EARTH AND LIFE SCIENCE

Ecology

Ecosystems, and their ecology, are essential matters of study as they

are pertinent to understanding life on earth. Ecology is a broad and
diverse field of study that concerns itself with the relationship between
organisms and their environment, as well as the relationships between
one environment and another. In this module, we will discuss the
principles of ecology in order to gain an understanding of how systems
work to keep and maintain the balance in the world today.

History of the Ecosystem Concept

One of the basic distinctions between autecology and synecology is
that autecology is considered the ecology of the individual organism
and populations. Autecology is mostly concerned with the biological
organisms themselves. Synecology, on the other hand, concerns itself
with the ecology of relationships among organisms and populations.
This is mostly concerned with the communication of material,
information, and energy of the entire system of components. In order
to study an ecosystem, one must have knowledge of the individual
parts. Thus, it is dependent on the fieldwork and experiments grounded
in autecology, but the focus is much more on how these parts interact
and relate to, and influence one another including the physical
environmental resources on which life depends. Therefore, ecosystem
ecology is the implementation of synecology.

The environment’s systems concepts have long played a rols in the

development of ecology as a discipline, but these came to a head in the
early 20th century. During this period, two dominant and competing
ecological paradigms were the organismisc and the individualistic
views. The organismic approach held that ecosystems and
communities were discernible objects that had an inherent and
organized complexity resulting in a cybernetic and self-governing
system, which is akin to the ways through which an organism regulates
itself.

On the other hand, the individualistic approach held that communities

had observer dependent boundaries and internal development was
stochastic and individual. In this paradigm, the internal relations were
synergistic, but not cybernetic since the individual parts functioned

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independently. The organismic paradigm grew out of the

understanding of whole systems, such as lakes and oceans, and also
from the discussions involving how communities changed over time
during the recession. These ideas were influenced by philosophers
during the time, such as Jan Smuts. There were both holists and
reductionists in this regard. The dialogue between them affected the
main currents of ecological thought during this period. It was, in part,
resolved by the introduction of the concept of ‘ecosystem,’ which is
aboth systemic and physical in nature.

The term ecosystem arose from this dialogue. It was first used by
Arthur Tansely and 1935, in a paper he published in the journal
Ecology. Tansley himself brought a systems perspective. The
underpinnings of the ecosystem have now become established.
However, the introduction of the term was theoretical, lacking
guidance as to how it might be useful as a field of study. A clear
application of the ecosystem concept was Lindeman’s study of Cedar
Bog Lake in Wisconsin. In addition to constructing the food cycle of
the aquatic system, Lindeman developed a metric, which is now called
the Lindeman efficiency. This metric was used to assess the efficiency
of energy movement from one trophic level to the next based on the
ecological feeding relations.

Defining an Ecosystem

As a unit of study, an ecosystem must be a bounded system. It can

range in scale from a puddle, to a lake, to a watershed, to a biome. The
scale of the ecosystem is defined more by the functioning of the
system rather than a checklist of constituent parts. The scale of the
analysis should also be determined by the problem being addressed.
Although individuals perish over time and even populations cannot
live indefinitely, every ecosystem contains the ecological community
necessary to sustain life. This includes the producers, the consumers,
the decomposers, and the physical environment for oikos. It is this
feature of the ecosystem that makes them the basic unit for sustaining
life over the long-term. The two main features of the ecosystem, which
are energy flow and nutrient biogeochemical cycling, comprise the
major areas of ecosystem ecology research.

Energy Flow in Ecosystems

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The recognition that an ecosystem is an open system is the starting

point of its thermodynamic assessment. The ecosystem receives
energy and matter input from outside its border and transfers output
back to this environment. Thus, every ecosystem must have a
boundary and must be embedded in an environment that provides low
entrpy energy input and can receive high entroy energy output. In
addition to the external sources of energy, there is another internal
environment with which organisms interact.

Patten proposed that there are two environments. One is external and
mostly unknowable (other than the input-output interactions) and the
second is internal and measurable (i.e. external to the specific
organismal component within system boundary).

Energy flow in the ecosystem begins with the capture of solar

radiation by photosynthetic processes in primary producers. Note,
however, that there are also chemoautotrophs that are able to capture
energy even without light. However, these chemoautotrophs contribute
negligible energy flux to the overall energy balance of the global
ecology.

Energy + 6CO2 + 6H2OC6H12O6 + 6O2

The accumulated organic matter, which starts out as sugars, then

combine with other elements to produce more complex molecules.
This represents the gross primary production in the system, some of
which is used for the growth and respiration of the primary producer.
The remaining energy, or net primary production, is available for the
rest of the ecosystem consumers, including decomposers. Secondary
production refers to the energetic availability of heterotrophic
organisms, which accounts for the energy uptake by heterotrophs, and
the energy used for their maintenance. Primary producers support the
overall ecosystem production. Additionally, ecosystem respiration
includes the metabolic activity of all the ecosystem biota. In this
manner, plants provide the essential base for all food webs.

The movement of energy goes through a network of dependencies,

which is known as the food web. In a simplified food chain, the
trophic concept is used to assess the distance away from the original
energy importation. However, in reality, the multiple feeding pathways
found in ecological food webs make discrete trophic levels a
convenient, yet inaccurate, simplification. Elton found that the number
of organisms decreases as one moves up from the food chain, from

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primary producers to herbivores, carnivores, and the top carnivores.

Thus, Elton proposed a pyramid of numbers. One can control the
variation of an individual’s body size by controlling for the biomass at
each trophic level, resulting in a pyramid of biomass.

The trophic pyramid is satisfying view of interactions since, according

to the second law of energy, energy must be lost at each
transformational step. In addition to this, energy is used at each trophic
level for the maintenance at that level. Under this paradigm, the
trophic levels cap out at five or six level. Fractional trophic levels have
been employed to account for organisms that feed at each different
trophic levels. However, these do not account for the role of detritus
and decomposition, which extends the feeding pathways to higher
numbers.

Energy sources flowing through the ecosystem are necessary to

maintain all development and growth activities. Organisms follow a
linear life history pattern and while the timelines vary from species to
species, early stage energy availability is generally used for growth,
what later energy surplus is used for reproduction or maintenance. A
similar pattern is observed in ecosystem growth and development. The
biomass and physical structure of the ecosystem is buit by net primary
production. The additional photosynthetic material allows for the input
of energy until saturation is reached at about 80% of the available solar
radiation. At this point, the overall growth of the ecosystem begins to
level off because although gross primary production is high, the
overall system supports more and more nonphotosynthetic organisms.
When the average gross production is utilized to support and maintain
the existing structure, net production is zero and the system has
reached a steady state regarding the growth of biomass. However, the
ecosystem continues to grow in terms of information capacity and
network organization. In addition to being in a dynamic steady state, it
does not persist indefinitely because disturbances occur which sets the
system to a previous successive state. In this manner, the disturbances
allow the ecosystem to develop along a different path.

Biogeochemical Cycles
Understanding how chemical elements are necessary for life is another
major focus of ecosystem ecology. The biosphere actively interacts
with the three abiotic spheres (hydrosphere, atmosphere, and
lithosphere) to provide the available concentration of each chemical

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element for life. This interaction has a significant impact on the

relative distribution of these elements. The products of photosynthesis,
which are simple sugars, are the bases for organic matter. Thus,
oxygen, hydrogen, and carbon dominate the composition of life.
Oxygen is available in the lithosphere and hydrogen in the
hydrosphere. However, carbon is quite scarce in the environment. A
hallmark of life is the disproportionate amount of carbon in the
biomass. There are about 20 elements used regularly in living
organisms, of which 9 are called macronutrients and are major
components of organic matter (hydrogen, oxygen, carbon, nitrogen,
calcium, potassium, silicon, magnesium, and phosphorous). Some of
these elements are easily available from the abiotic environment, in
which case conserving them through cycling is not important.
However, those that are scarce, such as phosphorous and nitrogen,
must be used many times before they are released from the system.
These biogeochemical cycles provide the foundation to understand
how human modification leads to eutriphica (N and P cycles) and
global climate change (C cycles). Therefore, much effort has been
made to study and understand these cycles, especially that of carbon,
nitrogen, and phosphorous.

Fundamental Laws in Ecology

There are, tentatively, 8 laws of ecology. However, it is pertinent to
split them into 10 because of the intricacies of each. Here, the 10 laws
of ecology will be presented.

1. All ecosystems are open systems embedded in an environment

from which they receive energy (matter) input and discharge
energy (matter) output.

-This first law is a prerequisite to ecological processes. If

ecosystems could be isolated, then they would be at
thermodynamic equilibrium without life and without gradients.
The first law is rooted in Prigogine’s law of thermodynamics.
The openness explains why systems can be maintained far
from thermodynamic equilibrium without violating the second
law of thermodynamics.

2. Ecosystems have many levels of organization and operate

hierarchically.

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-The law is based on differences in scale of interactions. The

distance between components becomes essential because it
takes time for events and signals to propagate. Ecological
complexity makes it necessary to distinguish between different
leves and with different local interactions.

3. Thermodynamically, carbon-based life has a viability domain

determined to be between 250 and 250 K.

- It is within the temperature range of 250 to 350 K that there is

a good balance between the opposing ordering and disordering
processes: decomposition of organic matter and building of
biochemically important compounds. The process rates are too slow
at lower temperatures. At higher temperatures, the enzymes that
catalyze the processes are destroyed. At increasing temperatures, as
well, the order creating processes increase, but the cost of
maintaining the structure in the face of disordering processes also
increases.

4. Mass, including biomass, and energy are conserved.

- This principle is used principally in ecological modeling.

5. The carbon based life on earth has a characteristic basic

biochemistry which all organisms share.

-It implies that many similar biochemical compounds can be found

in all living things. All living things have largely the same
composition, which can be represented using around 25 elements.
This principle also allows one to identify stoichiometric chemistry.

6. No ecological identity exists in isolation but is connected to

others.

-The theoretical minimum unit for any ecosystem is two

populations. One population fixes energy and the other decomposes
and cycles waste. In reality, however, ecosystems are composed of
complex interactions at all levels. These interactions provide
environmental niche in which each component acts. The network
also has a synergistic effect on its components. The ecosystem is
more than the sum of its parts.

7. All ecosystem processes are irreversible (this is the most useful

way to express the second law of thermodynamics).

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-Living organisms need energy to maintain, grow, and develop. This

energy is as heat to the environment, and cannot be recovered againt
as usable energy. Thus, evolution can also be understood in the
context of this principle that is rooted in the second law of
thermodynamics. Evolution is a stepwise development to based on
previously achieved solutions to survive in a dynamic and chaning
world. Due to the genetic and structural encapsulations of these
solutions, evolution has produced more and more complex
solutions.

8. Biological processes used captured energy (input) to move further

from thermodynamic equilibrium and maintain a state of low
entropy and high energy relative to its surrounding and to
thermodynamic equilibrium.

-This principle is another way of showing that ecosystems grow and

expand. It has been shown that eco exergy of an ecosystem
corresponds to the amount of energy that is needed to break down
the system.

9. After the initial intake of energy across a boundary, ecosystem

growth and development is possible by 1) an increase of physical
structure (biomass); 2) an increase of the network (more cycling); or
3) an increase of information embodies in the system.

-All three of these growth and development forms imply that the
system is moving away from thermodynamic equilibrium and all
three are associated with an increase of 1) the eco exergy stored in
the ecosystem, 2) the energy flow in the ecosystem (power), and 3)
the ascendency. When cycling increases, the space-time
differentiation, the energy use efficiency, and the eco exergy storage
capacity all increase. When information increases, the animal gets
bigger, the feedback control becomes more effective, which implies
that specific respiration decreases.

10. An ecosystem receiving solar radiation will attempt to maximize

eco exergt storage or maximize power such that if more than one
possibility is offered, then in the long run the one which moves
energy furthest from thermodynamic equilibrium will be selected.

-The eco exergy storage and energy flow increase during all three
growth and development forms. When an ecosystem evolves, it can
apply all three forms in a continuous Darwinian selection process.

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The nested space-time differentiation in organisms optimizes the

thermodynamic efficiency as expressed in this law, because it
allows the organism to simultaneously exploit equilibrium and non-
equilibrium energy transfer with minimum dissipation.

Population Coherence
In evolutionary biology, cost and benefit are measured in terms of
fitness. While mutation and natural selection represent the main forces
of evolutionary dynamics, cooperation is a fundamental principle that
is required for every level of biological organization. For instance,
individual cells rely on cooperation among their components.
Multicellular organisms exist because their cells exhibit cooperation.
Social insects, such as bees, are masters of cooperation. Whenever
evolution constructs something new, such as human language or
multicellularity, cooperation is needed. Thus, evolutionary
construction is based on cooperation. There are five rules for
cooperation.

1. Kin Selection

-Kin selection occurs when cooperation favors those from the

same family lineage. Kin selection is also more likely to work
in small groups, rather than large groups, unless highly inbred.

2. Direct Reciprocity

- All strategies of direct reciprocity can lead to evolution of

cooperation is the fundamental inequality is fulfilled.

3. Indirect reciprocity

-Whereas indirect reciprocity indicates that species will do for

others as others will do for them, indirect reciprocity suggests that
what one species does for another, then the other will do for others as
well.

4. Graph selection

-The traditional evolutionary game dynamics suggests that

populations are well-mixed. This means that interactions between any
two individuals in a population is equally likely to happen. More
realistically, however, the interactions between individuals are

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governed by spatial effects or social networks. Games, that are

presented on graphs, grew out of the tradition of evolutionary game
theory and spatial models in population genetics.

5. Group Selection

-Group selection is a powerful mechanism that promotes

cooperation. It only occurs, however, when all its basic requirements
are fulfilled in a particular situation.

Predator-Prey Interactions
A widespread population process, predation has evolved many times.
It can affect the distribution, abundance, and dynamics of species in
ecoystems. Predator-prey interactions have an inherent tendency to
fluctuate and show oscillary behavior. If predators are initially rare,
then the size of the prey population can increase. As prey population
increases, the predator population also begins to increase, which in
turn leads to a decrease in prey populations. As prey becomes scarce,
they the numbers of predators also decreases, and the cycle stars again.

Behavior and Patch Dynamics

Predators do not automatically respond to changes in prey density, nor

do they immediately convert prey to new predators. It takes time to
find, subdue, and consume prey. These behaviors require the
investment of time and energetic resources, and the foraging activities
of predators for prey have important consequences for the population
dynamics at a variety of different temporal and spatial scales.

For prey that are distributed in patches, patterns of distribution of

parasitism by insect parasitoids have been observed. The proportion of
hosts that have been parasitized can be a positive function, a negative
function, or independent of host density.

Intraspecific Competition
Individuals of the same species have very similar requirements for
their survival, growth, and reproduction. However, their combined
demand for resources may exceed the available supply. The
individuals then compete for the resource. Thus, some individuals
become deprived of resources.

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In many cases, competing individuals do not interact with each other

directly. Instead, individuals respond to the level of resource, which
has been depleted by the presence of other individuals. For instance,
grasshopers compete with one another for a plant. In such cases,
competition may be described as exploitation, in that each individual is
affected by the amont of resource that remains after that resource has
been exploited by others. Exploitation can occur if the resource that is
needed is of limited supply.

In many other cases, competition takes the form of interference. Here,

individuals interact directly with each other. Thus, one individual can
actually prevent another individual from accessing a resource. For
instance, this can be seen among animals who defend their own
territories, and among the sessile animals and plants that live on rocky
shores. Thus, interference competition may occur for a resource of real
value, such as space. In this case, interference is accompanied by a
degree of exploitation, or for a surrogate resource (territory, ownership
of a harem), which is only valuable because of the access it provides to
real resource (food, females). With exploitation, the intensity of
competition is closely linked to the level of resource present and the
level required. On the other hand, with interference, intensity might be
high even when the level of the real resource is not limiting.

Another case is of one-sided competition. Whether they compete

through exploitation or interference, individuals within a species have
many fundamental features in common, using the same resources and
reacting in a similar way to similar conditions. Nonetheless,
intraspecific competition may be one-sided. For instance, a plant may
shade another plant, causing it to die. Hence, the ultimate effect of
competition is far from being the same for every individual. Weak
competitors may make only small contributions to the next generation,
or no contribution at all.

Species Interactions
Individuals from different species can compete with each other. There
are two general points when considering interspecies competition.
First, careful, separate, ecological attention must be paid to both the
ecological and the evolutionary effects of interspecific competition.
The ecological effects are, broadly, that species may be eliminated
from a habitat by competition from individuals of other species. Or, if
competing species coexist, that at least one individual from the species

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undergoes deprivation. The evolutionary effects appear to be that

species differ more from one another that they would otherwise do.
Thus, they compete less.

The second point is that there are important and profound difficulties
in invoking competition as an explanation for observed patterns, and
especially invoking it as an evolutionary explanation.

There are two types of interspecific competition: symmetric and

assymetric. Asymmetric competition occurs when the effects of the
competition are not the same for both species. On a broade front, it
shows that highly asymmetric cases of interspecific competition
gradually outnumber symmetric cases. The fundamental point,
however, is that there is a continuum linking the perfectly competitive
cases to strongly asymmetric ones. Asymmtric competition results
from the differential ability of species to occupy higher positions in the
competitive hierarchy. In plants, for instance, this may result in height
differences, when one species is able to gain access to more light
compared to another. Asymmetric competition is also more likely
when the body sizes of species differ greatly.

Similary, the competition for one resource may influence the

competition for another resource. Examples are found among rooted
plants. If one species invades the canopy of another and deprives it of
light, the suppressed species will suffer directly from the decreased
light energy available. Therefore, they will also be less able to exploit
the water and nutrients in the soil. This, in turn, will reduce the rate at
which leaves and shoots grow. Therefore, when plant species compete,
repercussions flow backwards and forwards between roots and shoots.

The niche of a species without competition from other species is its

fundamental niche. On the other hand, the niche of a species in the
presence of competitors is its realized niche. The nature of the realized
niche is determined by the characteristics of the competing species that
are present. This can be summarized in the Competitive Exclusion
Principle, which states: 1) if two competing species coexist in a stable
environment, then they do so as a result of niche differentiation (i.e.
differentiation of their realized niches). If, however, such
differentiation does not occr, then one competing species will
eliminate or exclude the other. Thus, exclusion occurs when the
realized niche for a superior competitor completely fills those parts of
the inferior competitio’s fundamental niche that are provided by the
habitat.

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There is also the phenomenon of mutual antagonism. Mutual

antagonism can be seen in beetles, in which adult beetles may
sometimes cannibalize their own species. They also ate beetles from
other species. The important note here is that the beetles ate
individuals from other species more than they ate their own species.
Thus, a crucial mechanism for the interaction of these competing
species is reciprocal predation, and each species was more affected by
interspecific competition rather than intraspecific competition. Mutual
antagonism is strongest when the number of one species is greater than
the other.

The Nature of Predation

Predation is the consumption of one organism (prey) by another
organism (predator). In this relationship, the prey is alive when the
predator first attacks it. Thus, detrivory, the consumption of dead
matter, is excluded from the predator-prey relationship. Nevertheless,
the prey-predator relationship encompasses a wide variety of
relationships and interactions, with a wide variety of predators.

There are two main ways through which predators can be classified.
Neither one of these ways is perfect, but they are useful. The most
obvious classification is taxonomic. That is, carnivores consume
animals, herbivores consume plants and omnivores consume both.

An alternative way of classification is the functional classification.

Here, there are four types of predators. There are true predators,
parasitoids, grazers, and parasites (which are divisible into
macroparasites and microparasites).

True predators kill their prey immediately after attacking them. During
the lifetime of the true predator, it consumes several of many different
prey organisms, often consuming the prey in its entirety. The most
obvious true predators are lions, tigers, etc. However, rodents, ants,
and even plankton-consuming whales are also true predators.

Grazers, on the other hand, attack a large number of prey during their
lifetime. However, they do not consume their prey in its entirety, but
only parts of it. Their effect on the prey organism is rarely lethal in the
short-term, although it is typically harmful. The most obvious grazers
are sheep, cattle, and cows. However, flies that bite vertebrate prey,

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and leeches that suck their blood, are also grazers. Thus, grazers are
not limited to herbivores.

Parasites, like grazers, consume parts of their prey. Like grazers, they
are harmful, but they are not lethal in the short-term. Unlike grazers,
however, parasites only attack one or very few individuals during their
lifetime. Thus, there is an intimate relationship between parasites and
their hosts that is not seen in true predators and grazers. Measles virus,
tapeworms, liver flukes, and tuberculosis bacterium are all examples
of parasites. There are also many fungi, plants, and microorganisms
that act as parasites on plants, called plant pathogens. In addition,
many herbivores can be thought of as parasites. For instance, aphids
extract sap from plants with which they enter into intimate contact.
Even caterpillars can be thought of as parasites, as they often rely on
just one plant.

Parasitoids are a group of insects that belong mainly to the order

Hymenoptera, but it also includes individuals from Diptera. These
individuals are free-living as adults. However, they lay their eggs in,
on, or near other insects. The larval parasitoid develops either inside
its host or on its host. At first, it does little harm, but it eventually
consumes the host and therefore kills it. Thus, an adult parasitoid
emerges from a host. Often, just one parasitoid develops from one
host, but there are instances wherein several parasitoids develop from
one host. The rate of predation of parasitoid is dependent on the rate at
which the adult females lay eggs. Each egg, therefore, is an attack on
the host, even though it is the larva that emerges from the egg that
does the killing. Parasitoids seem to be of very little importance.
However, they account for 10% of the world’s species. Parasitoids
maya also be attacked by other parasitoids.

Human-Caused Changes on the Earth’s Ecosystem

Humans have effects on the earth’s ecosystems. Humans transform
land surfaces, add or remove species, and alter the biogeochemical
cycles. There are human activities that have direct impacts on the
earth’s ecosystem, such as land use changes and management. Other
activities, however, are indirect, such as atmospheric changes and
changes in hydrology. At least some of these anthropogenic activities
affect all the ecosystems on earth.

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The most substantial and direct effect of human activities that alter
ecosystems is the conversion of land for production of food, fiber, and
other goods used by humans. About 50% of the ice-free land on earth
has been altered by human activities. Agricultural fields and urban
areas cover 10-15% of land areas, whereas pastures cover 6-8% of the
land. Even more land is used for grazing and forestry. All portions of
the earth, except the most extreme environments, can experience
human impact.

Marine and freshwater ecosystems are also altered by human activities.

For instance, half of all the world’s accessible run-offs are used by
humans. In addition to this, humans use about 8% of the primary
production of the oceas. Commercial fishing reduces the size and
abundance of target species. It also alters the population characteristics
of species that are incidentally caught in the fishery. About 22% of all
marine fisheries are over-explooted or already depleted. Many coastal
areas are also being exploited. Nutrient enrichment of these areas has
led to the increased production of algae and created anaerobic
conditions in which fish cannot survive. This nutrient enrichment is
due largely to the transport of agricultural fertilizers and from human
sewage and livestock sewage.

Land use change causes a loss of habitat. It is the primary driving force
for the extinction of species and the loss of biological diversity. There
is also a time lag between ecosystem changes and species loss, which
makes it likely that species will continue to be driven to extinction
even when land use change have stabilized. Homogeneity of the
earth’s biota is also being caused by the transport of species around the
world. The frequency of these invasions is increasing, in large part due
to the globalization of the world’s economy. International commerce
breaks down biogeographic barriers, through both purposeful trade of
live organisms and inadvertent introductions. The former selects
species which are more likely to grow and reproduce in the new
environment. Many of these biological invasions are irreversible
because it is too expensive and difficult to remove species that have
invaded. Some of these species incur large economic losses or cause
damages to human health. Others alter to balance of ecosystems,
leading to further losses of species.

Human activities have also altered biogeochemical cycles in many

ways. Usage of fossil fuels and the intensification and expansion or
agriculture have altered the carbon cycles, nitrogen cycles, and sulfur
cycles on a global scale. These changes in biogeochemical cycles alter

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ecosystems and they also influence unmanaged ecosystems through

changes in lateral fluxes of nutrients and other materials through
surface waters and the atmosphere. Land use changes, which include
the intensive use of fertilizers and deforestations, have increased the
concentration of atmospheric gases that influence the climate. Land
transformations cause runoff and erosion of sediments that lead to
significant changes in lakes, rivers, and oceans.

Activities/Exercises

Assignment 1: Do you agree that a puddle can be considered an ecosystem? Why or why
not? Explain your answer in 300 words, with 2 references formatted in APA style.

Assignment 2: Choose an ecosystem from a foreign country and research this on the
internet. In 600 words, what is the main defining feature of that ecosystem that differentiates it
from our own ecosystems? How do energy transfers take place? Use 3 references formatted in
APA style.

Glossary
Autecology: the study of organisms or a particular species

Exergy: the available energy that can be used

Synecology: the study of whole communities of plants or animals

Mutual antagonism: occurs when interspecific competition has effects that

outweigh those of intraspecific competition

Thermodynamic equilibrium: the state at which an object with higher

temperatures transfers heat to an object with lower temperature, thereby
reaching a temperature equilibrium.

References
Gotelli, N. J. (1995). A primer of
ecology. Sinauer Associates
Incorporated.

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