Ololade Kolade

Dr. Kabak
AP Biology
Unit 8: Ecology (p. 1078-1231)
http://www.course-notes.org/Biology/Slides/Chapter_1_Exploring_LifeConcept 50.1:
Ecology is the study of interactions between organisms and the environment
Ecology and Evolutionary Biology: Events that occur in the framework of
ecological time translate into long-term effects in evolutionary time. An example of which,
the predator-prey relationship, has a direct effect in the size of the prey population
(ecological effect), which translates into the altering of the entire population’s gene pool
(an evolutionary effect).
Organisms and the Environment: An organism’s environment includes two things:
abiotic (nonliving; ex. light, temperature, water etc.) components and biotic (living)
components. The biota is all of the organisms that are present in that individual’s
environment. Organisms may compete with one another for food, prey on each other,
parasitize each other, become a food source for one another, or change the relative
physical and chemical environment.
Ecological Relationships: Distribution and Abundance
Geographic range, or Distribution, refers to the areas that a
particular type or species of organism may live. Abundance refers
to the amount of the organism in varied areas. Controls on
distribution and abundance may be both abiotic and biotic, or a
combination of the two.
Subfields of Ecology:
Organismal Ecology: how an organism’s structure, physiology, and (for
animals) behavior meet the challenges posed by the environment.
Population Ecology: Concentrates on factors that affect how many
individuals of a particular species live in a single geographic area (i.e. a
population).
Community Ecology: Focuses on how interactions such as predation,
competition and disease, as well as abiotic factors such as disturbance
(force that changes ecological community and usually removes organisms
from it) affect community structure and organization. A community
consists of all the organisms of all the species that inhabit a particular
area.
Ecosystem Ecology: Emphasizes energy flow and chemical cycle between
the various biotic and abiotic components in the ecosystem, all the abiotic
factors and the entire community of species that exist in a certain area.
Landscape Ecology: Deals with various ecosystems and how they are
arranged in a geographic region. All landscapes and seascapes consist of
different “patches”, a characteristic labeled patchiness. L.E. focuses on
factors controlling the exchanges of energy, materials, and organisms
among the ecosystems composing a landscape or seascape.
The biosphere is the global ecosystem, consisting of all the planet’s ecosystems.
The biosphere includes all of Earth that is inhabited by life, from the atmosphere at very
high altitudes to under-ground and sea caves. Research at the biosphere level studies how
global changes will affect all life.
Ecology and Environmental Issues:
 Ecology vs. Environmentalism: Environmentalism is the advocacy for the
protection or preservation of the natural environment. To address and cure
environmental problems, understanding of organisms and their
relationships with the environment is required, and thus, is acquired
through ecology. Rachel Carson’s book, Silent Spring, displays
environmentalism. Ecologists can aid legislators when creating laws that
will effect the environment.
The Precautionary Principle: The interconnectedness of the biosphere
leads to a fairly simple change meant to affect one population drastically
affecting countless others. The precautionary principle acts a guiding
principle when making decisions about the environment, stating that
because of this, potential consequences should be carefully considered.

Concept 50.2: Interactions between organisms and the environment limit the distribution of
species
Dispersal and Distribution: Dispersal is the movement of individuals away from
centers of high population density or from their area of origin.
Natural Range Expansions: The role of dispersal is demonstrated when
organisms expand their range by moving into areas where they did not
exist previously. Natural range expansions happen by natural means and
clearly show the influence of dispersal on distribution: when an organism
disperses, its global distribution is increased. Thus, expansion of
distribution relies on organism dispersal.
Species Transplants: Observing the accidental or intention transplant of a
species to an area it was previously absent is a way of determining if
dispersal is a limiting factor of distribution. For a transplant to be
successful, some organisms must both survive and reproduce in the new
environment. If a transplant is successful, it can be concluded that the
potential range (where the species could live) of the species is larger
than its actual range (where the species does live). Species transplants
often disrupt communities/ spread beyond the intended area of
transplantation.
Behavior and Habitat Selection: Some species do not occupy all of their potential
range even though they may be physically able to reach it. Distribution may be limited by
habitat selection behavior.
Biotic Factors: Biotic factors deal with the prospect of other species limiting the
distribution of a species through predation, parasitism, disease, herbivory, the
presence/absence of food, or competition. The role of biotic factors in limited distribution is
tested in removal and addition experiments, in which the alleged limiting factor is
removed and the growth of the species is recorded.
Abiotic Factors: The prospect of abiotic factors: temperature, sunlight, water, etc.
—limiting the widespread distribution of species. The environment is characterized by
spatial heterogeneity and temporal heterogeneity, the variations in both space and
time. Different region may experience fluctuation in abiotic condition, which may blur of
accentuate distinctions.
Temperature: May affect biological processes by freezing the water in cells/
affecting the heat exchange that regulates organisms’ internal environment. \
 Water: Organisms are distributed according to their water requirements.
Land-based organisms’ distribution reflects their ability to get and conserve
water. Some organisms have adapted for water acquisition and conservation
in dry environments.
Sunlight: Sunlight drives all ecosystems, is important to the
development/behavior of organisms sensitive to the relative times of night
and day. Sunlight may limit plant growth, and in marine environments, the
amount of sunlight limits the distribution of photosynthetic creatures.
Wind: Amplifies the effects of temperature by increasing heat loss,
contributes to water lo by increasing rate of evaporative cooling in
animals/transpiration in plants. Wind can “flag” tree limbs by inhibiting growth
on the windward side.
Rocks and Soil: Structure, pH, and mineral composition of rocks and soil
limit the distribution of plants and, in consequence, the animals that feed on
them. This is part of the “patchiness” of terrestrial ecosystems. Residue at
the bottom of bodies of water can affect water chemistry, influencing resident
organisms.
Climate: Temperature, water, sunlight, and wind are the four major components of
climate, the prevailing weather conditions in a particular area. Climate factors majorly
affect organism distribution. Macroclimate: patterns on the global, regional, and local
level. Microclimate: very fine scale patterns, in a very, very small controlled area.
Global Climate Patterns: Are largely determined by solar energy and
planetary movement in space. The sun’s warming effect on
land/water/atmosphere creates temperature variations, cycles of air
movement, and water evaporation that  latitudinal changes in climate.
Regional/Local/Seasonal Effects on Climate: Closeness to
water/topographic features like mountains/small landscape features
contribute to local climate variation. Regional and local variations contribute
to patchiness. Seasonal variation is another influence on climate.
 Bodies of Water: Ocean currents heat or cool overlying air masses,
which can then move across land at coastal regions. Coasts are usually
moister than inland regions at the same latitude. Oceans and large
lakes often moderate the climate of nearby terrestrial environments;
when this fails to happen it is called a Mediterranean climate, which is
a hot, rainless inland environment.
 Mountains: Mountains affect the amount of sunlight that reaches an
area, as well as temperature and rainfall. As elevation increases over
the mountains, temperature decreases, also in combination with the
decline of temperature with increasing latitude. When warm, moist air
meets a mountain, the air rises and cools, creating deep snow packs.
On the far side of the mountain, cooler, dry air falls, absorbing moisture
and producing a rain shadow with little precipitation.
 Seasonality: The angle at which sunlight hits the earth over the course
of the year affects local environments. Seasonal changes in wind
patterns create variations in ocean currents, sometimes sweeping
nutrient-rich water from the ocean floor, nourishing organisms at the
surface. Summer and winter cause thermal striation in lakes, in which
 they are layered according to temperature. As a result of changing
temperatures, the waters are mixed, bringing nutrients from the bottom to
the surface and oxygen from the surface to the both every spring and
autumn. This is the turnover.
Microclimate: Various things affect microclimates by casting shade, affecting
evaporation from soil, and changing wind patterns. All environments are
characterized by a mosaic of small-scale differences in abiotic factors that
affect organism distribution.
Long-Term Climate Change: Prediction of long term climate change is made
with the occurrences of past ice ages. Seed dispersal aids in tree migration.
Researchers can make predictions on how climatic warming with affect
distribution if they can determine the climatic limits of current distribution
ranges. Slow seed dispersal may contribute to extinction of species during
climatic changes, as they will not be able to move to their prediction ranges
at the needed speed.

Concept 50.3: Abiotic and biotic factors influence the structure and dynamics of aquatic
biomes
Aquatic Biomes: Biomes are any of the world’s major ecosystems, classified by
predominant vegetation and characterized by adaptations of organisms that live there.
Aquatic biomes (oceans in particular) account for 75% of the Earth and their evaporation
accounts for most of the planet’s rainfall; ocean temperatures have a major impact on
global climate and wind patterns. A large amount of marine algae and photosynthetic
bacteria provide the world’s oxygen and take in CO2.
Aquatic Biome Zonation: Aquatic Biomes are physically & chemically stratified.
The photic zone is the top of the lake, where there is enough light for photosynthesis. The
aphotic zone has little light. The bottom of all aquatic biomes is called the benthic zone,
which is made of sand and inorganic sediments (ooze), and inhabited by organisms called
benthos. Benthos eat dead organic matter called detritus that rains down from the photic
zone. The deepest part of the ocean is called the abyssal zone. In the ocean and most
lakes, a thermocline (a narrow stratum of rapid temperature change) separates the warm
upper layer from the cold deeper waters.
Community Distribution in Aquatic Biomes: In aquatic biomes, communities are
distributed according to water depth, intensity of light penetration, distance from shore, and
open water vs. bottom. Marine biomes often lack a vast amount of life in the aphotic zone,
and harbor a variety of fish and plankton in the photic zone.
Lakes—standing bodies of water *Salinity/Oxygen-concentration/nutrient
surrounded by land. Light decreases with content varies in lakes
depth, creating stratification. Temperate *Oligotrophic lakes are nutrient-poor
lakes have seasonal thermocline; tropical and oxygen-rich; decomposable organic
lowland lakes have one year-round. material in sediments is low, less SA to
Littoral zone, the shallow, well-lighted depth, may become eutrophic w/ runoff
waters close to shore, houses *Eutrophic lakes are nutrient-rich and
floating/rooted plants. The limnetic often depleted of oxygen if ice-covered in
zone (deep water), has phytoplankton, winter and up in summer, decomp.
cyanobacteria, zooplankton. Benthic organic material is high.
zone has invertebrates; all have fish is Pollution leads to nutrient
oxygen concentration supports. enrichmentalgae blooms, oxygen

Wetlands—an area covered with water
for a long enough period to support
aquatic plants. Water and soils are
periodically low in dissolved oxygen,
have high capacity to filter dissolved
nutrients and chemical pollutants. Water-
saturated soil favors growth of many
plants; woody plants=swamps,
bogs=mosses. Home to
invertebratessupport birds. Herbivores
consume algae/detritus/plants.
Carnivores include insects, mammals,
reptiles, & birds.
Streams and Rivers—Streams are
characterized by cold temperature, clear
water, and turbulent/swift current. Rivers
are warmer and muddy, because rivers
carry more sediment than their
headwaters. Streams/rivers are stratified
into vertical cones. Headwater streams
are often narrow with rocky bottoms.
Downstream rivers are usually wide and
winding, bottoms are silty from
sediments.
Estuaries—Transition areas between
rivers and sea. Complex flow patterns.
Grasses and algae; crustaceans, human-
consumed fish, worms. Invertebrates/fish
use as breading ground.
Intertidal Zones—A periodically
submerged or exposed area by the tides
on marine shores. Oxygen/nutrient levels
are high; renewed by tides. Rarely
support rooted photosynthetic
organisms. Small animals, worms,
crustaceans and fish live in intertidal
zones.
Oceanic Pelagic Biome—A vast realm
of open blue water, constantly mixed by
wind-driven ocean currents. Oxygen
depletion, fish kills.
*Basin Wetlands develop in shallow
basins, include filled in lakes &
ponds/upland depressions
*Riverine Wetlands develop along
shallow and periodically flooded banks of
rivers/streams.
*Fringe Wetlands occur on coasts of
big lakes/seas because of rising lake
levels or tidal actions.
Draining/Filling destroys large amounts of
wetlands.
*Salinity and nutrient content of
streams/rivers increases from
headwaters to mouth. Stream
headwaters are rich in oxygen; rivers
may be the same.
*Headwater streams are rich in
algae/rooted aquatic plants. Lots of fish.
*Salinity varies
*Some of most productive biomes
*Flow patterns created tidal channels,
islands, levees, and mudflats
*Pollution has disrupted
*Variations in temperature make it
difficult for organisms to survive
*Variations from upper intertidal zone to
lower limits the distribution of several
organisms
*Animals often exhibit adaptations that
allow them to attach to rocky bottom.
*Oil spills have disrupted; decline in
birds/turtles
*Thermally stratified all year
*Covers about 70% of Earth’s surface
*Most animals are zooplankton, many
levels high; nutrients low.
Coral Reefs—Are limited to areas with
higher water clarity High oxygen levels
and nutrients. Fairly temperature
sensitive.
Marine Benthic Zone—The ocean floor.
Oxygen is sufficient, except in areas of
organic enrichment. Soft sediments
cover, some rocky bases, underwater
volcanoes, and oceanic crust. Many
invertebrates, arthropods, and fish live in
the benthic zone.
free-swimming animals
*Over-fishing kills fish; pollution through
waste dumping and oil spills
*Require solid substrate for attachment
*Reef corals form symbiotic relationships
with unicellular algae, providing selves
with nutrients
*Collection of coral skeletons/over
fishing/aquarium trade have reduced
coral populations. Global warming and
pollution maybe be contributing to coral
death.
*the Nertic zone—sea floor below
surface waters of the coast
*Most of the Benthic zone gets no
sunlight
*Water temperatures decline with depth
*Deep-sea hydrothermal vents—dark,
hot, low oxygen areas; chemoautotrophic
prokaryotes obtain energy through a
reaction w/ hot water dissolving sulfate
*Dumping has created low- oxygen zones

Concept 50.4: Climate largely determines the distribution and structure of terrestrial
biomes
Climate and Terrestrial Biomes: Impacts of climate on distribution can be seen on
climographs, plots of temperature and precipitation of a certain area. Averages for
temperature & rainfall are correlated with biomes that exist in different regions.
Correlation vs. Causation: Climographs show that rainfall & temperature are
important to the distribution of biomes, but they do not confirm they govern
biome location. There are region where biomes overlap, which means other
factors play a role in determining where biome exist. Variations may be
accounted for in climatic change patterns, not merely averages.
General Features of Terrestrial Biomes: Terrestrial biomes are name for physical
& climate features, as well as predominant vegetation, also characterized by
microorganism, fungi, & animals. Biomes grade into each other; the area of intergradation,
called an ecotone, may vary in width. Disturbance in biomes is more common that
stability; leads to patchiness within biomes. Dominant plants depend on periodic
disturbance. Human activity has altered natural patterns of disturbance.
Vertical stratification in Terrestrial Biomes: Forests—upper canopy, low-tree
stratum, shrub understory, ground layer of herbaceous (without woody
stems) plants, forest floor (litter layer), root layer. Non-forests—herbaceous,
litter layer, root layer. Stratification provides many different habitants for
animals.
Tropical Forest—Distributed in *Little temperature variation, warm all
equatorial and subequatorial regions, year-round
with relatively constant rainfall in *Stratified, intense competition for
tropical rainforests. Tropical dry light
forests have seasonal precipitation & a *Great amount of animal diversity, all
6-7 month dry season. Display typical adapated
forest striation. Fewer strata in dry *Population growth is destroying
forests. tropical forests
Deserts—Occur in a band near 30 *Temperature is variable seasonally
degrees north/south latitudes, or in and daily, hot deserts may exceed 50C
the interior of continents. Low and cold below 30C
precipitation, highly variable. Human *Low, widely scattered vegetation,
intervention has made it possible for adapted for heat/desiccation
populations to survive in deserts; tolerance, water storage, reduced leaf
destroy natural biodiversity. SA.
*Many nocturnal animal species,
water conservation=common
adaptation.
Savanna—Occur in *Scattered trees are thorny w/ reduced
equatorial/subequatorial regions. leaf SA (adaptation).
Rainfall is seasonal, dry season *Dry season fires fire-adapted plants
averages 8-9 months. Typically warm *Large mammalian herbivores and
year round, more season variation predators are common, dominant
than tropical forests. Earliest humans herbivores are termites
are said to have lived on savannas.
Chaparral—Occurs in mid-latitudinal *Shrubs & small trees, high diversity of
coastal regions on several continents. grasses & herbs
Precipitation is seasonal, rainy winters;*High plant diversity, species confined
long dry summers. Fall, winter, spring to small are
are cool, summers are very hot. *Adaptations to drought in plants:
Humans have converted chaparral tough leaves which reduce water loss.
areas to settlements, agricultural Fire-adaptations: seeds that only
areas, urban areas, contribute to fires germinate after hot fires etc.
that sweep across chaparral. *Support deer/goats, small mammals,
amphibians, birds, reptiles, and insects
Temperate Grassland—South Africa, *Dominant plant are grasses/forbs
Hungary, Argentina/Uruguay, Russia, (broad-leaved herbaceous plants)
North America. Precipitation is *Main adaptations of plants are of
seasonal; dry winter, wet summers. droughts/fire
Periodic drought is common. Winters *Large grazers (horses/bison),
are generally cold, summers are burrowing mammals
generally hot. *Ideal for agriculture, landconverted
to farm land
Coniferous Forest—A broad band *Cone-bearing trees, conical shape
around northern North America and prevents snow from accumulating +
Euraisa to the edge of the artic tundra; breaking branches
the largest terrestrial biome on Earth. *Diversity lower than in temperate
Precipitation is moderate; periodic broadleaf forests
droughts are common. Cold, long *Migratory birds, moose, brown bears,
winters/Hot summers. Siberian tigers
*Insects kill tracts of trees

Temperate Broadleaf Forests—
Found mainly at mid-latitudes in
Northern Hemisphere, in NZ and
Australia. Precipitation averages from
70-200 cm, significant amounts all
seasons. Winter temps average at
freezing; summers are hot & humid.
Temperate broadleaf forest has been
settled on all forest; logging/land
clearing has destroyed forests.
Tundra—covers expansive areas of
the arctic. High winds/ cold
temperatures create similar plant
territories call alpine tundra on
mountain peaks. Precipitation varies
from 20-60 cm in arctic, may exceed
100 in alpine. Winters are long and
cold, summers are shot and cool.
Ololade Kolade
*Being logged at alarming rate
*Have distinct, highly diverse layers
(closed canopy, one or two strata of
understory of trees, shrub layer,
herbaceous stratum)
*Dominant plants are deciduous trees,
drop leaves before winter
*Mammals hibernate in winter; birds
migrate
*Insects, birds, mammals make use of
all layers
*herbaceous plants, many bryophytes
*permafrost, a permanently frozen
layer of soil, generally prevents water
infiltration
*Oxen are present, caribou and
reindeer migrate, predators are bears,
wolves, foxes. Birds nest in summer.
*Humans extract oil and minerals
Dr. Kabak
AP Biology
Chapter 51: Behavioral Ecology

Concept 51.1: Behavioral ecologists distinguish between proximate and ultimate causes of
behavior
What is Behavior?: Behavioral traits are considered parts of an animals
phenotype. Most of behavior is what the visible result of an animal’s muscular activity is in
a given situation. Some non-muscular activities, such as secreting a pheromone, are
considered behavior. Learning is considered to be a behavioral process. Behavior can be
thought of as everything an animal does and how it does it.
Proximate and Ultimate Questions: Proximate questions about behavior focus
on environmental stimuli and the genetic, physiological, and anatomical mechanism
underlying a behavioral act. They are also referred to as “how” questions. Ultimate
questions address the evolutionary success of a behavior. They are often “why”
questions, proposing that the behavior increases fitness in a way. Proximate mechanisms
produce behaviors that have evolved because they reflect fitness in a way.
Ethology: Ethology is the scientific study of how animals behave in their natural
environments. Niko Tinbergen suggested the four questions that must be answered to
fully understand any one behavior:
1. What is the mechanistic basis of the behavior, including chemical
anatomical and physiologic mechanisms? (proximate)
2. How does development of the animal, from zygote to mature individual,
3. influence the behavior. (proximate)
4. What is the evolutionary history of the behavior? (ultimate)
5. How does the behavior contribute to survival and reproduction (fitness)?
(ultimate)
Fixed Action Patterns: A fixed action pattern (FAP) is a sequence of
unlearn behavior that is essentially unchangeable and, once initiated, is
usually carried to completion. FAPs are triggered by an external sensory
stimulus known as a sign stimulus.
Imprinting: Imprinting is a type of behavior that includes learning & innate
components that is generally irreversible. It is different from other learning by
having a sensitive period, a limited phase in an animal’s development that
is the only time certain behaviors can be learned. In species that provide
parent care, the young imprint of the parent and learn basic behaviors. The
tendency to respond is innate to the birds; the outside world provides the
imprinting stimulus, something to which the response will be directed. The
respond to and identify with the first object they encounter that has certain
key characteristics. By utilizing imprinting, species can be conserved.

Concept 51.2: Many behaviors have a strong genetic component

Introduction: Behaviors are the result of genetic and environmental factors. An
approach to studying the influence of different factors on a particular behavior is to view
the behavior in terms of the norm of reaction. Sometimes nearly all members of a
population exhibit the same behavior, despite internal & external environmental differences
during development/throughout life. Behavior that is developmentally fixed in this way is
called innate behavior. Innate behaviors are under strong genetic influence.
Directed Movements: Many animal movements are under substantial genetic
influence; because of this, they are referred to as directed movements.
Kinesis: A kinesis is a simple change in activity or turning rate in response to
a stimulus.
Taxis: A taxis is an automatic oriented movement toward (positive taxis) or
away (negative taxis) from some stimulus.
Migration: Many features of migratory behaviors are genetic. Migratory
restlessness (absent flapping of wings, hopping restlessly) occurs in
populations that do not need to migrate to survive, and thus remain “captive”
during the season in which their migrating counterparts migrate. Studies
crossing captive birds and migratory birds have lead to the conclusion that
migratory restlessness is a polygenic trait.
Animal Signals and Communication: A signal (display) is a behavior that causes
a change in another animal’s behavior. Much of social interaction between animals
involves transmitting information through signals. The transmission of, reception of, and
response to signals composes animal communication, an essential element of
interactions between individuals. Many signals are very efficient in terms of energy
expenditures. Animals communicate using visual, auditory, chemical (olfactory), tactile,
and electrical signals; the type of signal used depends on an animal’s lifestyle &
environment.
Chemical Communication: Many animals that communicate through odors
emit pheromones, chemical substances that function in communication. In
several cases, the production of pheromones and animal responses to them
are controlled genetically. The context of a chemical signal (eg. with
honeybees) can be as important as the chemical itself. Pheromones can also
function in non-reproductive behaviors, such as when issuing panic
responses.
Auditory Communication: The songs of most birds species are at least partly
learned. In insects, the songs are generally under direct genetic control. A
variety of evidence shows that the song is genetically controlled and under
strong selective pressure. Some insect species can only be identified through
their courtship songs. Hybrids of lacewings with different mating songs had
properties of both parents in one mating song, leading to the conclusion that
songs are generally genetically controlled.
Genetic influences on Mating and Parental Behavior: Behavioral research
proves that many mammalian behaviors are under strong genetic control. Monogamy, non-
aggression, and parental behaviors in prairie voles are linked to arginine-vaspressin, a
neurotransmitter released during matting, binding to V1a receptors. The distribution of the
receptors causes the behavior. Thomas R. Insel injected the prairie vole V1a gene into
mice, which then developed brains similar to voles and exhibited similar mating/parental
behaviors.

Concept 51.3: Environments, interaction with an animal’s genetic makeup, influences the
development of behaviors
 Introduction: Environmental factors such as quality of diet, nature of social
interactions, and opportunities for learning can influence the development of behaviors in
every group of animals.
Dietary Influence on Mate Choice Behavior: Flies lay their eggs on the rotting
tissues of cactus plants, where the cacti can serve as food for the developing larvae. When
flies from two different areas, which lay young on two different breed of cacti, were bred on
the same artificial banana-based medium, females from Sonora (organ pipe cactus)
avoided males from Baja California (agria). When raised on a cactus medium, the larvae
chose mates from both reasons with approximate equal frequency. The diet caused
differences in the hydrocarbons in the exoskeletons of the flies, which caused an aversion
within Sonora females toward Baja males.
Social Environment and Aggressive Behavior: Researchers have done cross-
fostering studies in California mice (monogamous, aggressive, parental behavior) and
white-footed mice (non-aggressive, non-monogamous, non-parental) This has uncovered
an influence of social environment on the aggressive and parental behaviors of these
mice.

Learning: Learning is the modification of behavior, based on specific experiences.

Learned behaviors can range from very simple to very complex.
Habituation: Habituation is a loss of responsiveness to stimuli that convey
little or no information. It may ultimately increase fitness because it will allow
the animal to focus on stimuli that are directly relevant to its survival, and not
waste time or energy on those that are not.
Spatial Learning: Spatial learning is the modification of behavior based on
experience with the spatial structure of the environment. Tinbergen did a
study using pinecones and a wasp’s nest, in which he came to the
conclusion that the wasps learned to view the pinecones as a landmark, or
location indicator. The experiment also shows that for spatial learning to be a
reliable way to navigate through the environment, the landmarks must be
stable. The degree of environmental stability influences the spatial learning
strategies of animals.
Cognitive Maps: A cognitive map is an internal representation of the spatial
relationships between objects in an animal’s surroundings, and thus a more
 efficient mechanism in locating things. Birds have been found by researchers
that could find the midway point between landmarks, suggesting that they are
capable of using general geometric rules to locate seed caches.
Associative Learning: In associative learning, animals associate one
feature of their environment with another. Classical conditioning is a type
of associative learning, in which an arbitrary stimulus is associated with a
reward or punishment. Animals become conditioned to associate one
substance with another. Operant conditioning is another type of associative
learning in which an animal learns to associate one of its behaviors with a
reward or punishment. For example, predators learn to associate potential
pretty with painful experiences and modify their behaviors accordingly.
Cognition and Problem Solving: Cognition is the ability of an animal’s
nervous system to perceive, store, process, and use information gathered by
sensory receptors. Cognitive ethology examines the connection between
an animal’s nervous system & its behavior. In cognitive ethology, it has been
found that animals can categorize things as “same” and “different”. Problem
solving can be learned by observing the behavior of other animals.
Genetic and Environmental Interaction in Learning: Genetics and the
environment can interact to influence the learning process. Some birds have
learned songs, some are partially learned, others are completely genetic—
even birds that are raised away from others of their species develop an adult
song similar to others of it. Some birds have songs that are learned then
“crystallized” into one song; others are relearned and add sections.
Concept 51.4: Behavioral traits can evolve by natural selection
Introduction: Because of the influence of genes on behavior, natural selection can
result in the evolution of behavioral traits in populations.
Behavioral Variation in Natural Populations: When behavioral variation within a
species corresponds to variation in the environment, it may be evidence of past evolution.
Variation in Prey Selection: Differences in prey selection in populations of
garter snakes are due to prey availability and are evidence of behavioral
evolution. Inland snakes, which live where slugs are rare, refused them,
while coastal snakes of the same species readily ate them. Studies with
laboratory-born snakes supported that when inland snakes colonized coastal
regions, some snakes could recognize slugs as food, and thus were more fit
than their counterparts, and therefore were chosen by the environment to
reproduce.
Variation in Aggressive Behavior: Funnel spiders living in different habitats
exhibit differing degrees of aggressiveness in defense and foraging
(behavior associated with recognizing, searching for, capturing, and
consuming food) behavior. Spiders that live in arid climates are more
aggressive, and those that live in forests, rich with both food and predators,
are more timid, which was proven to be genetic through studies of laboratory-
born spiders. The respective traits were selected by the environment in which
the spiders live.
Experimental Evidence for Behavioral Evolution: Laboratory and field
experiments can demonstrate the evolution of behavior.
Laboratory Studies of Drosophila Foraging Behavior: Studies of Drosophila
populations raised in high- and low-density conditions show a clear
 divergence in behavior linked to specific genes. They prove that the sitter
allele had increased in low-density populations where short distance foraging
would yield enough food, and the rover allele had increased in high-density
areas, where larvae would need to move beyond areas of food depletion.

Migratory Patterns in Blackcaps: Field and laboratory studies of Blackcap birds

have documented a change in their migratory behavior. Birds placed in funnel
cages left marks indicating the direction they were trying to migrate, indicating
that migration was a genetic behavior selected by the environment the birds
were from. Migratory orientation of wintering adult birds captured in Britain was
very similar to that of laboratory-raised birds.

Concept 51.5: Natural selection favors behaviors that increase survival and reproductive
success
Introduction: The genetic components of behavior evolve through natural selection.
Behavior can affect fitness through its influence on foraging and mate choice.
Foraging Behavior: The optimal foraging theory views foraging behavior as a
compromise between the benefits of nutrition and the costs of obtaining food. According to
the theory, natural selection should favor foraging behavior that minimizes that costs of
foraging and maximizes the benefits.
Energy Costs and Benefits: Reto Zach conducted a cost-benefit analysis of feeding
behavior in crows. The crows eat mollusks called whelks but must drop them from
the air to crack the shells. Zach determined that the optimal flight height in foraging
behavior correlated with a fewer number of drops, indicating a trade-off between
energy gained (food) and energy expended. This supported the OF theory. The
feeding behavior of the bluegill sunfish does the same. In bluegill sunfish prey
selection behavior is related to prey density. When prey density is high, sunfish will
focus on larger prey because it is more efficient; at low prey densities, fish should
eat all prey to meet energy needs.
Rise of Predation: Research on mule deer populations has shown that predation
risk affects where the deer choose to feed. The deer that maximize gains &
 minimize losses as well as avoid predation have become more efficient when
foraging for food.
Mating Behavior and Mate Choice: Mating behavior is the product of a form of
natural selection call sexual selection
Mating Systems and Parental Care: The mating relationship between males
and females varies a great deal from species to species. In many species,
mating is promiscuous, with no strong pair-bonds or lasting relationships. In
monogamous relationships, one male mates with one female; in
polygamous relationships, an individual with one sex mates with several of
the other. In a system called polygyny, one male mates with many females
and the males are often more showy and larger than the females. In
polyandrous systems one female mates, with many males and the females
are often more showy than the males.
 Parent Care: The needs of the young are an important factor
constraining the evolution of mating systems. The certainty of paternity
influences parental care and mating behavior. In species that produce large
numbers of offspring. Parental care is at least as likely to be carried out by
males as females. The term certainty of paternity does not mean the
animals are aware of these factors; parental behavior correlated with CoP
has been reinforced by generations of natural selection.
Sexual Selection and Mate Choice: In intersexual selection members of
one sex choose mates on the basis of particular characteristics. Intrasexual
selection involves competition among members of one sex for mates.
 Mate Choice by Females: Mate preferences by females may play a
central role in the evolution of male behavior and anatomy through
intrasexual selection. Male zebra finches are more ornate than females, a
trait that may affect mate choice by the females. Results from studies
suggest that females imprint on fathers and that mate choice by females has
played a role in the evolution of male zebra finches. In stalk-eyed flies, the
size of eyestalks affects which males the females choose to mate with;
proving that female choice has been a selective pressure for the selection of
long eyestalks, because males with the inability to develop them often have
genetic disorders. In general, ornaments suggest health, and females choose
healthy mates in hope of creating healthy offspring.
 Mate Competition for Males: Male competition for mates is a source of
intrasexual selection that can reduce variation among males. Such
competition may involve agonistic behavior, an often ritualized contest that
determines which competitor gains access to a resource. Victories in such
contests may be psychological rather than physical. In species where more
than one mating behavior can result in successful reproduction, intrasexual
selection has led to the evolution of alternative male mating behavior and
morphology. Morphology affects the mating behavior in isopods of the same
species that are genetically distinct.
Applying Game Theory: Game theory evaluates alternative behavioral strategies
in situations where the outcome depends on each individual’s strategy and the strategy of
other individuals. Game theory is a way of thinking about evolution in situations where the
fitness of a particular behavior phenotype is influenced by other behavioral phenotypes in
the population. Mating success of male side-blotched lizards was found to be influenced by
male polymorphism and the abundance of different males in a given area. Game theory
deals with relative, not absolute, performance in terms of the efficiency of a behavior.
Concept 51.6: The concept of inclusive fitness can account for most altruistic behavior
Introduction: Many social behaviors are selfish. Natural selection favors behavior
that maximizes an individual’s survival and reproduction
Altruism: Some animals behave in ways that reduce their individual fitness but
increase the fitness of others. This kind of behavior is called altruism, or selflessness.
This is present in squirrels, bees, and naked moles rats. In naked mole rat populations,
non-reproductive individuals may sacrifice their lives protecting the reproductive individuals
from predators.
Inclusive Fitness: Altruistic behavior can be explained by inclusive fitness,
the total effect an individual has on proliferating its genes by producing its own offspring
and by providing aid that enables close relatives to produce offspring.
Hamilton’s Rule and Kin Selection: Hamilton proposed a quantitative
measure for predicting when natural selection would favor altruistic acts
among related individuals. The three key variables in an altruistic act are:
– The benefit to the recipient (B)
– The cost to the altruist (C)
– The coefficient of relatedness (r).
The coefficient of relatedness is the probability that two relatives may
share the same genes. Natural selection favors altruism when the benefit to
the recipient multiplied by the coefficient of relatedness exceeds the cost to
the altruist (rB > C). This inequality is called Hamilton’s rule. Kin selection
is the natural selection that favors this kind of altruistic behavior by
enhancing reproductive success of relatives. An example of kin selection and
altruism is the warning behavior observed in Belding’s ground squirrels.
Reciprocated Altruism: Altruistic behavior toward unrelated individuals can be
adaptive if the aided individual returns the favor in the future. This type of
altruism is called reciprocal altruism.
Social Learning: Social learning forms the roots of culture. Culture can be
defined as a system of information transfer through observation or teaching that
influences the behavior of individuals in a population. Culture transfer of information
can alter behavioral phenotypes and influence fitness.
Culturally based change happens much faster than
natural selection-based change.
Mate Choice Copying: Mate choice
copying is a behavior in which individuals in
a population copy the mate choice of others.
This type of behavior has been extensively
studied in the guppy Poecilia reticulata.
 Social Learning of Alarm Calls: Vervet monkeys produce a complex set of
alarm calls depending on the particular predator. Infant monkeys give
undiscriminating alarm calls at first but learn to fine-tune them by the time
they are adults. Infants probably learn to give the right call by observing other
members and social confirmation. Still, no other species comes close to
matching the social learning and cultural transmission that occurs among
humans.
Evolution and Human Culture: Human culture is related to evolutionary theory in
the distinct discipline of sociobiology. Sociobiology’s main premise is that certain
behavioral characteristics exist because they are expressions of genes that have been
perpetuated by natural selection. Human behavior, like that of other species is the result
of interactions between genes and environment. However, our social and cultural
institutions may provide the only feature in which there is no continuum between humans
and other animals.
Ololade Kolade
Dr. Kabak
AP Biology
Chapter 52: Population Ecology

Concept 52.1: Dynamic biological processes influence population density, dispersion, and
demography
Overview: To understand human population growth we must consider the general
principles of population ecology. Population ecology is the study of populations in relation
to the environment, including environmental influences on population density and
distribution, age structure, and variations in population size
Introduction: A population is a group of individuals of a single species living in a
specific area. Members of populations rely on the same resources & have a high likelihood
of interacting/breeding with one another. Populations can evolve.
Density and Dispersion: Density is the number of individuals per unit area or
volume. Dispersion is the pattern of spacing among individuals within the boundaries of
the population. Populations can be defined by both their density and dispersion, once
boundaries have been defined.
Density: A Dynamic Perspective: Determining the density of natural
populations is possible, but difficult to accomplish. In most cases, it is
impractical or impossible to count all individuals in a population. A method
used to estimate wildlife populations is the mark-recapture method.
 M-RC Method: Researcher places traps, marks animals, and then releases
them. After time, traps are set again, catching both unmarked and marked
individuals. From this, researchers can estimate total number of members in a
population. The method assumes that each marked individual has the same
probability to be capture as unmarked, which is not always true.
 Density as a Property: Density is the result of a dynamic interplay
between processes that add individuals to a population and those that remove
individuals from it. Additions to populations occur through birth and
immigration, the influx of new individuals from other areas. Decreases occur
through death and emigration, the movement of individuals out of a
population.
Patterns of Dispersion: Environmental and social factors influence the spacing of
individuals in a population.
--A clumped dispersion: is one in which individuals aggregate in patches,
most common, may be influenced by resource availability and behavior.
--A uniform dispersion: is one in which individuals are evenly distributed,
may be influenced by social interactions such as territoriality. Often a result of
territoriality, the defense of a bounded physical space against encroachment
by other individuals.
-- A random dispersion is one in which the position of each individual is
independent of other individuals. Occurs when there are no strong attractions
 or repulsions among individuals in an area where conditions are fairly
homogenous.
Demography: Factors that influence population density and dispersion also
influence other characteristics of a population. Demography is the study of the vital
statistics of a population and how they change over time. Death rates and birth rates are of
particular interest to demographers.
Life Tables: A life table is an age-specific summary of the survival pattern of
a population, and is best constructed by following the fate of a cohort, a
group of individuals of the same age, from birth until death. To build a life
table, the number of individuals that die in each age group needs to be
determined and the proportion of the cohort surviving from each age to the
next needs to be calculated. The life table of Belding’s ground squirrels
reveals many things about this population.
Survivorship Curves: A survivorship curve is a graphic way of representing
the data in a life table. It is a plot of the proportion or numbers in a cohort still
alive Type I Curve *Flat at start, low death
at each age. rates in early/ middle
Survivorship life
curves can *Decline steeply as
be classified death increases in older
into three age groups
general *Happens in
types: humans/large mammals
Type I, Type who care for few
II, and Type offspring
III. Most Type II Curve *Intermediate
species fall *Constant death rate
in between during life span
patterns, or *Occurs in Belding’s
show ground squirrels
more Type III Curve *Drops steeply at start,
complex high death rates for
ones. In young
populations *Flattens out as death
with out rates decrease for those
who survived to critical
age
emigration/immigration, survivorship is a key factor in determining
changes in population size.
 Reproductive Rates: Demographers focus on females in populations
because females produce offspring. They view populations in terms of
females giving rise to new females. To describe the reproductive pattern of a
population, one must determine how the reproductive output varies with the
ages of the females. A reproductive table or fertility schedule is an age-
specific summary of the reproductive rates in a population. It also describes
the reproductive patterns of a population and is best conducted following a
cohort. Multiplying the weaning proportion of females of a given age and the
number of female offspring gives the average number of female offspring for
each female in a given age class. Reproductive tables vary depending on
species.

Concept 52.2: Life history traits are products of natural selection

Introduction: Life history traits are evolutionary outcomes reflected in the
development, physiology, and behavior of an organism. The traits that affect an organism’s
schedule of reproduction and survival (birth-reproduction-death) compose its life history.
Life histories have 3 basic variables: when reproduction begins (age at first reproduction,
age at maturity), how often the organism reproduces and how many offspring are
produced in each reproductive episode.
Life History Diversity: Life histories are very diverse. Species that exhibit
semelparity, or “big-bang” reproduction reproduce a single time and die (pacific salmon,
agave plants), can be the result of an erratic climate. Species that exhibit iteroparity, or
repeated reproduction produce offspring repeatedly over time (lizards). Evolution of life
history patterns is largely due to offspring survival rate. When rate is low, semelparity is
favored; in dependable environments where competition may be intense, iteroparity is
prefer
“Trade-offs” and Life Histories: Organisms have finite resources which may lead to
trade-offs between survival and reproduction. Some plants produce a large number of
small seeds ensuring that at least some of them will grow and eventually reproduce. Other
types of plants produce a moderate number of large seeds that provide a large store of
energy that will help seedlings become established. Parental care of smaller broods may
also facilitate survival of offspring.

Concept 52.3: The exponential model describes population growth in an idealized,

unlimited environment
 Introduction: It is useful to study population growth in an idealized situation in
order to understand the capacity of species for increase and the conditions that may
facilitate this type of growth
Per Capita Rate of Increase: If immigration and emigration are ignored, a
population’s growth rate (per capita increase) equals birth rate minus death rate. Change
in population during a fixed time period can be determined with (N=population size,
t=time):

To covert the model into one where births and deaths are averages, per captia birth &
death rates are used. The per capita birth rate is the # of offspring produced per unit
of time by an average member of the population. To calculate the expected number of
births per certain measure of time, use the per capita birth rate of that time measure
and multiply it by the population size: B=bN. The per capita death rate allows the
expected number of deaths per unit time to be calculated (M=mN). Per capita birth and
death rates can be calculated from life tables and reproductive tables.

The per capita rate of increase is the difference between the per capita birth and
death rate (r=b-m). Zero population growth occurs when the per capita birth rate
equals the per capita death rate. Change in population size using the per capita rate of
increase:

Exponential Growth: Exponential population growth is population increase

under idealized conditions. Under these conditions the rate of reproduction is at its
maximum, called the intrinsic rate of increase. Under these conditions the rate of
reproduction is at its maximum, called the intrinsic rate of increase. The equation of
exponential population growth is:

Exponential population growth results in a J-shaped curve. The J-shaped curve of

exponential growth is characteristic of some populations that are rebounding.

Concept 52.4: The logistic growth model includes the concept of carrying capacity
Introduction: Exponential growth cannot be sustained for long in any population. A
more realistic population model limits growth by incorporating carrying capacity.
Carrying capacity (K) is the maximum population size a particular environment can
support. Carrying capacity is not fixed, and varies over space, time, and abundance of
limiting to factors.
The Logistic Growth Model: In the logistic population growth model, the per
capita rate of increase declines as carrying capacity is reached. We construct the
logistic model by starting with the exponential model and adding an expression that
reduces the per capita rate of increase as N increases. The logistic growth equation
includes K, the carrying capacity.

When N is small compared to K, the term (K-N)/K is large and the per capita rate of
increase is close to the maximum rate of increase. When N is large then (K-N)/K is
small, so is the rate of increase. When N equals K the population stops growing. New
individuals are added most rapidly at intermediate population sizes, where there is not
only a substantial breeding population but available space and resources. The logistic
model of population growth produces a sigmoid (S-shaped) curve. Growth rate slows
as N approaches K.
The Logistic Model and Real Populations: The growth of laboratory populations
of paramecia fits an S-shaped curve. Some populations overshoot K before settling
down to a relatively stable density. Some populations fluctuate greatly around K,
making it difficult to define a carrying capacity. The logistic model fits few real
populations but is useful for estimating possible growth.

The logistic model assumes that each individual added to a population has the same
negative effect on population growth; however some populations display an Allee
effect, in which individuals may have a more difficult time surviving or reproducing if
the population size is too small.
The Logistic Model and Life Histories: Life history traits favored by natural
selection may vary with population density and environmental conditions. K-selection,
or density-dependent selection, selects for life history traits that are sensitive to
population density; r-selection, or density-independent selection, selects for life history
traits that maximize reproduction. The concepts of K-selection and r-selection are
somewhat controversial and have been criticized by ecologists as oversimplifications.

Concept 52.5: Populations are regulated by a complex interaction of biotic and abiotic
influences.
Introduction: There are two general questions we can ask about regulation of
population growth:
 1. What environmental factors stop a population from growing?
2. Why do some populations show radical fluctuations in size over time, while
others remain stable?
Population Change and Population Density: If emigration and immigration offset
each other, then a population grows when birth rate exceeds death rate and declines
when the death rate exceeds birth rate. In density-independent populations birth rate
and death rate do not change with population density. In density-dependent
populations, birth rates fall and death rates rise with population density. Determining
equilibrium for population density:

Density-Dependent Population Regulation: Density-dependent birth and death

rates are an example of negative feedback that regulates population growth and are
affected by many different mechanisms.
Competition for Resources: In crowded populations, increasing population
density intensifies intraspecific competition for resources, resulting in a lower
birth rate. Crowding can reduce seed production of plants, and food can limit
the amount of eggs laid by songbirds in high density populations.
Territoriality: In many vertebrates and some invertebrates territoriality may
limit density. Territory space is the resource for which individuals compete.
Cheetahs are highly territorial, using chemical communication to warn other
cheetahs of their boundaries. Oceanic birds exhibit territoriality in nesting
behavior. Evidence of non-breeding individuals in a population is an
indication that territoriality is restricting population growth.
Health: Population density can influence the health and survival of
organisms. In dense populations, pathogens can spread more rapidly
Predation: As a prey population builds up, predators may feed preferentially
on that species. Predation can be a cause of density-dependent mortality if a
predator encounters and captures more food.
Toxic Wastes: The accumulation of toxic wastes can contribute to density-
dependent regulation of population size. Metabolic by-products accumulate
as populations grow, poisoning organisms’ environment.
Intrinsic Factors: For some populations intrinsic (physiological) factors
appear to regulate population size. In populations of white-footed mice,
 aggressive interactions will increase with population density, even when food
and shelter are in abundance. A stress syndrome causes reproductive
organs to shirk, which ultimately causes an increase in mortality and a
decreased birth rate.
Population Dynamics: All populations show numerical fluctuations; the changes
are more interesting than averages. The study of population dynamics focuses on the
complex interactions between biotic and abiotic factors that cause variation in
population size.
Stability and Fluctuation: Long-term population studies have challenged the
hypothesis that populations of large mammals are relatively stable over time.
Extreme fluctuations in population size are typically more common in
invertebrates than in large mammals.

VS

Metapopulations and Immigration: A metapopulation is a subdivided

population of a single species. They can be linked through emigration and
immigration. High levels of immigration combined with higher survival can
result in greater stability in populations. The metapopulation concept
underscores the significance of immigration and emigration in understanding
populations in patchy habitats.
Population Cycles: Many populations undergo regular boom-and-bust cycles.
Boom-and-bust cycles are influenced by complex interactions between biotic and
abiotic factors. In the case of snowshoe hares and lynxes, the 10 year boom-and-bust
cycles may be due to winter food shortage and predator-prey interaction.

Concept 52.6: Human population growth has

slowed after centuries of exponential increase
 Introduction: No population can grow indefinitely and humans are no exception.
The Global Human Population: The human population increased relatively slowly
until about 1650 and then began to grow exponentially. Though the global population is
still growing, the rate of growth began to slow approximately 40 years ago.

To

Regional Patterns of Population Change: To maintain population stability, a

regional human population can exist in one of two configurations:
Zero population growth = High birth rate – High death rate
or
Zero population growth = Low birth rate – Low death rate
The demographic transition is the move from the first toward the second
state. The demographic transition is associated with various factors in
developed and developing countries, including the education of women,
voluntary contraception/family planning, and strict birthing policies. Reduced
family size is the key to demographic transition.
Age Structure: One important demographic factor in present and future
growth trends is a country’s age structure, the relative number of individuals
at each age. Age structure is commonly represented in pyramids. Age
structure diagrams can predict a population’s growth trends, illuminate
social conditions, and help us plan for the future.
Infant Mortality and Life Expectancy: Infant mortality is the number of infant
deaths per 1,000 live births and life expectancy is at birth is the predicated
average length of life at birth. Infant mortality and life expectancy at birth vary
 widely among developed and developing countries but do not capture the
wide range of the human condition.
Global Carrying Capacity: Just how many humans can the biosphere support?
Estimates of Carrying Capacity: The carrying capacity of Earth for humans is
uncertain, and many things must be taken into consideration. Some
scientists use curves like that produced by the logistic equation; others
generalize using maximum population density and multiply this by the area of
habitable land.
Ecological Footprint: The ecological footprint concept summarizes the
collective land and water area needed to sustain the people of a nation and
is one measure of how close we are to the carrying capacity of Earth. Six
ecologically productive areas are distinguished when calculating ecological
foot print: arable land, pasture, forest, built-up land and fossil energy land. All
measures are converted to hectares (2.47 acres) per person. Ecological
footprints for 13 countries show that the countries vary greatly in their
footprint size and their available ecological capacity (the resource base of
each country). At more than 6 billion people, the world is already in
ecological deficit.

Ololade Kolade
Dr. Kabak
AP Biology

Chapter 53: Community Ecology

Concept 53.1: A community’s interactions include competition, predation, herbivory,
symbiosis, and disease
Overview: A biological community is a group of populations of various species
living close enough for potential interaction. For example, the various animals and plants
surrounding a watering hole are all members of a savanna community in southern Africa.
Introduction: Populations are linked by interspecific interactions that affect the
survival and reproduction of the species engaged in the interaction. Interspecific
interactions are interactions with other species within a community. Interspecific
interactions can have differing effects on the populations involved.

Competition: Interspecific competition occurs when species compete for a

particular resource that is in short supply. Strong competition can lead to competitive
exclusion: the local elimination of one of the two competing species.
The Competitive Exclusion Principle: The competitive exclusion principle
states that two species competing for the same limiting resources cannot
coexist in the same place. One species will use the resources more
efficiently and thus reproduce more rapidly than the other, eliminating the
inferior species.
Ecological Niches: The ecological niche is the total of an organism’s use of
the biotic and abiotic resources in its environment. The niche concept allows
restatement of the competitive exclusion principle two species cannot coexist
in a community if their niches are identical. However, ecologically similar
species can coexist in a community if there is one or more significant
difference in their niches. As a result of competition a species’ fundamental
niche (the niche it may potentially occupy) may be different from its realized
niche (the niche it does).
Resource Partitioning: Resource partitioning is the differentiation of niches
that enables similar species to coexist in a community. Resource partitioning
is indirect evidence of earlier interspecific competition solved by the evolution
of niche differentiation.
Character Displacement: In character displacement, there is a tendency for
characteristics to be more divergent in sympatric (geographically
overlapping) populations of two species than in allopatric (geographically
separate) populations of the same two species. Allopatric species tend to be
morphologically similar and use similar resources; sympatric species could
potentially compete, but tend to be different and use different resources.
Predation: Predation refers to a interaction where one species, the predator, kills
and eats the other, the prey. Feeding adaptations of predators include: claws, teeth,
 fangs, stingers, acute senses, and poison; animals also display a great variety of
defensive adaptations. In some cases, one prey species may gain significant protection
by mimicking the appearance of another

Cryptic Coloration *Camouflage makes prey difficult to

spot
Aposematic Coloration *Animals with effective chemical
defenses often exhibit bright warning
coloration to warn predators to stay
away
Batesian Mimicry *A palatable or harmless species
mimics an unpalatable or harmful
model

Müllerian Mimicry *Two or more unpalatable species

resemble each other
*Each species gains an
advantageThe greater number of
that prey with that
appearancepredators learn to
avoid it type of aposematic
coloration

Herbivory: Herbivory is the process in which an herbivore eats parts of a plant. It

has led to the evolution of plant mechanical and chemical defenses and consequent
adaptations by herbivores. Most herbivores are small invertebrates, while there are some
large mammalian species.
Parasitism: In parasitism, one organism, the parasite, derives its nourishment
from another organism, its host, which is harmed in the process. Parasites that live within
the body of their hosts are endoparasites. Ectoparasites feed on the external surface of
a host. In parasitoidism, insects lay eggs on or in living hosts. The larvae feed on the host
and kill it. Parasitism exerts substantial influence on the survival, reproduction, and density
of populations and the structure of communities. Parasites can cause hosts to engage in
atypical behaviors, and have a complex life cycle including many hosts.
Disease: The effects of disease on populations and communities are similar to that
of parasites. Pathogens, disease-causing agents, cause effects similar to parasites in
their hosts. Pathogens are typically bacteria, viruses, or protists. Pathogens have been the
study of few ecological studies, but prove lethal more often than parasites, drastically
lowering the numbers of populations.
Mutualism: Mutualistic symbiosis, or mutualism, is an interspecific interaction that
benefits both species. Mutualistic relationships sometimes involve the evolution of related
adaptations in both species. Changes in one species is likely to affect the survival and
reproduction of the other (ex: nectar in plants).
Commensalism: In commensalism, one species benefits and the other is not
affected. Commensal interactions have been difficult to document in nature because any
close association between species likely affects both species.
 Interspecific Interactions and Adaptation: Evidence for coevolution, which
involves reciprocal genetic change by interacting populations, is scarce. A change in one
species acts as a selective pressure in one species, which then serves to act as a
selective pressure on the first species. However, generalized adaptation of organisms to
other organisms in their environment is a fundamental feature of life. Much evidence
proves that competition and predation are key process driving community dynamics. Few
interspecific studies have been conducted in tropical environments.

Concept 53.2: Dominant and keystone species exert strong controls on community
structure
Introduction: In general, a small number of species in a community exert strong
control on that community’s structure.
Species Diversity: The species diversity of a community is the variety of different
kinds of organisms that make up the community. It has two components: species
richness, the total number of different species in the community and relative abundance,
the proportion each species represents of the total individuals in the community. Two
different communities can have the same species richness, but a different relative
abundance. A community with an even species abundance is more diverse than one in
which one or two species are abundant and the remainder rare.
Trophic Structure: Trophic structure is the feeding relationships between
organisms in a community and is a key factor in community dynamics. Food chains link
the trophic levels from producers to top carnivores. Decomposers “feed” on organisms
from all trophic levels.

Food Webs: A food web is a branching food chain with complex trophic
interactions. They are composed of linked food chains, with arrows showing
who eats who. Food webs can be simplified by isolating a portion of a
community that interacts very little with the rest of the community or grouping
several species under one title. Food links are linked into food chains
because a given species may weave into the web at more than one trophic
level.
 Limits on Food Chain Length: Each food chain in a food web is usually only a
few links long. There are two hypotheses that attempt to explain food chain
length. The energetic hypothesis suggests that the length of a food chain is
limited by the inefficiency of energy transfer along the chain. The dynamic
stability hypothesis proposes that long food chains are less stable than
short ones. Population fluctuations at lower trophic levels are magnified at
higher levels, causing the possible extinction of top predators. Most of the
available data supports the energetic hypothesis.
Species with a Large Impact: Certain species have an especially large impact on
the structure of entire communities either because there is a lot of them or obecause they
play a pivotal role in community dynamics.
Dominant Species: Dominant species are those species in a community
that are most abundant or have the highest biomass (total mass of all
individuals in a population). They exert powerful control over the occurrence
and distribution of other species. One hypothesis suggests that dominant
species are most competitive in using limited resources. Another hypothesis
says that they are most successful at avoiding predators/disease. This idea
explains the high biomass of invasive species (species that take hold
outside their native range) can get in environments lacking their natural
predators & pathogens. One way to study the impact on a dominant species
on a community is to remove it.
Keystone Species: Keystone species are not necessarily abundant in a
community but exert strong control on a community by their ecological roles,
or niches. Field studies of sea stars exhibit their role as a keystone species in
intertidal communities. Observation of sea otter populations and their
predation shows the effect the otters have on ocean communities.
Ecosystem “Engineers” (Foundation Species): Some organisms exert their
influence by causing physical changes in the environment that affect
community structure. For example, beaver dams can transform landscapes
on a very large scale, creating flooded wetlands. Some foundation species
act as facilitators that have positive effects on the survival and reproduction
of some of the other species in the community.
Bottom-Up and Top-Down Controls: There are two possible methods of
community organization.
Bottom-Up: The bottom-up model of community organization proposes a
unidirectional influence from lower to higher trophic levels. In this case, the
presence or absence of abiotic nutrients determines community structure,
including the abundance of primary producers. If you add/remove predators
in from a bottom-up community, the effects will not extend to lower trophic
levels.
Top-Down: The top-down model of community organization proposes that
control comes from the trophic level above. In this case, predators control
through consumption herbivores which in turn control primary producers. The
top-down model is also called the trophic cascade model. The effects of
any manipulation moves down the trophic structure as a series of +/- effects.
Intermediates between the two models may exist. Also, communities may shift from
bottom-up to top-down over time. Pollution can affect community dynamics but through
biomanipulation and use of community models, polluted communities can be restored.
Concept 53.3: Disturbance influences species diversity and composition
Introduction: Most ecologists once thought that communities were a state of
equilibrium, and remained so, unless humans disturbed them. Emphasis on change has
led to a nonequilibrium model which describes communities as constantly changing after
being buffeted by disturbances.
What Is Disturbance? A disturbance is an event that changes a community. It
removes organisms from a community and alters resource availability. Fire is a significant
disturbance in most terrestrial ecosystems and is often a necessity in some communities.
The intermediate disturbance hypothesis suggests that moderate levels of disturbance
can foster higher species diversity than low or high levels of disturbance. Studies show
that several communities are nonequilibrium communities.
Human Disturbance: Humans are the most widespread agents of disturbance.
Human disturbance to communities usually reduces species diversity. Humans also
prevent some naturally occurring disturbances which can be important to community
structure.
Ecological Succession: Ecological succession is the sequence of community
and ecosystem changes after a disturbance. When the process begins on a lifeless area or
where no soil exists it is called primary succession. During primary succession, often the
only life-forms present are autotrophic prokaryotes, which are overtaken by bryophytes,
which are then overtaken by shrubs and grasses after soil develops. Secondary
succession begins in an area where soil remains after a disturbance, such as after a large
fire. The area often returns to its original state. Early-arriving species may: facilitate the
appearance of later species, inhibit the growth of the later species, or tolerate later species
and have no impact on their establishment. Retreating glaciers provide a valuable field-
research opportunity on succession.

Concept 53.4: Biogeographic factors affect community biodiversity

Introduction: Two key factors correlated with a community’s species diversity are
its geographic location and its size.
Equatorial-Polar Gradients: The two key factors in equatorial-polar gradients of
species richness are probably evolutionary history and climate. Species richness generally
declines along an equatorial-polar gradient and is especially great in the tropics. The
greater age of tropical environments may account for the greater species richness
because speciation occurs about five times faster.
Climate in Equatorial-Polar Gradients: Climate is likely the primary cause of
the latitudinal gradient in biodiversity. The two main climatic factors
correlated with biodiversity are solar energy input and water availability.
These factors can be considered together by measuring the community’s
rate of evapotranspiration, the evaporation of water from the soil plus the
transpiration of water from plants. Actual evapotranspiration is determined
by solar radiation, temperature, and water availability and is very high in hot
areas with abundant rainfall. Potential evapotranspiration is a measure of
energy availability and is determined by solar radiation and temperature and
is highest in regions of high solar radiation/temperature.
Area Effects: The species-area curve quantifies the idea that all other factors
being equal, the larger the geographic area of a community, the greater the number of
species.
 Island Equilibrium Model: Species richness on islands depends on island size,
distance from the mainland, immigration, and extinction. The equilibrium model of island
biogeography maintains that species richness on an ecological island levels off at some
dynamic equilibrium point.

Concept 53.5: Contrasting views of community structure are the subject of continuing
debate
Introduction: Two different views on community structure emerged among
ecologists in the 1920s and 1930s.
Integrated and Individualistic Hypotheses:
Integrated: The integrated hypothesis of community structure describes a
community as a group of closely linked species, locked into association by
mandatory biotic interactions. This ultimately causes the community to
function as one single unit. The integrated hypothesis predicts that the
presence or absence of particular species depends on the presence or
absence of other species.
Individualistic: The individualistic hypothesis of community structure
proposes that communities are loosely organized associations of
independently distributed species with the same abiotic requirements. The
individualistic hypothesis predicts that each species is distributed according
to its tolerance ranges for abiotic factors. In most actual cases the
composition of communities seems to change continuously, with each
species more or less independently distributed.
Rivet and Redundancy Models: The rivet model of communities suggests that
all species in a community are linked together in a tight web of interactions. It also
states that the loss of even a single species has strong repercussions for the entire
community. The redundancy model of communities proposes that if a species is
lost from a community, other species will fill the gap. Generally, community
hypotheses and models represent extremes, and that most communities probably
lie somewhere in the middle.
Ololade Kolade
Dr. Kabak
AP Biology

Chapter 54: Ecosystems

Concept 54.1: Ecosystem ecology emphasizes energy flow and chemical cycling
Overview: An ecosystem consists of all the organisms living in a community as
well as all the abiotic factors with which they interact. Ecosystems can vary greatly in size,
and can be natural or human dominated. Ecosystem dynamics involve two main
processes: energy flow and chemical cycling Energy flows through ecosystems while
matter cycles within them.
Introduction: Ecosystem ecologists view ecosystems as transformers of energy
and processors of matter.
Ecosystems and Physical Laws: The laws of physics and chemistry apply to
ecosystems, especially in regard to the flow of energy. Energy is conserved but degraded
to heat during ecosystem processes (principle of conservation, 2nd law of
thermodynamics).
Trophic Relationships: Energy and nutrients pass from primary producers
(autotrophs) to primary consumers (herbivores) and then to secondary consumers
(carnivores). Primary producers are autotrophs, the trophic level that supports all others.
Herbivores, that eat primary producers, are primary consumers. Carnivores that eat
herbivores are secondary consumers; carnivores that eat other carnivores are tertiary
consumers. Detritivores/decomposers are consumers that get their energy from
detritus, non-living organic material. Nutrients cycle within an ecosystem. Energy flows
through an ecosystem entering as light and exiting as heat.

Decomposition: Decomposition connects all trophic levels. Detritivores, mainly

bacteria and fungi, recycle essential chemical elements by decomposing organic material
and returning elements to inorganic reservoirs. Decomposition by prokaryotes and fungi
accounts for most of the conversion of organic materials from all trophic levels to inorganic
compounds usable by primary producers.
Concept 54.2: Physical and chemical factors limit primary production in ecosystems
Introduction: Primary production in an ecosystem is the amount of light energy
converted to chemical energy by autotrophs during a given time period.
Ecosystem Energy Budgets: The extent of photosynthetic production sets the
spending limit for the energy budget of the entire ecosystem.
The Global Energy Budget: The amount of solar radiation reaching the
surface of the Earth limits the photosynthetic output of ecosystems. Only a
small fraction of solar energy actually strikes photosynthetic organisms.
Gross and Net Primary Production: Total primary production in an
ecosystem is known as that ecosystem’s gross primary production (GPP).
This is the amount of light energy converted to chemical energy by
photosynthesis per unit of time. Not all of this production is stored as organic
material in the growing plants because some is used in cellular respiration.
Net primary production (NPP) is equal to GPP minus the energy used by
the primary producers for respiration (R). Only NPP is available to
consumers. NPP=GPP-R. Different ecosystems vary considerably in their net
primary production and in their contribution to the total NPP on Earth.
Overall, terrestrial ecosystems contribute about two-thirds of global NPP and
marine ecosystems about one-third.
Primary Production in Marine and Freshwater Ecosystems: In marine and
freshwater ecosystems, light and nutrients are important in controlling primary production.
Light Limitation: The depth of light penetration affects primary production
throughout the photic zone of an ocean or lake.
Nutrient Limitation: More than light, nutrients limit primary production in
different geographic regions of the ocean and in lakes. A limiting nutrient is
the element that must be added in order for production to increase in a
particular area. Nitrogen and phosphorous are typically the nutrients that
most often limit marine production.
 The Addition of Nutrients: Experiments in another ocean region
showed that iron limited primary production. The iron factor in marine
ecosystems is related to the nitrogen factor because adding iron stimulates
the growth of cyanobacteria that fix nitrogen. The addition of large amounts
of nutrients to lakes has a wide range of ecological impacts. In some areas,
sewage runoff has caused eutrophication (increase of phosphorus and
nitrogen, causing a growth of algae) of lakes, which can lead to the eventual
loss of most fish species from the lakes.
Primary Production in Terrestrial and Wetland Ecosystems: In terrestrial and
wetland ecosystems climatic factors, such as temperature and moisture, affect primary
production on a large geographic scale. The contrast between wet and dry climates can be
represented by a measure called actual evapotranspiration. Actual evapotranspiration is
the amount of water annually transpired by plants and evaporated from a landscape and is
related to net primary production. It increases with precipitation and amount of solar
energy. On a more local scale, soil nutrient is often the limiting factor in primary
production. Nitrogen and phosphorus are often limiting factors in primary production; a
certain nutrient may limit production even if another is in abundance.
Concept 54.3: Energy transfer between trophic levels is usually less than 20% efficient
Introduction: The secondary production of an ecosystem is the amount of
chemical energy in consumers’ food that is converted to their own new biomass during a
given period of time (growth).
Production Efficiency: The production efficiency of an organism is the fraction
of energy stored in food that is not used for respiration. Net secondary production is the
energy stored in biomass represented by growth and reproduction. Assimilation is the
total energy taken in and used for growth, reproduction, and respiration.
The efficiency of animals as energy transformers can be calculated with:

Trophic Efficiency and Ecological Pyramids: Trophic efficiency is the

percentage of production transferred from one trophic level to the next. It
usually ranges from 5% to 20%. Trophic efficiency is less than production
efficiency because it takes into account energy lost through respiration and
contained through feces as well as the energy in a lower trophic level that is
not consumed by the next trophic level.
 Pyramids of Production: This loss of energy with each transfer in a
food chain can be represented by a pyramid of net production. Each
block is proportional to net production.

 Pyramids of Biomass: One important ecological consequence of low

trophic efficiencies can be represented in a biomass pyramid. Each tier
represents the standing crop (total dry weight of all organisms) in a given
trophic level. Most biomass pyramids show a sharp decrease at
successively higher trophic levels. Certain aquatic ecosystems have
inverted biomass pyramids due to a short turnover time in phytoplankton
—they have a small standing crop biomass compared to their production.

 Pyramids of Numbers: A pyramid of numbers represents the number

of individual organisms in each trophic level. The dynamics of energy
flow through ecosystems have important implications for the human
 population. Eating meat is an inefficient way of tapping photosynthetic
production. Worldwide agriculture could successfully feed many more
people if humans all fed more efficiently, eating only plant material.
The Green World Hypothesis: The green world hypothesis states that terrestrial
herbivores consume relatively little plant biomass because they are held in check by a
variety of factors. Most terrestrial ecosystems have large standing crops despite the large
numbers of herbivores. The green world hypothesis proposes several factors that keep
herbivores in check:
– Plants have defenses against herbivores. For example, spines and
noxious chemicals.
– Nutrients, not energy supply, usually limit herbivores. Animals need
protein, which plants supply in small amounts.
– Abiotic factors limit herbivores. Seasonal changes can set a low carry
capacity in an environment.
– Intraspecific competition can limit herbivore numbers. Competition can
maintain population densities much lower than a number that would strip
vegetation.
– Interspecific interactions check herbivore densities. Predators, parasites,
and disease are the most important factors in limiting the growth of herbivore
populations.
Concept 54.4: Biological and geochemical processes move nutrients between organics
and inorganic parts of the ecosystem
Introduction: Life on Earth depends on the recycling of essential chemical
elements. Nutrient circuits that cycle matter through an ecosystem involve both biotic and
abiotic components and are often called biogeochemical cycles.
A General Model of Chemical Cycling: Gaseous forms of carbon, oxygen, sulfur,
and nitrogen occur in the atmosphere and cycle globally. Less mobile elements, including
phosphorous, potassium, and calcium cycle on a more local level, mainly in soil. A general
model of nutrient cycling includes the main reservoirs of elements and the processes that
transfer elements between reservoirs. Reservoirs are defined by two characteristics:
organic/inorganic materials and whether or not the materials are directly available for use
by organisms. All elements cycle between organic and inorganic reservoirs.
Ecosystems often exchange elements with other ecosystems; ecologists can trace
elements by adding a small amount of a radioactive isotope and tracking it, or tracking a
stable, naturally occurring isotope.
Biogeochemical Cycles:
Water moves in a global cycle driven by solar energy. The carbon cycle reflects the
reciprocal processes of photosynthesis and cellular respiration.
Most of the nitrogen cycling in natural ecosystems involves local cycles between
organisms and soil or water. The phosphorus cycle is relatively localized.

Decomposition and Nutrient Cycling Rates: Decomposers (detritivores) play a

key role in the general pattern of chemical cycling. The rates at which nutrients cycle in
different ecosystems are extremely variable, mostly as a result of differences in rates of
decomposition. The rate of decomposition in terrestrial ecosystems increases with actual
evapotranspiration.