Introduction - What is an Ecosystem?

An ecosystem consists of the biological community that occurs in some locale, and the
physical and chemical factors that make up its non-living or abiotic environment. There are
many examples of ecosystems -- a pond, a forest, an estuary, a grassland. The boundaries
are not fixed in any objective way, although sometimes they seem obvious, as with the
shoreline of a small pond. Usually the boundaries of an ecosystem are chosen for practical
reasons having to do with the goals of the particular study.
The study of ecosystems mainly consists of the study of certain processes that link the
living, or biotic, components to the non-living, or abiotic, components. Energy
transformations andbiogeochemical cycling are the main processes that comprise the
field of ecosystem ecology. As we learned earlier, ecology generally is defined as the
interactions of organisms with one another and with the environment in which they occur.
We can study ecology at the level of the individual, the population, the community, and the
ecosystem.
Studies of individuals are concerned mostly about physiology, reproduction, development
or behavior, and studies of populations usually focus on the habitat and resource needs of
individual species, their group behaviors, population growth, and what limits their
abundance or causes extinction. Studies of communities examine how populations of many
species interact with one another, such as predators and their prey, or competitors that
share common needs or resources.
In ecosystem ecology we put all of this together and, insofar as we can, we try to
understand how the system operates as a whole. This means that, rather than worrying
mainly about particular species, we try to focus on major functional aspects of the system.
These functional aspects include such things as the amount of energy that is produced by
photosynthesis, how energy or materials flow along the many steps in a food chain, or what
controls the rate of decomposition of materials or the rate at which nutrients are recycled in
the system.

Components of an Ecosystem
You are already familiar with the parts of an ecosystem. You have learned about climate and
soils from past lectures. From this course and from general knowledge, you have a basic
understanding of the diversity of plants and animals, and how plants and animals and
microbes obtain water, nutrients, and food. We can clarify the parts of an ecosystem by
listing them under the headings "abiotic" and "biotic".
ABIOTIC COMPONENTS
Sunlight
Temperature
Precipitation
Water or moisture
Soil or water chemistry (e.g., P, NH4+)
etc.
All of these vary over

BIOTIC COMPONENTS
Primary producers
Herbivores
Carnivores
Omnivores
Detritivores
etc.
space/time

By and large, this set of environmental factors is important almost everywhere, in all
ecosystems.
Usually, biological communities include the "functional groupings" shown above.
A functional group is a biological category composed of organisms that perform mostly
the same kind of function in the system; for example, all the photosynthetic plants or
primary producers form a functional group. Membership in the functional group does not
depend very much on who the actual players (species) happen to be, only on what function
they perform in the ecosystem.

Processes of Ecosystems
This figure with the plants, zebra, lion, and so forth illustrates the two main ideas about how
ecosystems function: ecosystems have energy flows and ecosystems cycle materials.
These two processes are linked, but they are not quite the same (see Figure 1).

Figure 1. Energy flows and material cycles.

Energy enters the biological system as light energy, or photons, is transformed into
chemical energy in organic molecules by cellular processes including photosynthesis and
respiration, and ultimately is converted to heat energy. This energy is dissipated, meaning it
is lost to the system as heat; once it is lost it cannot be recycled. Without the continued
input of solar energy, biological systems would quickly shut down. Thus the earth is
an open system with respect to energy.
Elements such as carbon, nitrogen, or phosphorus enter living organisms in a variety of
ways. Plants obtain elements from the surrounding atmosphere, water, or soils. Animals
may also obtain elements directly from the physical environment, but usually they obtain
these mainly as a consequence of consuming other organisms. These materials are

transformed biochemically within the bodies of organisms, but sooner or later, due to
excretion or decomposition, they are returned to an inorganic state. Often bacteria complete
this process, through the process called decomposition or mineralization (see previous
lecture on microbes).
During decomposition these materials are not destroyed or lost, so the earth is a closed
systemwith respect to elements (with the exception of a meteorite entering the system
now and then). The elements are cycled endlessly between their biotic and abiotic states
within ecosystems. Those elements whose supply tends to limit biological activity are
called nutrients.

The Transformation of Energy

The transformations of energy in an ecosystem begin first with the input of energy from the
sun. Energy from the sun is captured by the process of photosynthesis. Carbon dioxide is
combined with hydrogen (derived from the splitting of water molecules) to produce
carbohydrates (CHO). Energy is stored in the high energy bonds of adenosine triphosphate,
or ATP (see lecture on photosynthesis).
The prophet Isaah said "all flesh is grass", earning him the title of first ecologist, because
virtually all energy available to organisms originates in plants. Because it is the first step in
the production of energy for living things, it is called primary production (click here for a
primer on photosynthesis). Herbivores obtain their energy by consuming plants or plant
products,carnivores eat herbivores, and detritivores consume the droppings and
carcasses of us all.

Figure 2 portrays a simple food chain, in which energy

from the sun, captured by plant photosynthesis, flows
fromtrophic level to trophic level via the food chain.
A trophic level is composed of organisms that make a
living in the same way, that is they are all primary
producers (plants),primary consumers (herbivores)
or secondary consumers (carnivores). Dead tissue
and waste products are produced at all levels.
Scavengers, detritivores, and decomposers collectively
account for the use of all such "waste" -- consumers of
carcasses and fallen leaves may be other animals, such
as crows and beetles, but ultimately it is the microbes
that finish the job of decomposition. Not surprisingly,
the amount of primary production varies a great deal
from place to place, due to differences in the amount of
solar radiation and the availability of nutrients and
water.
For reasons that we will explore more fully in
subsequent lectures, energy transfer through the
food chain is inefficient. This means that less energy
is available at the herbivore level than at the primary

producer level, less yet at the carnivore level, and so on. The result is a pyramid of energy,
with important implications for understanding the quantity of life that can be supported.
Usually when we think of food chains we visualize green plants, herbivores, and so on.
These are referred to asgrazer food chains, because living plants are directly consumed.
In many circumstances the principal energy input is not green plants but dead organic
matter. These are called detritus food chains. Examples include the forest floor or a
woodland stream in a forested area, a salt marsh, and most obviously, the ocean floor in
very deep areas where all sunlight is extinguished 1000's of meters above. In subsequent
lectures we shall return to these important issues concerning energy flow.
Finally, although we have been talking about food chains, in reality the organization of
biological systems is much more complicated than can be represented by a simple "chain".
There are many food links and chains in an ecosystem, and we refer to all of these linkages
as a food web. Food webs can be very complicated, where it appears that "everything is
connected to everything else", and it is important to understand what are the most
important linkages in any particular food web.

Biogeochemistry
How can we study which of these linkages in a food web are most important? One obvious
way is to study the flow of energy or the cycling of elements. For example, the cycling of
elements is controlled in part by organisms, which store or transform elements, and in part
by the chemistry and geology of the natural world. The term Biogeochemistry is defined
as the study of how living systems influence, and are controlled by, the geology and
chemistry of the earth. Thus biogeochemistry encompasses many aspects of the abiotic and
biotic world that we live in.
There are several main principles and tools that biogeochemists use to study earth
systems. Most of the major environmental problems that we face in our world toady can be
analyzed using biogeochemical principles and tools. These problems include global warming,
acid rain, environmental pollution, and increasing greenhouse gases. The principles and
tools that we use can be broken down into 3 major components: element ratios, mass
balance, and element cycling.
1. Element ratios
In biological systems, we refer to important elements as "conservative". These elements
are often nutrients. By "conservative" we mean that an organism can change only slightly
the amount of these elements in their tissues if they are to remain in good health. It is
easiest to think of these conservative elements in relation to other important elements in
the organism. For example, in healthy algae the elements C, N, P, and Fe have the following
ratio, called theRedfield ratio after the oceanographer who discovered it:
C : N : P : Fe = 106 : 16 : 1 : 0.01
Once we know these ratios, we can compare them to the ratios that we measure in
a sample of algae to determine if the algae are lacking in one of these limiting

nutrients.

2. Mass Balance
Another important tool that biogeochemists use is a simple mass balance equation to
describe the state of a system. The system could be a snake, a tree, a lake, or the entire
globe. Using a mass balance approach we can determine whether the system is changing
and how fast it is changing. The equation is:
NET CHANGE = INPUT + OUTPUT + INTERNAL CHANGE
In this equation the net change in the system from one time period to another is
determined by what the inputs are, what the outputs are, and what the internal change in
the system was. The example given in class is of the acidification of a lake, considering the
inputs and outputs and internal change of acid in the lake.

3. Element Cycling
Element cycling describes where and how fast elements move in a system. There are two
general classes of systems that we can analyze, as mentioned above: closed and open
systems.
A closed system refers to a system where the inputs and outputs are negligible compared
to the internal changes. Examples of such systems would include a bottle, or our entire
globe. There are two ways we can describe the cycling of materials within this closed
system, either by looking at the rate of movement or at the pathways of movement.

1.
2.

Rate = number of cycles / time * as rate increases, productivity increases

Pathways-important because of different reactions that may occur

In an open system there are inputs and outputs as well as the internal cycling. Thus we
can describe the rates of movement and the pathways, just as we did for the closed system,
but we can also define a new concept called the residence time. The residence time
indicates how long on average an element remains within the system before leaving the
system.

1.
2.
3.

Rate
Pathways
Residence time, Rt

Rt = total amount of matter / output rate of matter

(Note that the "units" in this calculation must cancel properly)

Controls on Ecosystem Function

Now that we have learned something about how ecosystems are put together and how
materials and energy flow through ecosystems, we can better address the question of "what
controls ecosystem function"? There are two dominant theories of the control of
ecosystems. The first, called bottom-up control, states that it is the nutrient supply to the
primary producers that ultimately controls how ecosystems function. If the nutrient supply
is increased, the resulting increase in production of autotrophs is propagated through the
food web and all of the other trophic levels will respond to the increased availability of food
(energy and materials will cycle faster).
The second theory, called top-down control, states that predation and grazing by higher
trophic levels on lower trophic levels ultimately controls ecosystem function. For example, if
you have an increase in predators, that increase will result in fewer grazers, and that
decrease in grazers will result in turn in more primary producers because fewer of them are
being eaten by the grazers. Thus the control of population numbers and overall productivity
"cascades" from the top levels of the food chain down to the bottom trophic levels.
So, which theory is correct? Well, as is often the case when there is a clear dichotomy to
choose from, the answer lies somewhere in the middle. There is evidence from many
ecosystem studies that BOTH controls are operating to some degree, but that NEITHER
control is complete. For example, the "top-down" effect is often very strong at trophic levels
near to the top predators, but the control weakens as you move further down the food
chain. Similarly, the "bottom-up" effect of adding nutrients usually stimulates primary
production, but the stimulation of secondary production further up the food chain is less
strong or is absent.
Thus we find that both of these controls are operating in any system at any time, and we
must understand the relative importance of each control in order to help us to predict how
an ecosystem will behave or change under different circumstances, such as in the face of a
changing climate.

The Geography of Ecosystems

There are many different ecosystems: rain forests and tundra, coral reefs and ponds,
grasslands and deserts. Climate differences from place to place largely determine the types
of ecosystems we see. How terrestrial ecosystems appear to us is influenced mainly by the
dominant vegetation.
The word "biome" is used to describe a major vegetation type such as tropical rain forest,
grassland, tundra, etc., extending over a large geographic area (Figure 3). It is never used
for aquatic systems, such as ponds or coral reefs. It always refers to a vegetation category
that is dominant over a very large geographic scale, and so is somewhat broader than an
ecosystem.

Figure 3: The distribution of biomes.

We can draw upon previous lectures to remember that temperature and rainfall patterns for
a region are distinctive. Every place on earth gets the same total number of hours of
sunlight each year, but not the same amount of heat. The sun's rays strike low latitudes
directly but high latitudes obliquely. This uneven distribution of heat sets up not just
temperature differences, but global wind and ocean currents that in turn have a great deal
to do with where rainfall occurs. Add in the cooling effects of elevation and the effects of
land masses on temperature and rainfall, and we get a complicated global pattern of
climate.
A schematic view of the earth shows that, complicated though climate may be, many
aspects are predictable (Figure 4). High solar energy striking near the equator ensures
nearly constant high temperatures and high rates of evaporation and plant transpiration.
Warm air rises, cools, and sheds its moisture, creating just the conditions for a tropical rain
forest. Contrast the stable temperature but varying rainfall of a site in Panama with the
relatively constant precipitation but seasonally changing temperature of a site in New York
State. Every location has a rainfall- temperature graph that is typical of a broader region.

Figure 4. Climate patterns affect biome distributions.

We can draw upon plant physiology to know that certain plants are distinctive of certain
climates, creating the vegetation appearance that we call biomes. Note how well the
distribution of biomes plots on the distribution of climates (Figure 5). Note also that some
climates are impossible, at least on our planet. High precipitation is not possible at low
temperatures -- there is not enough solar energy to power the water cycle, and most water
is frozen and thus biologically unavailable throughout the year. The high tundra is as much
a desert as is the Sahara.

Figure 5. The distribution of biomes related to temperature and precipitation.

Summary

Ecosystems are made up of abiotic (non-living, environmental) and biotic

components, and these basic components are important to nearly all types of
ecosystems. Ecosystem Ecology looks at energy transformations and biogeochemical
cycling within ecosystems.
Energy is continually input into an ecosystem in the form of light energy, and some
energy is lost with each transfer to a higher trophic level. Nutrients, on the other
hand, are recycled within an ecosystem, and their supply normally limits biological
activity. So, "energy flows, elements cycle".
Energy is moved through an ecosystem via a food web, which is made up of
interlocking food chains. Energy is first captured by photosynthesis (primary
production). The amount of primary production determines the amount of energy
available to higher trophic levels.
The study of how chemical elements cycle through an ecosystem is termed
biogeochemistry. A biogeochemical cycle can be expressed as a set of stores (pools)
and transfers, and can be studied using the concepts of "stoichiometry", "mass
balance", and "residence time".
Ecosystem function is controlled mainly by two processes, "top-down" and "bottomup" controls.
A biome is a major vegetation type extending over a large area. Biome distributions
are determined largely by temperature and precipitation patterns on the Earth's
surface.