members of the same species and different species

Abiotic Factors:
 Any nonliving geological, geographical and climatological factors
 Examples: water, air, soil, light, temperature, and natural disasters
Both abiotic and biotic factors can limit or enhance a population’s success in a particular environment.
Changes in Ecosystems
 The survival of organisms depends greatly on physical factors in their environment

Forest Ecology Prepared by J. del

Rosario
  Any changes to the biotic or abiotic factors can impact an ecosystem
 Even minor changes can have a large impact
 Changes in the environment have ongoing effects

Energy Flow in Ecosystems

 Energy flows from the sun through ecosystems from one organism to another
 The sun’s energy cycles through ecosystems from producers to consumers and back into the nutrient
pool through decomposers
 Trophic levels describe the feeding levels of organisms: producers, primary/secondary/tertiary
consumers, and decomposers

Trophic levels
Trophic levels can be represented by numbers, starting at level 1 with plants. Further trophic levels are
numbered subsequently according to how far the organism is along the food chain.
 Level 1: Plants and algae make their own food and are called primary producers.
 Level 2: Herbivores eat plants and are called primary consumers.
 Level 3: Carnivores which eat herbivores are called secondary consumers.
 Level 4: Carnivores which eat other carnivores are called tertiary consumers.
 Level 5: Apex predators which have no predators are at the top of the food chain.

Producers
 Organisms able to make their own food from inorganic compounds using photosynthesis
 Plants, protista (algae), and some bacteria
 At the bottom of all food pyramids, all other organisms depend on producers for energy

Consumers
 Consumers get energy by feeding on producers or other consumers
 Classification depends on location within the food chain:
 Primary = animals that eat producers (herbivores), ex. deer
 Secondary = animals that eat primary consumers (carnivores) or primary consumers and
producers (omnivores), ex. wolves
 Tertiary = eat secondary consumers, can still be carnivores or omnivores

Decomposers
 Organisms that consume dead organisms, releasing nutrients back into the soil, water, and
atmosphere
 Crucial to ecosystems, playing important roles in the carbon, nitrogen, phosphorus, and oxygen
cycles; ex. Fungi
 Decomposers may feed on organisms at any stage of the food chain.
 Decomposers recycle nutrients back into the ecosystem. Ex. worms

Food Chains
 Describe the energy flow between species within an ecosystem
 Producers use energy from the sun to make food and therefore start the chain
 The arrows represent the direction of energy flow, pointing from the organism being consumed to the
organism receiving the energy

Food Chain sample

Figure 2: example of a food chain

Food Webs
 Group or series of interconnected food chains

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  Organisms within a food web can belong to more than one trophic level, or feeding level.

Figure 3: Example of a food web

Energy Pyramid
 A diagram that shows the relative amounts of energy located within each trophic level
 Most of the energy in an energy pyramid is used or lost as heat energy as it moves up the pyramid,
therefore each level in an energy pyramid has less energy available to it than the level below (only
about 10% of the energy produced at each level is available to the one above it)
 Producers are the foundation of all pyramids

Figure 4:
Symbiosis
 An interaction between individuals of different biological species
 One of the organisms receive a benefit from the interaction, the other can either receive a benefit, be
harmed, or not be affected in any way
 Three main kinds of symbiotic relationships: commensalism, mutualism, and parasitism

Symbiosis Chart
Interaction Species A Species B
Commensalism Receives Benefits Not Affected
Mutualism Receives Benefits Receives Benefits
Parasitism Receives Benefits Harmed

Commensalism: One organism benefits and the other is neither helped nor harmed in a commensalistic
relationship.

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Mutualism: Both organisms benefit in a mutualistic relationship. Ex. flowers and pollinators
Parasitism: One organism benefits and the other is harmed in a parasitic relationship. Ex. Mosquitoes
Other Relationships
Predator/Prey: predator hunts, kills, and eats prey
Competition: two organisms compete for the same resources, ex. food, water, shelter, space
Cooperation: an interaction where organisms work together, ex. wolf packs

Biogeochemical Cycles

 Predictable pathways followed by chemical elements or molecules as the elements or molecules travel
through the living and nonliving parts of an ecosystem
 Move among the Earth's biosphere, atmosphere, lithosphere, and hydrosphere
 For example: the carbon-oxygen cycle through photosynthesis and cellular respiration

Nitrogen Cycle
 Nitrogen is an essential component of amino acids (proteins) and nucleic acids (DNA and RNA); all
organisms require nitrogen to survive
 A few microscopic organisms and natural processes, such as lightening, that can convert unusable
nitrogen in the atmosphere to usable forms of nitrogen

Figure 5: Nitrogen Cycle

Primary production

Productivity is the accrual of matter and energy in biomass.

The first step in this process (termed primary productivity) is performed by green plants, which are
the only organisms capable of capturing the electro-magnetic energy of the sun and converting it to the
chemical energy of reduced car-bon compounds (i.e., photosynthates). Secondary productivity results
when heterotrophic organisms consume plant tissues and convert some proportion of that matter and energy
to their own biomass. Secondary producers, which are associated with the detrital and the grazing energy
transfer pathways, compose a small proportion of total forest productivity, but are critically important
regulators of ecosystem processes, particularly nutrient cycling. Gosz et al. (1978) give a relatively thorough
balance sheet for energy transfers in a temperate deciduous forest.
Climate, water and nutrients as a main driving force of primary productivity
The availability of water and nutrients is the major factor governing carbon input to ecosystems.
Photosynthesis is the process by which most carbon and chemical energy enter ecosystems. The proximate
controls over photosynthesis by a single leaf are the availability of reactants such as light energy and CO2;
temperature, which governs reaction rates; and the availability of nitrogen, which is required to produce
photosynthetic enzymes.

Global Carbon Cycle and Productivity

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 The global carbon cycle is a complex set of processes involving three main components: the land; the
oceans; and the atmosphere. Through natural flows, hundreds of billions of tons of carbon are exchanged
with the atmosphere. However, this number pales in comparison to the amount of carbon stored in stocks
(Houghton 2001). The carbon cycle is the biogeochemical cycle by which carbon is exchanged among the
biosphere, pedosphere, geosphere, hydrosphere, and atmosphere of the Earth. Along with the nitrogen cycle
and the water cycle, the carbon cycle comprises a sequence of events that are key to making the Earth
capable of sustaining life; it describes the movement of carbon as it is recycled and reused throughout the
biosphere.
Carbon is also present in the Earth's atmosphere, soils, oceans, and crust. When viewing the Earth as a
system, these components can be referred to as carbon pools (sometimes also called stocks or reservoirs)
because they act as storage houses for large amounts of carbon. Any movement of carbon between these
reservoirs is called a flux. In any integrated system, fluxes connect reservoirs together to create cycles and
feedbacks. An example of such a cycle is seen in the following figure on which the carbon in the atmosphere
is used in photosynthesis to create new plant material.

Figure 7: A sub-cycle within the global carbon cycle. Carbon continuously moves between the atmosphere,
plants and soils through photosynthesis, plant respiration, harvesting, fire and decomposition.

On a global basis, this processes transfers large amounts of carbon from one pool (the atmosphere) to
another (plants). Over time, these plants die and decay, are harvested by humans, or are burned either for
energy or in wildfires. All of these processes are fluxes that can cycle carbon among various pools within
ecosystems and eventually releases it back to the atmosphere. Viewing the Earth as a whole, individual cycles
like this are linked to others involving oceans, rocks, etc. on a range of spatial and temporal scales to form an
integrated global carbon cycle.

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Figure 8: A simplified diagram of the global carbon cycle.
Source: http://earthobservatory.nasa.gov/Library/CarbonCycle/carbon_cycle4.html

On the shortest time scales, of seconds to minutes, plants take carbon out of the atmosphere through
photosynthesis and release it back into the atmosphere via respiration. On longer time scales, carbon from
dead plant material can be incorporated into soils, where it might reside for years, decades or centuries before
being broken down by soil microbes and released back to the atmosphere. On still longer time scales, organic
matter1 that became buried in deep sediments (and protected from decay) was slowly transformed into
deposits of coal, oil and natural gas, the fossil fuels we use today. When we burn these substances, carbon
that has been stored for millions of years is released once again to the atmosphere in the form of carbon
dioxide (CO2)

The carbon cycle has a large effect on the function and well-being of our planet. Globally, the carbon
cycle plays a key role in regulating the Earth’s climate by controlling the concentration of carbon dioxide in
the atmosphere. Carbon dioxide (CO2) is important because it contributes to the greenhouse effect, in which
heat generated from sunlight at the Earth’s surface is trapped by certain gasses and prevented from escaping
through the atmosphere. The greenhouse effect itself is a perfectly natural phenomenon and, without it, the
Earth would be a much colder place. But as is often the case, too much of a good thing can have negative
consequences, and an unnatural buildup of greenhouse gasses can lead to a planet that gets unnaturally hot.
In recent years CO2 has received much attention because its concentration in the atmosphere has risen to
approximately 30% above natural background levels and will continue to rise into the near future. Scientists
have shown that this increase is a result of human activities that have occurred over the last 150 years,
including the burning of fossil fuels and deforestation. Because CO2 is a greenhouse gas, this increase is
believed to be causing a rise in global temperatures. This is the primary cause of climate change and is the
main reason for increasing interest in the carbon cycle. The Earth’s carbon reservoirs naturally act as both
sources, adding carbon to the atmosphere, and sinks, removing carbon from the atmosphere. If all sources
are equal to all sinks, the carbon cycle can be said to be in equilibrium (or in balance) and there is no change
in the size of the pools over time. Maintaining a steady amount of CO2 in the atmosphere helps maintain
stable average temperatures at the global scale. However, because fossil fuel combustion and deforestation
have increased CO2 inputs to the atmosphere without matching increases in the natural sinks that draw CO2
out of the atmosphere (oceans, forests, etc.), these activities have caused the size of the atmospheric carbon
pool to increase. This is what has been responsible for the present buildup of CO2 and is believed to cause
the observed trend of increasing global temperatures. How far will CO2 levels rise in the future? The answer
depends both on how much CO2 humans continue to release and on the future amount of carbon uptake and
storage by the Earth's natural sinks and reservoirs. In short, it depends on the carbon cycle.

Photosynthesis
Photosynthesis is a process used by plants and other organisms to convert light energy, normally from
the sun, into chemical energy that can be used to fuel the organisms' activities. Carbohydrates, such as
sugars, are synthesized from carbon dioxide and water. Oxygen is also released, mostly as a waste product.

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Most plants, most algae, and cyanobacteria perform the process of photosynthesis, and are called
photoautotrophs. Photosynthesis maintains atmospheric oxygen levels and supplies all of the organic
compounds and most of the energy necessary for all life on Earth.
Although photosynthesis is performed differently by different species, the process always begins when
energy from light is absorbed by proteins called reaction centres that contain green chlorophyll pigments. In
plants, these proteins are held inside organelles called chloroplasts, which are most abundant in leaf cells,
while in bacteria they are embedded in the plasma membrane. In these light-dependent reactions, some
energy is used to strip electrons from suitable substances such as water, producing oxygen gas. Furthermore,
two further compounds are generated: reduced nicotinamide adenine dinucleotide phosphate (NADPH) and
adenosine triphosphate (ATP), the "energy currency" of cells.
In plants, algae and cyanobacteria, sugars are produced by a subsequent sequence of light-
independent reactions called the Calvin cycle, but some bacteria use different mechanisms, such as the
reverse Krebs cycle. In the Calvin cycle, atmospheric carbon dioxide is incorporated into already existing
organic carbon compounds, such as ribulose bisphosphate (RuBP). Using the ATP and NADPH produced by
the light-dependent reactions, the resulting compounds are then reduced and removed to form further
carbohydrates such as glucose.

Figure 3.4. A simplified diagram of photosynthesis. Photosynthesis changes sunlight into chemical energy,
splits water to liberate O2, and fixes CO2 into sugar.
(Source: http://en.wikipedia.org/wiki/File:Simple_photosynthesis_overview.svg)

Carbon is a constituent of all terrestrial life. Carbon begins its cycle through forest ecosystems when
plants assimilate atmospheric CO2 through photosynthesis into reduced sugars. Usually about half the gross
photosynthetic products produced (GPP) are expended by plants in autotrophic respiration (Ra) for the
synthesis and maintenance of living cells, releasing CO2 back into the atmosphere. The remaining carbon
products (GPP - Ra) go into net primary production (NPP): foliage, branches, stems, roots, and plant
reproductive organs. As plants shed leaves and roots, or are killed, the dead organic matter forms detritus, a
substrate that supports animals and microbes, which through their heterotrophic metabolism (Rh) release
CO2 back into the atmosphere. On an annual basis, undisturbed forest ecosystems generally show a small
net gain in carbon exchange with the atmosphere. This represents net ecosystem production (NEP). The
ecosystem may lose carbon if photosynthesis is suddenly reduced or when organic materials are removed as a
result of disturbance. Soil humus represents the major accumulation of carbon in most ecosystems because it
remains unoxidized for centuries. It is the most important long-term carbon storage site in ecosystems.

Water Cycle
 Much more water stored in the cycle than moving in the cycle
 Water may be stored for a short time as water vapor in the atmosphere, for days or weeks in a lake, or
for thousands of years in a polar ice cap
 Sun drives the cycle

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Figure 6: Water Cycle

Water cycle in forest ecosystem

The water cycle, also known as the hydrologic cycle or the H2O cycle, describes the continuous
movement of water on, above and below the surface of the Earth. The mass water on Earth remains fairly
constant over time but the partitioning of the water into the major reservoirs of ice, fresh water, saline water
and atmospheric water is variable depending on a wide range of climatic variables. The water moves from one
reservoir to another, such as from river to ocean, or from the ocean to the atmosphere, by the physical
processes of evaporation, condensation, precipitation, infiltration, runoff, and subsurface flow. In so doing,
the water goes through different phases: liquid, solid (ice), and gas (vapor). The water cycle involves the
exchange of energy, which leads to temperature changes. For instance, when water evaporates, it takes up
energy from its surroundings and cools the environment. When it condenses, it releases energy and warms
the environment. These heat exchanges influence climate. The evaporative phase of the cycle purifies water
which then replenishes the land with freshwater. The flow of liquid water and ice transports minerals across
the globe. It is also involved in reshaping the geological features of the Earth, through processes including
erosion and sedimentation. The water cycle is also essential for the maintenance of most life and ecosystems
on the planet.
The hydrologic cycle is an important feature of all ecosystems, and particularly forests, which
generally grow in climates where precipitation provides more water than the vegetation can use or soils can
store. The excess water contributes to stream flow, which provides for irrigation and urban needs far from the
source of precipitation. Vegetation is a major factor in the hydrologic cycle. Before precipitation reaches the
soil, water is intercepted and evaporated from the surface of vegetation and the litter layer. The rate at which
water infiltrates into the soil, runs off the surface, or percolates through to the water table is affected by the
density and depth of root channels and organic residue incorporated into the soil.

Energy and Water

Water and solar energy are essential for the functioning of the Earth System. Since neither is
distributed evenly around the globe, the mechanisms by which they are redistributed (the global hydrologic
cycle and energy budget) are important. These processes are so tightly intertwined that they cannot be treated
separately. Solar energy drives the hydrologic cycle through the vertical transfer of water from Earth to the
atmosphere via evapotranspiration, the sum of evaporation from surfaces and transpiration, which is the
water loss from plants. Conversely, evapotranspiration accounts for 75% of the turbulent energy transfer from
Earth to the atmosphere and is therefore a key process in Earth’s energy budget (see Fig. 4.1). The hydrologic
cycle also controls Earth’s biogeochemical cycles by influencing all biotic processes, dissolving nutrients, and
transferring them within and among ecosystems. These nutrients provide the resources that support growth
of organisms. The movement of materials that are dissolved and suspended in water links ecosystems within
a landscape.

Main components of water balance

Storage - water in soil and aboveground and belowground biomass.

Inputs - Precipitation, Interception, Throughfall, Stemflow, Infiltration, Percolation, Underground water
Outputs - Transpiration, Evaporation Surface Runoff, Base Flow
Precipitation - Condensed water vapor that falls to the Earth's surface . Most precipitation occurs as rain,
but also includes snow, hail, fog drip, graupel, and sleet.
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Canopy interception - The precipitation that is intercepted by plant foliage, eventually evaporates back to
the atmosphere rather than falling to the ground.
Snowmelt - The runoff produced by melting snow.
Runoff - The variety of ways by which water moves across the land. This includes both surface runoff and
channel runoff. As it flows, the water may seep into the ground, evaporate into the air, become stored in lakes
or reservoirs, or be extracted for agricultural or other human uses.
Infiltration - The flow of water from the ground surface into the ground. Once infiltrated, the water becomes
soil moisture or groundwater.
Subsurface flow - The flow of water underground, in the vadose zone and aquifers. Subsurface water may
return to the surface (e.g. as a spring or by being pumped) or eventually seep into the oceans. Water returns
to the land surface at lower elevation than where it infiltrated, under the force of gravity or gravity induced
pressures. Groundwater tends to move slowly, and is replenished slowly, so it can remain in aquifers for
thousands of years.
Evaporation - The transformation of water from liquid to gas phases as it moves from the ground or bodies of
water into the overlying atmosphere. The source of energy for evaporation is primarily solar radiation.
Evaporation often implicitly includes transpiration from plants, though together they are specifically referred
to as evapotranspiration.
Sublimation - The state change directly from solid water (snow or ice) to water vapor.
Deposition - This refers to changing of water vapor directly to ice.
Advection - The movement of water in solid, liquid, or vapor states through the atmosphere. Without
advection, water that evaporated over the oceans could not precipitate over land.
Condensation - The transformation of water vapor to liquid water droplets in the air, creating clouds and fog.
Transpiration - The release of water vapor from plants and soil into the air. Water vapor is a gas that cannot
be seen.
Percolation - Water flows horizontally through the soil and rocks under the influence of gravity

Figure 9. Water balance of an ecosystem (Waring and Running 1998).

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LESSON 3: ANALYSIS OF CHANGES IN FOREST STRUCTURE AND FUNCTION

Objectives:

At the end of the lesson, you should be able to:

1. discuss the resources present in the ecosystem;
2. identify the physical and the chemical properties that affect the activities of organisms in the
forest ecology; and
3. discuss the impact of human activities in the ecosystem.
Observe the picture below (Figure 6). In the given activity sheet, list the function of a forest which you
can observe on the given picture.

Figure 6. Function of a forest

Ecosystem ecology addresses the interactions between organisms and their environment as an
integrated system. The ecosystem approach is fundamental in managing Earth’s resources because it
addresses the interactions that link biotic systems, of which humans are an integral part, with the physical
systems on which they depend. This applies at the scale of Earth as a whole, a continent, or a farmer’s field.
An ecosystem approach is critical to resource management, as we grapple with the sustainable use of
resources in an era of increasing human population and consumption and large, rapid changes in the global
environment.
The flow of energy and materials through organisms and the physical environment provides a
framework for understanding the diversity of form and functioning of Earth’s physical and biological
processes.

Ecosystems analysis
Ecosystem analysis seeks to understand the factors that regulate the pools (quantities) and fluxes
(flows) of materials and energy through ecological systems. These materials include carbon, water, nitrogen,
rock-derived minerals such as phosphorus, and novel chemicals such as pesticides or radionuclides that
people have added to the environment. These materials are found in abiotic (nonbiological) pools such as
soils, rocks, water, and the atmosphere and in biotic pools such as plants, animals, and soil microorganisms.
Francis C. Evans (1956) used the first original definition of ecosystems analyses: Ecosystem analysis is a mix
of biogeo-chemistry, ecophysiology, and micrometeorology that emphasizes "the circulation, trans-formation,
and accumulation of energy and matter through the medium of living things and their activities".
For example, rather than concentrating on the growth of individual trees, the ecosystem ecologist often
expresses forest growth as net primary production in units of kilograms per hectare per year. Ecosystem

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ecology is less concerned with species diversity than with the contribution that any complex of species makes
to the water, carbon, energy, and nutrient transfer on the landscape (Waring, Running 1996).
An initial step in ecosystem analysis is to measure the amount of material stored in different
components of the system, for example, the carbon stored in stem biomass, water stored in the snowpack,
and nutrients stored in the soil. In systems terminology, these are the state variables that can be directly
measured at any given time. Innumerable studies have been published measuring the current state of forest
ecosystems. Frequently, however, the rates of change of these system states, or flows of material, are of
greatest interest. What is the rate of snowmelt, stem biomass accumulation, or nutrient leaching in a
particular system? These questions require study of the processes controlling energy and matter transfer, a
much more difficult undertaking. In these process studies, we wish to identify the cause-effect relationships
controlling system activity, which is often called a mechanistic approach. This identification of system states
and multiple cause-effect relationships that operate in a forest ecosystem to regulate material flows can be
quantified and organized with an ecosystem simulation model. This type of model becomes the starting point
of our space/time scaling of ecosystem principles.
Ecosystem processes can be studied at many spatial scales. How big is an ecosystem? The appropriate
scale of study depends on the question being asked (Fig. 7).

Figure 7. Examples of questions appropriate to each scale (Chapin et al. 2011).

The impact of zooplankton on the algae that they eat might be studied in the laboratory in small
bottles. Other questions such as the controls over productivity might be studied in relatively homogeneous

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patches of a lake, forest, or agricultural field. Still other questions are best addressed at the global scale. The
concentration of atmospheric CO2, for example, depends on global patterns of biotic exchanges of CO2 and
the burning of fossil fuels, which are spatially variable across the globe. The rapid mixing of CO2 in the
atmosphere averages across this variability, facilitating estimates of long-term changes in the total global flux
of carbon between Earth and the atmosphere.

Component Processes of a Comprehensive Ecosystem (Waring, Running 1996).

Energy balance
 Short-wave radiation balance (incoming—outgoing)
 Long-wave radiation balance (incoming—outgoing)
 Sensible heat flux
 Latent heat flux
 Soil heat flux

Water balance
 Precipitation partitioning (snow versus rain)
 Canopy and litter interception and storage
 Soil surface infiltration
 Soil water content
 Subrooting zone outflow
 Hili slope hydrologie routing
 Evaporation
 Transpiration
Carbon balance
 Photosynthesis, gross primary production
 Maintenance respiration
 Growth respiration
 Photosynthate storage
 Net primary production
 Carbon allocation - Leaves, stem/branches, roots, defensive compounds, reproduction
 Phenological timing - Canopy growth/senescence Litlerfall of leaves, turnover of stems and roots
 Decomposition
 Net ecosystem production
Elemental balance
 Sources (atmosphere, rock weathering, biological fixation)
 Soil solution transformation
 Immobilization, nitrification, denitrification
 Mineralization
 Root uptake
 Tissue storage
 Internal recycling
 Volatilization
 Leaching
 Export through harvesting and erosion

Dynamic of processes in forest ecosystems - energy and material transfer

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 Ecosystem structure and functioning are governed by at least five independent control variables.
These state factors, as Jenny and co-workers called them, are climate, parent material (i.e., the rocks that
give rise to soils), topography, potential biota (i.e., the organisms present in the region that could potentially
occupy a site), and time (Jenny 1941, Amundson, Jenny 1991).

Figure 8. The relationship between state factors (outside the circle), interactive controls (inside the circle), and
ecosystem processes. The circle represents the boundary of the ecosystem.

This figure shows in a simplified way how these major factors affect a terrestrial ecosystem and
resulting effects. Of prime importance for the organisms and the ecosystem is the climate, in terms of its
physical and chemical components. Light as a component of the physical climate is necessary for organisms,
in particular plants. Further, the energy coning from light and expressed in temperature or heat is
fundamental as a rate regulator of all biological activities. A part of the physical climate is also water. With its
double importance through its physiological action and its function as a carrier of substances in the plants,
as well as in the whole ecosystem. There is also a chemical dimension to the climate. The air contains not
only gases such as oxygen, carbon dioxide and nitrogen, but also acids such as carbonic and sulfuric acid.
The soil contains mineral or nutrient elements essential to the organisms. Over time there are also changes as
a consequence of Man's actions or from natural causes. The topography or slope determines the incoming
radiation to the ecosystems and also affects the ways water passes through the ecosystem. In addition, there
are mechanical factors acting in and on the ecosystem: wind, fire, grazing and Man's activities, such as
harvesting in fields and forests. Finally, time is an essential factor, sometimes forgotten. It is always a
question of the time perspective in which different factors should be considered - short term vs. long term.
The soil has a key role in terrestrial ecosystems as its properties determine the type of species and
ecosystem that can and will develop under specific climatic regimes.
Ecosystem processes both respond to and control the factors that directly govern their activity. For
example, plants both respond to and influence their light, temperature, and moisture environment. Interactive
controls are factors that both control and are controlled by ecosystem characteristics (Chapin et al. 2011).
Important interactive controls include the supply of resources to support the growth and maintenance of
organisms, modulators that influence the rates of ecosystem processes, disturbance regime, the biotic
community, and human activities.

Resources are the energy and materials in the environment that are used by organisms to support
their growth and maintenance (Field et al. 1992). The acquisition of resources by organisms depletes their
abundance in the environment. In terrestrial ecosystems these resources are spatially separated, being
available primarily either aboveground (light and CO2) or belowground (water and nutrients). Resource supply

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is governed by state factors such as climate, parent material, and topography. It is also sensitive to processes
occurring within the ecosystem. Light availability, for example, depends on climatic elements such as
cloudiness and on topographic position, but is also sensitive to the quantity of shading by vegetation.
Similarly, soil fertility depends on parent material and climate but is also sensitive to ecosystem processes
such as erosional loss of soils after overgrazing and inputs of nitrogen from invading nitrogen-fixing species.
Soil water availability strongly influences species composition in dry climates. Soil water availability also
depends on other interactive controls, such as disturbance regime (e.g., compaction by animals) and the types
of organisms that are present (e.g., the presence or absence of deep-rooted trees such as mesquite that tap
the water table). In aquatic ecosystems, water seldom directly limits the activity of organisms, but light and
nutrients are just as important as on land. Oxygen is a particularly critical resource in aquatic ecosystems
because of its slow rate of diffusion through water.

Modulators are physical and chemical properties that affect the activity of organisms but, unlike
resources, are neither consumed nor depleted by organisms (Field et al. 1992). Modulators include
temperature, pH, redox state of the soil, pollutants, UV radiation, etc. Modulators like temperature are
constrained by climate (a state factor) but are sensitive to ecosystem processes, such as shading and
evaporation. Soil pH likewise depends on parent material and time but also responds to vegetation
composition.
Landscape-scale disturbance by fire, wind, floods, insect outbreaks, and hurricanes is a critical
determinant of the natural structure and process rates in ecosystems (Pickett, White 1985, Sousa 1984). Like
other interactive controls, disturbance regime depends on both state factors and ecosystem processes.
Climate, for example, directly affects fire probability and spread but also influences the types and quantity of
plants present in an ecosystem and therefore the fuel load and flammability of vegetation. Deposition and
erosion during floods shape river channels and influence the probability of future floods. Change in either the
intensity or frequency of disturbance can cause long-term ecosystem change. Woody plants, for example,
often invade grasslands when fire suppression reduces fire frequency. The nature of the biotic community
(i.e., the types of species present, their relative abundances, and the nature of their interactions) can
influence ecosystem processes just as strongly as do large differences in climate or parent material. These
species effects can often be generalized at the level of functional types, which are groups of species that are
similar in their role in community or ecosystem processes. Most evergreen trees, for example, produce leaves
that have low rates of photosynthesis and a chemical composition that deters herbivores. These species make
up a functional type because of their ecological similarity to one another. A gain or loss of key functional types
for example, through introduction or removal of species with important ecosystem effects can permanently
change the character of an ecosystem through changes in resource supply or disturbance regime.
Introduction of nitrogen-fixing trees onto British mine wastes, for example, substantially increases nitrogen
supply and productivity and alters patterns of vegetation development. Invasion by exotic grasses can alter
fire frequency, resource supply, trophic interactions, and rates of most ecosystem processes (D’Antonio,
Vitousek 1992). Elimination of predators by hunting can cause an outbreak of deer that overbrowse their food
supply. The types of species present in an ecosystem depend strongly on other interactive controls, so
functional types respond to and affect most interactive controls and ecosystem processes.
Human activities have an increasing impact on virtually all the processes that govern ecosystem
properties (Vitousek 1994). Our actions influence interactive controls such as water availability, disturbance
regime, and biotic diversity. Humans have been a natural component of many ecosystems for thousands of
years. Since the Industrial Revolution, however, the magnitude of human impact has been so great and so
distinct from that of other organisms that the modern effects of human activities warrant particular attention.
The cumulative impact of human activities extend well beyond an individual ecosystem and affect state
factors such as climate, through changes in atmospheric composition, and potential biota, through the

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introduction and extinction of species. The large magnitude of these effects blurs the distinction between
“independent” state factors and interactive controls at regional and global scales. Human activities are
causing major changes in the structure and functioning of all ecosystems, resulting in novel conditions that
lead to new types of ecosystems. The major human effects are summarized in the next section.

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