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Chapter 51: Animal Behavior

Behavior: action carried out by muscles under control of the nervous system
Niko Tinbergens 4 Questions:
1. What stimulus elicits the behavior, and what physiological mechanisms mediate the
2. How does the animals experience during growth and development influence the
3. How does the behavior aid survival and reproduction?
4. What is the behaviors evolutionary history?
Proximate causation: how a behavior occurs or is modified (1 & 2)
Ultimate causation: why a behavior occurs in the context of natural selection (3 & 4)
*Concept of ultimate causation is central to behavioral ecology: study of the ecological
and evolutionary basis for animal behavior
Fixed Action Patterns
- A sequence of unlearned acts directly linked to a simple stimulus (e.g. Male
sticklebacks aggressive behavior & red underbellies)
o Unchangeable and carried to completion
o Sign stimulus: external cue that triggers behavior (e.g. red color)
- A regular, long-distance change in location
o Environmental stimuli can trigger behaviors and provide cues that animals use
to carry out behaviors
o Circadian clock: internal mechanism that maintains a 24-hour activity
rhythm/cycle (e.g. animals track position relative to sun)
o Detection of magnetic field: cells in pigeons brainstem encode info about
magnetic field direction, intensity, and polarity
Magnetoreceptors in eye, beak, and inner ear
Behavioral Rhythms
- Circannual rhythms: behavioral rhythms linked to the yearly cycle of seasons (e.g.
migration and reproduction)
o Influenced by the periods of daylight and darkness in the environment (some
linked to phases of the moon e.g. fiddler crab)
Animal Signals and Communication
- Signal: a stimulus transmitted from one organism to another
- Communication: the transmission and reception of signals between animals
Forms of Animal Communication
*Four modes of animal communication: visual, chemical, tactile, and auditory
Fruit fly
- Courtship constitutes a stimulus-response chain: the response to each stimulus is
itself the stimulus for the next behavior
- (1) Orienting; (2) Tapping; (3) Singing
- The form of communication that evolves is closely related to an animals lifestyle and
environment (e.g. most terrestrial animals are nocturnal so they use olfactory +
auditory signals instead of visual)
1. Workers cluster around a recently returned bee
2. Round dance: Food is near bees leave hive and search nearby for food
3. Waggle dance: Food is distant

a. Waggle dance resembles a figure eight. The number of waggles in straight the
run indicates distance & direction is indicated by the angle of the straight run.
Pheromones: chemical substances that can be used for communication
- Especially common among mammals and insects and relate to reproductive behavior
o Bees: queen + daughters produce pheromones
Attracts workers to queen, inhibits development of ovaries in workers,
attracts males
- Serve as alarm signals
o Injured fish releases substances that induce fright response in other fish
Minnows aggregate near bottom of environment after introduction of
alarm substance and reduce their movement
Innate Behavior
Behavior that is developmentally fixed (e.g. when all individuals in a population behave
- Cross-fostering study: the young of one species are placed in the care of adults from
another species
o Extent of which the offsprings behavior changes provides measure of how the
social and physical environment influences behavior (e.g. mouse studies of
aggressiveness and paternal behavior)
Finding: influence of experience on behavior can be passed on to
Experience during development can modify physiology that alters
parental behavior
- Twin study in humans
- The modification of behavior as a result of specific experiences; depends on nervous
system organizations
o Involves the formation of memories by changes in neuronal connectivity
- Imprinting: the establishment of a long-lasting behavioral response to a particular
individual or object
o Takes place only during specific time period in development: sensitive
During this period, young imprint on their parent and learn basic
o Formation of pair-bond (strong attachment) is crucial
Spatial Learning and Cognitive Maps
- Spatial learning: establishment of a memory that reflects the environments spatial
structure (e.g. female digger wasp)
o Animals use landmarks to find places such as nests
- Cognitive Map: a representation in animals nervous system of the spatial
relationships between objects in its surroundings (e.g. birds keep track of half-way
point between landmarks rather than distance to find food stores)
Associative Learning
- The ability to associate one environmental feature (e.g. color) with another (e.g. foul
o Classical conditioning: an arbitrary stimulus becomes associated with a
particular outcome
Dog would salivate to ringing of bell, which was associated w/food

Operant conditioning (trial-and-error): an animal first learns to associate one

of its behaviors with a reward or punishment and ten tends to repeat or avoid
that behavior
Rat learns through trial and error to obtain food by pressing a lever

Cognition and Problem Solving

- Cognition: the process of knowing that involves awareness, reasoning, recollection
and judgment
- Abstract thinking maze:
o Bees were trained in a color maze and one group was rewarded for choosing
the same color as the stimulus
o Bees were tested in a pattern maze. If previously rewarded for choosing the
sane color, bees most often chose lines oriented the same was as the stimulus
o Bees able to distinguish between same and different
- Problem solving: the cognitive activity of devising a method to proceed from one
state to another in the face of real or apparent obstacles
Development of Learned Behaviors
- Song learning in birds
o White-crowned sparrow: different stages of song learning; 1st stage takes place
early in life
o Memorizes song during sensitive period; then, final song crystallizes
- Social Learning: type of learning through observing others (e.g. vervet monkey calls)
- Culture: system of information transfer through social learning or teaching that
influences the behavior of individuals in a population
Evolution of Foraging Behavior
- Foraging: food-obtaining behavior
- Drosophila: Rover and sitter allele; rover allele higher in denser populations
whereas sitter allele higher in low density populations
Optional Foraging Model: proposes that foraging behavior is a compromise between the
benefits of nutrition and the costs of obtaining food
- Costs include energy expenditure of foraging and risk of being eaten
- Natural selection should favor a foraging behavior that minimizes the costs of
foraging and maximizes the benefits (e.g. mule deer)
Mating Systems and Sexual Dimorphism
- Mating systems: length and number of relationships between males and females
- Promiscuous mating: no strong pair-bonds
- Monogamous: one male mating with one female
- Polygamous: an individual of one sex mating with several of the other
o Polygyny: single male and many females
o Polyandry: single female and multiple males
- Sexual dimorphism: the extent to which males and females differ in appearance
o Monogamous: females and males look very similar
o Polygamous: the sex that attracts multiple partners is showier
o Results from sexual selection
Intersexual selection: members of one sex chooses mates on the basis
of the other sex (e.g. courtship songs)
Intrasexual selection: competition between members of one sex
- Certainty of paternity: relatively low in most species w/internal fertilization
o High in species w/external fertilization (egg layer + mating occurs together)
Mate choice

Females choose healthier, more showy males

Mate choice can be influenced by imprinting (e.g. zebra finches)
Mate-choice copying: a behavior in which individuals in a population copy the mate
choice of others (e.g. guppies and male orange color choice)
Male vs. male: competition can reduce variation among males b/c it involves
o Agonistic behavior: often-ritualized contest that determines which competitor
gains access to a resources, such as food or mates

Game Theory
- Evaluates alternative strategies in situations where the outcome depends on the
strategies of all the individuals involved
- Example: side-blotched lizard w/orange, blue, and yellow throats
Genetic Basis of Behavior:
- Drosophila and fru gene association w/courtship ritual
- Male meadow voles vs. male prairie voles in vasopressin brain receptors and pairbond connections
Genetic Variation and the Evolution of Behavior
- Case Study: Garter Snakes and Banana Slugs
- Case Study: Blackcaps Variation in Migratory Patterns
- A behavior that reduces an animals fitness but increases the fitness of other
individuals in the population
o E.g. squirrels, honeybees, and naked-mole rats
Inclusive Fitness: the total effect an individual has on proliferating its genes by producing
its own offspring and providing aid that enables other close relatives to produce offspring
Hamiltons Rule and Kin Selection
- Hamiltons hypothesis: a way to measure the effect of altruism on fitness
o Benefit of the recipient (B)
Average number of extra offspring that the recipient of an altruistic act
o Cost to the altruist (C)
How many fewer offspring the altruist produces
o Coefficient of relatedness
The fraction of genes that, on average, are shared
- Natural selection favors altruism when rB>C (Hamiltons Rule)
- Kin selection: natural selection that favors altruism by enhancing the reproductive
success of relatives; weakens with hereditary distance
Reciprocal Altruism
- An exchange of aid (where the aided individual returns the favor in the future)
commonly invoke to explain altruism that occurs between unrelated humans
- Tit-for-tat mechanism
Sociobiology: certain behavioral characteristics exist because they are expressions of
genes that have been perpetuated by natural selection

Chapter 52: Ecology

Ecology: the scientific study of the interactions between organisms and the environment

Global Ecology: examines how the regional exchanges of energy and materials
influences the functioning and distribution of organisms across the biosphere
o Biosphere: the global ecosystemthe sum of all the planets ecosystems and
- Landscape Ecology: focuses on the factors controlling exchanges of energy,
materials, and organisms across multiple ecosystems
o Landscape: mosaic of connected ecosystems
- Ecosystem ecology: emphasizes energy flow and chemical cycling between
organisms and the environment
o Ecosystem: the community of organisms in an area and the physical factors
with which those organisms interact
- Community Ecology: examines how species interactions, such as predation and
competitions, affect community structure and organization
o Community: a group of populations of different species in an area
- Population Ecology: analyzes factors that affect population size and how and why it
changes through time
o Population: a group of individuals of the same species living in an area
- Organismal Ecology: concerned with how an organisms structure, physiology, and
behavior meet the challenges posed by its environment
The most significant influence on the distribution of organisms on land and in the oceans is
climate: the long-term prevailing weather conditions in a given area
- Four physical factors: temperature, precipitation, sunlight, and wind
- Macroclimate: patterns on the global, regional, and landscape level
- Microclimate: very fine, localized patterns
Global Climate Patterns
- Are determined largely by the input of solar energy and Earths movement in space
- Latitudinal Variation in Sunlight Intensity
o More heat and light per unit of surface area at tropics compared to higher
latitudes where sunlight strikes Earth at an oblique angle
- Global Air Circulation and Precipitation Patterns
o Warm, wet air masses rise in tropics and move towards poles
o As they cool, the ascending moist air releases moisture and create abundant
precipitation in tropical regions
o The now descending dry air moves across land and absorbs moisture, creating
arid climates
Regional and Local Effects on Climate
- Can be modified by factors such as large bodies of water and mountain ranges
- Seasonality:
o Earths tilted axis + rotation around sun cause strong seasonal cycles in
middle to high latitudes
o Changing angle of sun moves belts of wet and dry air slightly north and
southward, producing dry and wet seasons
o Changes in wind patterns alter ocean currents, causing upwelling of cold
water from deep ocean layers
Nutrient-rich water stimulates growth of surface-dwelling
- Bodies of Water
o Ocean currents influence climate along coasts of continents by heating or
cooling overlying air masses that pass across land
o Oceans + large lakes can moderate climate

Air over land heats up and rises, drawing a cool breeze from water
across land
At night, air over the now warmer water rises, drawing cooler air from
land back out over the water and replacing it w/warmer water offshore
o Mediterranean climate: cool, dry ocean breezes in summer are warmed when
they contact the land, absorbing moisture and creating a hot, arid climate just
a few km inland
o Rain-shadow effect:
Cool air flows inland from the water, moderating temps near the shore
Air that encounters mountains flows upwards, cools at higher altitudes,
and releases water as precipitation
Less moisture is left in the air reaching the leeward side, which
therefore has little precipitation. This rain-shadow can create a desert
on the back side of the mountain range
o Effect amount of sunlight reaching an area and thus the local temps and
rainfall (e.g. south sides of mountains are drier and warmer)
o Every 1,000-m increase in elevation = avg. temp drop of 6 degrees Celsius

- Every environment on Earth is characterized by a mosaic of small-scale differences in
chemical and physical attributes
- Abiotic Factors: nonliving factors that influence the distribution and abundance of
organisms (e.g. temperature, light, water, and nutrients)
- Biotic Factors: the living factors (the other organisms that are part of the
Global Climate Change
- Northern expansion of tree distributions due to warming of climate
Biomes: major life zones characterized by vegetation type in terrestrial biomes or by the
physical environment in aquatic biomes
Climograph: a plot of the annual mean temperature and precipitation in a particular region
Terrestrial Biomes
- Most biomes are named for major physical or climatic features and for their
predominant vegetation
- Usually grade into neighborhood biomes
o Ecotone: area of intergradation
- Vertical layering of vegetation is an important feature
o Canopy, low-tree layer, shrub understory, ground layer of herbaceous plants,
forest floor (litter layer), and the root layer
- Disturbance: event such as a storm, fire, or human activity that changes a
community, removing organisms from it and altering resource availability
Tropical Rainforest
Temperate Grassland
Northern Coniferous Forest (taiga)
Temperate Broadleaf Forest

Aquatic Biomes
- Characterized primarily by their physical environment
- Show far less latitudinal variation
- Salt concentration >3% = marine; salt concentration <0.1% = freshwater
- Water evaporated from the oceans provide most of the planets rainfall
- Freshwater biomes are closely linked to the soils and biotic components of the
surrounding terrestrial biome
- Many aquatic biomes are physically and chemically stratified (layered), vertically and
- Light intensity decreases rapidly with depth
- Pelagic zone
o Photic zone: where there is sufficient light for photosynthesis
o Aphotic zone: where little light penetrates
Abyssal zone: part of the ocean 2,000 6,000 m below the surface
- Benthic zone: at the bottom of all these aquatic zones
o Made up of sand and inorganic sediment
o Occupied by communities of organisms called benthos
Major food source for benthic species is detritus: dead organic matter,
which rains down from the productive surface waters of the photic
- Neritic zone: surface waters of the coastal zone
- Thermocline: a narrow later of abrupt temperature change that separates the more
uniformly warm upper layer from more uniformly cold deeper waters
o Lakes tend to be particularly layered w/respect to temperature esp. summer +
o Turnover: semiannual mixing of the water as a result of changing temperature
Sends oxygenated water from a lakes surface to the bottom and
brings nutrient-rich water from the bottom to the surface in spring +
Streams and Rivers
Intertidal Zones
Ocean Pelagic Zone
Coral Reefs
Marine benthic zone: deep-sea hydrothermal vents
Species distribution: a consequence of both ecological and evolutionary interactions
through time
- Ecological time: minute-to-minute time frame of interactions between organisms and
the environment
o The differential survival and reproduction of individuals that lead to evolution
occur in this time
- Evolutionary time: through natural selection, organisms adapt to their environment
over the time frame of many generations
- Dispersal: the movement of individuals or gametes away from their area of origin or
from centers of high population density
o Can lead to adaptive radiation: the rapid evolution of an ancestral species into
new species that fill many ecological niches

Species transplant and potential vs. actual range

Habitat selection behavior
Biotic Factors: negative interactions with predators or herbivores restrict the ability of
a species to survive and reproduce (Limpet, urchin, and seaweed study)
Abiotic Factors:
o Temperature
o Water and oxygen
o Salinity
o Sunlight
o Rocks and soil

Chapter 53
Population: a group of individuals of a single species living in the same general area
- Density: the number of individuals per unit area of volume; not a static property
o Immigration: influx of new individuals from other areas
o Emigration: movement of individuals out of a population and into other
- Dispersion: pattern of spacing among individuals within the boundaries of the
- Mark-recapture method: what ecologists use to estimate the size of wild-life
Patterns of Dispersion:
- Clumped: most common pattern in which individuals are aggregated in patches
- Uniform: evenly spaced pattern of dispersion that may result from direct interactions
between individuals in the population
o Territoriality: the defense of a bounded physical space against encroachment
by other individuals
- Random: position of each individual in a population is independent of other

Demography: the study of vital statistics of populations and how they change over
o Birth rates and death rates
Life tables: age-specific summaries of the survival pattern of a population; follow fate
o Cohort: a group of individuals of the same age, from birth until all of the
individuals have died

Survivorship Curves:
- A plot of the proportion or numbers in a cohort still alive at each stage; graph method
of representing data in a life table
- Type I Curve: flat at the start, reflecting low death rates during early and middle life,
and then drops steeply as death rates increase among older age groups
- Type III Curve: drops sharply at the start, reflecting very high death rates for the
young, but flattens out as death rates decline for those few individuals that survive
the early period of die-off
- Type II curve: intermediate, with a constant death rate over the organisms life span
Reproductive Rates:
- How reproduction output varies with the number of females and their ages
- Reproductive table: age-specific summary of the reproductive rates in a population
o Constructed by measuring the reproductive output of a cohort from birth until
Population Growth:
- Change in population size = births + immigrants deaths emigrants
o Per capita birth/death rate; per capita rate of increase
- Zero population growth: when the per capita birth and death rates are equal
- Exponential population growth: occurs when r is greater than zero and is constant at
each instant in time (J shape)
- Carrying capacity (K): the maximum population size that a particular environment
can sustain
o Crowding + resource limitation
- Logistic Population growth: the per capita rate of increase approaches zero as the
population size nears the carrying capacity
o Based on assumption that regardless of population density, each individual
added to a population has the same negative effect on population growth rate
o Allee effect: individuals may have a more difficult time surviving or
reproducing if population size is too small
Life History: evolutionary outcomes reflected in its development, physiology, and behavior
- Three variables: when reproduction begins, how often organism reproduces, and how
may offspring are produced per reproductive episode
- Semelparity: big-bang reproduction (salmon)
- Iteroparity: repeated reproduction
o Factors: survival rate of offspring and likelihood that the adult will survive to
reproduce again
o Trade Off
- K-selection: density-dependent selection; selection for traits that are sensitive to
population density and are favored at high densities
- R-selection: density-independent selection; selections for traits that maximize
reproductive success in uncrowded environments
Population Change:

Density independent: birth rate or death rate that does not change with population
density (death rate)
Density dependent: death rate that increases with population density or a birth rate
that falls with rising density (birth rate)
o Provides feedback regulation
o Mechanisms: competition for resources; disease; predation; intrinsic factors;
territoriality; toxic wastes
Population dynamics: population fluctuations from year to year or place to place
Population cycles: many undergo regular boom-and-bust cycles
Metapopulation: a linked number of local populations when immigration and
emigration are particularly important
Demographic transition: movement from high birth and death rates toward low birth
and death rates, which tends to accompany industrialization and improved living
Age structure: relative number of individuals of each age in the population
o Can predict a populations growth trends and illuminate social conditions
Infant mortality: number of infant deaths per 1,000 live births
o Life expectancy at birth: predicted average length of life at birth
Ecological footprint: summarizes the aggregate land and water area required by each
person, city, or nation to produce all of the resources it consumes and to absorb all of
the waste it generates

Chapter 54
Community: a group of populations of different species living close enough to interact
- Interspecific interactions: interactions with individuals of other species in the
community: competition, predation, herbivory, symbiosis (parasitism, mutualism, and
commensalism), and facilitation
- Interspecific competition: a -/- interaction that occurs when individuals of different
species compete for a resource that limits their growth and survival
- Competitive exclusion: two species competing for the same limiting resources cannot
coexist permanently in the same space; inferior competitor will be eliminated
Ecological niches
- Ecological niche: the sum of a species use of the biotic and abiotic resources in its
- Resource partitioning: the differentiation of niches that enables certain species to
coexist in a community
- Fundamental niche: the niche potentially occupied by a species
- Realized niche: the portion of its fundamental niche that the species actually
occupies due to competition
- Character displacement: tendency for characteristics to diverge more in sympatric
and in allopatric populations of two species
- Refers to a +/- interaction between species in which one species, the predator, kills
and eats the other, the prey
- Aposematic coloration: warning coloration causing predators to avoid prey
- Cryptic coloration: camouflage that makes prey difficult to see
- Batesian mimicry: a palatable or harmless species mimics and unpalatable or harmful

Mullerian mimicry: two or more unpalatable species resemble each other

Herbivory: refers to a +/- interaction in which an organism eats part of a plant or alga
- Plants have evolved to develop mechanisms to counteract predation
Symbiosis: when individuals of two or more species live in direct and intimate contact with
one another
- Parasitism: a +/- symbiotic interaction in which one organism, the parasite, derives
its nourishment from another organism, its host, which is harmed in the process
o Ectoparasites and endoparasites
- Mutualism: interspecific interaction that benefits both species (+/+)
o Obligate mutualism: at least one species has lost the ability to survive on its
o Facultative mutualism: both species can survive alone
- Commensalism: interaction that benefits one species but neither harms nor helps the
other (+/0)
Facilitation: species can have positive effects (+/+ or 0/+) on the survival and reproduction
of other species without necessarily living in the direct and intimate contact of a symbiosis
- Particularly common in plant ecology
Species Diversity
- The variety of different kinds of organisms that make up the community; has two
o Species richness: the number of different species in the community
o Relative abundance: the proportion each species represents of all individuals
in the community
o Shannon diversity
Diversity and Community Stability
- Higher-diversity communities generally are more productive and are better able to
withstand and recover from environmental stresses
- Biomass: the total mass of all organisms in a habitat
- Invasive species: organisms that become established outside their native range
Trophic Structure: the feeding relationships between organisms of the community
- Food chain: the transfer of food energy up the trophic levels from its source in plants
and other autotrophs through herbivores to carnivores and eventually to
o Energetic hypothesis: idea that suggests the length of a food chain is limited
by the inefficiency of energy transfer along the chain
- Food webs: linked food chains
- Dominant species: the species that are the most abundant or that collectively have
the highest biomass
- Keystone species: not usually abundant in a community, but they exert strong control
on community structure not by numerical might but by their pivotal ecological roles
or niches
- Ecosystem engineers/foundation species: organisms that exert their influence on a
community not through trophic interactions but by changing their physical
Bottom-Up and Top-Down Controls

Bottom-up model: postulates a unidirectional influence from lower to higher trophic

o Mineral nutrients controls plants numbers, which control herbivore numbers,
which control predator numbers
o To change structure, you need to alter biomass at the lower trophic levels
Top-down model: postulates that predation mainly controls community organization
because predators limit herbivores, herbivores limit plants, and plants limit nutrient
levels through nutrient uptake
o Also called trophic cascade model
o Effects move down as alternation (+/-) effects
Biomanipulation: application of the top=down model to improve water quality in
polluted lakes

- Stability: a communitys tendency to reach and maintain a relatively constant
composition of species
- Climax community: stable community of plants controlled solely by climate
- Disturbance: an event that changes a community by removing organisms from it or
altering resource availability
- Nonequilibrium model: describes most communities as constantly changing after
- Intermediate disturbance hypothesis: moderate levels of disturbance foster greater
species diversity than do high or low levels of disturbance
Ecological Succession
- Primary (begins in a lifeless area where soil has not yet formed) vs. secondary
succession (existing community has been cleared by some disturbance that leaves
soil intact)
- Linkage of early-arriving and late-arriving species
o Facilitate appearance of later species by making environment more favorable
o Inhibit establishment of later species
o Early species completely independent of later species, which tolerate
conditions already created
- Stages:
o Pioneer stage, dryas stage, alder stage, spruce stage
Latitudinal Gradients: plant and animal life appear to be more abundant and diverse in the
tropics than in any other parts of the globe
- Evapotranspiration: evaporation of water from soil and plants
o Potential evapotranspiration: a measure of potential water loss that assumes
that water is readily available
Area Effects
- Species-area curve: the larger the geographic community, the more species it has, in
part because larger areas offer a greater diversity of habitats and microhabitats
Island Equilibrium Model
- Identified key determinants of species diversity on an island given set of physical
- Factors that determine number of species:
o Rate at which new species immigrate and rate at which species become
- Two physical features affecting immigration + extinction rates:
o Size + distance from mainland

Equilibrium will eventually be reached where the rate of species immigration equals
the rate of species extinction

Pathogens: disease-causing microorganisms

- Zoonotic pathogens: those that are transferred to humans from other animals
o Vector: intermediate species carrying pathogens

Chapter 54
Ecosystem: the sum of all the organisms living in a given area and the abiotic factors with
which they interact
Conservation of Energy:
- Energy cannot be created or destroyed but only transferred or transformed
- Law of conservation of mass: matter cannot be created or destroyed
Energy, Mass, and Trophic Levels
- Species are grouped into trophic levels based on their main source of nutrition and
- Primary producers: photosynthetic organisms that use light energy to synthesize
sugars and other organic compounds, which they use as fuel for cellular respiration
and as building material for growth
o Supports all other trophic levels
- Consumers: primary, secondary, tertiary
- Detritivores: decomposers/ consumers that get their energy from detritus
- Detritus: nonliving organic material; help recycle organic material back into nonorganic material
Primary production: the amount of light energy converted to chemical energyin the form of
organic compoundsby autotrophs during a given time period.
- 1% of visible light that strikes photosynthetic organisms is converted to chemical
Gross and Net Production
- Gross Primary Production: the amount of energy from light converted to the chemical
energy or organic molecules per unit time
o Not all is stored as organic material in primary producers
- Net primary production: gross primary production minus energy used by primary
consumers for their autotrophic respiration
o Usually GPP
o Most valuable b/c represents the storage of chemical energy available to
consumers in the ecosystem
o J/(m^2 x yr);
o Amount of new biomass added in a given period of time
- Standing crop: the total biomass of photosynthetic autotrophs present
- Net ecosystem production: measure of the total biomass accumulated during a given
period of time
o GPP total respiration of all organisms in the system
o Value determines whether an ecosystem is gaining or losing carbon over time
o Measure flux of carbon dioxide and oxygen entering or leaving the ecosystem
Primary Production in Aquatic Ecosystems:

Light Limitation: key variable in controlling primary productivity

Nutrient Limitation: limit primary production in most oceans and lakes more than light
- Limiting nutrient: element that must be added for production to increase
o Nitrogen or phosphorus
- Upwelling: deep, nutrient rich waters circulate to the ocean surface
- Eutrophication: when primary producers die and detritivores decompose them,
depleting the water of much or all of its oxygen loss of many fish species
o Phosphorus responsible!?
Primary Production in Terrestrial Ecosystems
- Temperature and moisture are main factors controlling primary production
- Mean annual precipitation and evapotranspiration (total amount of water transpired
by plants and evaporated from a landscape) are useful second predictors
o Evapotranspiration increases w/ temp and amount of solar energy
Nutrient Limitations and Adaptations that Reduce Them
- Soil nutrients limit primary productivity
- Nitrogen limits plant growth the most
- Symbiosis of mycorrhizae (fungi) and plant roots
Secondary Production: the amount of chemical energy in consumers food that is converted
to their own new biomass during a given period
- Production efficiency: net secondary production (energy stored in biomass
represented by total growth and reproduction)/assimilation of primary production
(total energy taken in)
Trophic Efficiency: percentage of production transferred from one trophic level to the next
- Always less than production efficiencies
- 90% of energy available is not transferred to the next
- Limits abundance of top-level carnivores and food chains
- Pyramid of net production: trophic levels are arranged in tiers
- Biomass pyramid: each tier represents the standing crop in one trophic level
o Some aquatic ecosystems have inverted biomass pyramids
Occur b/c producers grow, reproduce, and are consumed so quickly by
zooplankton that they never develop a large population size
Phytoplankton have short turnover time:
Standing crop/production
Biological and geochemical cycles
- Global (gases) vs. local (nutrients)
- Scientists use isotopes to determine cycle
- Water cycle, Carbon cycle, Phosphorus cycle, Nitrogen cycle
- Controlled by the same factors that limit primary productivity: temperature, moisture,
and nutrient availability
Biological Restoration
Bioremediation: using organismsusually prokaryotes, fungi, or plants to detoxify polluted
- Removes harmful substances from an ecosystem
Biological augmentation: uses organisms to add essential materials to a degraded

Chapter 56
Conservation biology: integrates ecology, physiology, molecular biology, genetics, and
evolutionary biology to conserve biological diversity at all levels
Three Levels of Biodiversity:
- Genetic diversity: comprises not only the individual genetic variation w.in a
population, but also the genetic variation between populations that is often associate
with adaptations to local conditions
- Species Diversity: the variety of species in an ecosystem or across the biosphere
o Endangered species: one that is in danger of extinction throughout all or a
significant portion of its range
o Threatened species: those considered likely to become endangered in the
near future
- Ecosystem Diversity
Ecosystem Services: encompass all the processes through which natural ecosystems help
sustain human life
- Ecosystems purify air and water
- Detoxify and decompose wastes and reduce impacts of extreme weather and
Threats to Biodiversity:
- Habitat Loss
- Introduced Species: species that humans move intentionally or accidentally from the
species native locations to new geographic regions
- Overharvesting
- Global Change: acid precipitation
Population Conservation
- Small Population Approach
o Extinction vortex: due to inbreeding and genetic drift, the population becomes
smaller and smaller until no individuals survive
Loss of genetic variation
o Minimum Viable Population: the minimal population size at which a species is
able to sustain its numbers
o Effective population size: based on the breeding potential of the population
- Declining Population Approach: focuses on threatened and endangered populations
that show a downward trend, even if the population is far above its minimum viable
Edges: boundaries between ecosystems
- Movement corridor: a narrow strip or series of small clumps of habitat connecting
otherwise isolated patches, can be extremely important for conserving biodiversity
- Biodiversity hotspot: relatively small area with numerous endemic species (species
found nowhere else in the world) and a large number of endangered and threatened
- Zoned reserve: extensive region that includes areas relatively undisturbed by
humans surrounded by areas that have been changed by human activity and are
used for economic gain
- Urban ecology: examines organisms and their environment in urban settings
- Critical load: the amount of added nutrient that can be absorbed by plants without
damaging ecosystem integrity
- Biological magnification: when toxins become more concentrated in successive
trophic levels

Greenhouse effect
Assisted migration: the translocation of a species to a favorable habitat beyond its
native range to protect the species from human-caused threats
Sustainable development: economic