BIOB50 - 2023 - Lecture 3

BIOB50H3F
Ecology
Fall 2023, Week 3

Learning Goals, Lecture 3
- Understand limiting factors and how they determine an organism’s niche.
- Understand the role of energy as a limiting factor, including the usage of energy by
organisms (Principle of Allocation), the ways organisms acquire energy (optimal foraging
theory), and an example using these principles to understand how climate change impacts
polar bears
- Understands the role of temperature as a limiting factor, including how temperature affects
physiological and demographic rates (thermal performance curves) and an example using
these principles to understand how climate change impacts disease spread
LIMITING FACTORS
&
AN ORGANISM’S NICHE
Limiting Factors & An Organism’s Niche
Limiting Factors, Example: Biomes
A limiting factor is a resource or environmental

condition that limits the growth, distribution or
abundance of an organism or population within
an ecosystem.
Example: In Lecture 2, we have seen how the limiting

factors temperature and precipitation determine what
plant groups will establish in an area.
The Niche
A species’ tolerances to its limiting

factors define its niche, which can
broadly be understood as the set
of environmental conditions in
which an organism can live and
Limiting Factor 2
reproduce.
Formally, a (Hutchinsonian) niche

is defined as an n-dimensional
hypervolume, where n indicates
the number of different
environmental conditions that
determine where an organism can
survive and reproduce.
Limiting Factor 1
Example: The Niche of Devil’s Hole Pupfish
The Niche: Performance Curves
An organism’s performance, often quantified as the rate of

a given physiological process (e.g., rates of movement,
reproduction, mortality) varies along the axes of its niche
space. It is low near the extremes of a limiting factor and
optimal at some intermediate value. Performance curves
describe this variation.
Limiting Factor 2
Stress Stress
Limiting Factor 1
Measuring the Niche
Variation in the preferences of a species Variation in physiological performance due

along one or more axes of their niche space variation in limiting factors can often be measured
can be measured by studying their experimentally (e.g., rearing individuals in thermal
behavior. incubators for studying temperature effects)
Fundamental Niche vs Realized Niche
In nature, the realized niche of a species (the set of

environmental conditions where the species occurs) is typically
smaller than its fundamental niche (the set of environmental
conditions that a species could tolerate), as species are typically Stress Stress
excluded from the extremes of their niche by better adapted,
superior competitors and other factors.
Example: the barnacle Semibalanus balanoides only occurs

in the northern part of its potential range; it is excluded from
the southern part by other barnacle species that are superior
competitors
ENERGY,
A KEY LIMITING FACTOR
Energy, a Key Limiting Factor All metabolic processes require energy,
Energy, the Currency of Life including for the maintenance of cells, the
production of enzymes, growth, movement,
reproduction & thermoregulation. Without energy,
enzymes fail, cell membranes degrade, organelles
cease to operate. Life is impossible.
Heterotrophs obtain energy by consuming organic

compounds from other organisms
Autotrophs convert the energy of sunlight or

inorganic compounds into chemical energy stored in
the carbon-carbon bonds of organic compounds.
ENERGY,
Using Energy:
The Principle of Allocation
Energy, A Key Limiting Factor
The Principle of Allocation
To understand the acquisition and use of energy by an organism, it is important to consider its energy budget, as
defined through the Principle of Allocation: Individuals must prioritize how to use the limited energy that is
available to them. As a results, energy allocated to one of life’s necessary physiological functions will reduce the
amount of energy that can be allocated to other such functions. Given that energy cannot be created or
destroyed, an individual’s energy budget can be described by:
Eintake = Erespiration + Eassimilation + Ereproduction + Ewaste
total energy that total energy devoted total energy converted to total energy used for total energy lost
an individual to getting oxygen living tissue in the reproduction in waste or heat
ingests (in plants: into the body and organism, either to grow or inefficiency
total energy cells for metabolism its structure (tissues that (in plants: total
captured by cannot be broken down energy lost as
photosynthesis for energy; e.g. organs) or dead plant
its storage (tissues that tissue)
can be tapped into later
when the individual’s
energy intake does not
meet its energy needs)
ENERGY,
Acquiring Energy:
Optimal Foraging Theory
Energy, a Key Limiting Factor
Animals can adjust their energy budgets by adjusting their behavior. Applying the Principle of
Allocation to animal foraging allows answering questions, such as ‘How much time and energy
should an individual expend foraging?’, ‘What search strategy should it employ?’, ‘What
types of food should it choose?’
Optimal Foraging Theory aims to predict the best strategies for maximizing energy intake by
considering the energetic profitability of a food item
P = E / C, where E = energy gained from a food item, and

C = energy costs associated with acquiring and eating the food.
Higher values of P indicate more profitable prey items (i.e., high energetic value, low acquisition
costs, or both); P = 1 is a breakeven point where energy gained equals the energy invested; and
P < 1 indicates food items where the costs of acquiring them outweighs the energy gain.
Energy, a Key Limiting Factor
The simple energetic profitability model, P = E / C, is typically extended to separate the costs of a food
item into its search costs (dependent on resource density and search strategy) and its handling
costs (the combined the time it takes to capture, manipulate, consume, and digest the food):
P = E / (S + H), where S = energy costs associated with searching for the food,
H = energy costs associated with handling the food.
P is maximized when energy

intake is large, relative to
search and handling costs.
Title
Subtitle
Optimal Foraging Theory makes quantifiable predictions that can be tested experimentally, for example, (1)
foragers should prefer resources that have higher profitability, and (2) as the abundance of higher-value
resources increases, consumption of lower-value resources should decrease.
Percentage large
prey selected
Ratio of profitability of large prey to

profitability of small prey
ENERGY,
Application: Understanding Climate Change

Impacts on Polar Bears
Application: Understanding Climate Change Impacts on Polar Bears (Energy as a Limiting Factor)
Can we use our understanding of energy budgets

to predict how climate change may impact the
reproduction and survival of polar bears?
Eintake =
Erespiration + Eassimilation + Ereproduction + Ewaste
Molnár et al. 2020. Nature Climate Change 10: 732-738 https://www.nytimes.com/2020/07/20/climate/polar-bear-extinction.html

Polar Bears Need Sea Ice
Polar bears cannot catch seals in open water. They need sea ice to hunt them.
Declining Sea Ice Results in a Cascade of Demographic Impacts
climate change
reduced sea ice
less food, longer fasting
reduced energy stores fewer & lighter cubs
increased cub mortality
higher adult & population declines

subadult mortality
Approach: Energy Budget Modelling
Energy budget models can be used to estimate how an individual partitions its energy among
physiological processes.
Respiration
Storage Cubs Reproduction
Assimilation: Storage
Intake
Waste
Production
Assimilation: Structure
When energy intake is plentiful, resources can be allocated to all processes, including growth and reproduction.
Approach: Energy Budget Modelling
Energy budget models can be used to estimate how an individual partitions its energy among
physiological processes, as well as how long it can survive on its energy stores without feeding.
Respiration
Storage Cubs Reproduction
Assimilation: Storage
Intake
Waste
Production
Assimilation: Structure
When energy intake is insufficient or absent, energy needs can be met from storage. Growth and/or reproduction may
cease to save energy. When the storage is depleted, the individual cannot meet its needs anymore and dies. Comparing
the amount of energy that an individual has stored against its necessary daily respiration and waste costs allows
estimating how long the individual can survive before it starves to death.
How Long Will Polar Bears Have to Fast?
Green: Seasonal Ice Ecoregion (SIE)

Red: Divergent Ice Ecoregion (DIE)
Blue: Convergent Ice Ecoregion (CIE)
Yellow: Archipelago Ecoregion (AE)
The length of the summer ice-free season, and thus, how long polar bears have to survive without food, varies
among the 19 recognized polar bear populations. In some populations, the sea ice is perennial, guaranteeing
access to food year-round (CIE); in others, the sea ice melts each summer, stranding bears on land (SIE / DIE),
or on ice shelves over the deep, unproductive polar basin (DIE), where they have to survive for months without
food.
Example: increasingly long ice-free seasons in the

Western Hudson Bay subpopulation
Sea ice extent in Western Hudson Bay subpopulation (SIE)

(blue, 1980s; green, 1990s; yellow, 2000s; red 2010s
Longer ice-free seasons →

Less time spent on the sea ice to feed on seals (→ less energy reserves when fast begins)
AND
More time spent on land fasting (→ more energy reserves are needed to survive the fast)
January December
Warmer climate
Feeding On-shore fast

On-shore fast Feeding
Feeding On-shore fast Feeding
ice melts freeze-up

ice melts freeze-up
Length of fasting period [days]
Past fast durations:

Red line, estimated from passive microwave
satellite imagery
Projected fast durations (based on Community

Earth System Model version 1):
- RCP4.5 scenario (emissions peak around

2040 and then decline): grey line
- RCP8.5 scenario (emissions continue to rise
through 21st century): black line
How Long Are Polar Bears Able to Fast?
How long polar bears are able to fast depends on how fat they are at the beginning of the fast, and on
how much energy they require every day.
Survival: maybe
Reproduction: no
Survival: almost certain

Reproduction: almost certain
How Long Are Polar Bears Able to Fast?
How long polar bears are able to fast depends on how fat they are at the beginning of the fast, and on
how much energy they require every day .
Example: An adult male polar bear that weighs 476.7 kg and has a body length of 2.34 m,
could theoretically fast for 296 days before death by starvation occurs. This is calculated
using an energy budget model that compares the amount of energy that an
individual has stored against its necessary daily respiration and waste costs.
How Long are Polar Bears Able to Fast?
The energy budget model suggest that surpassing the thresholds for the number of ice-free days outlined below
is expected to result in steep declines in adult survival and the successful rearing of offspring.
Note: cub survival is impacted first, then yearling survival, followed by adult male survival, the survival of mother bears,
and the survival of solitary females
Looking backwards in time. Do the models match reality?
Fast duration in relation to impact

thresholds for cub recruitment &
adult male survival
1980 1990 2000 2010
- Declines in cub recruitment started in late 1990s/early
2000s, as projected by the model (intersection of fasting
period length, black line, with estimated threshold for
recruitment impacts, magenta line); the survival rates of
adults remained unaffected until at least the late 2000s,
also as projected by the model (no intersection between
black line and threshold for impacts on adult male
Stirling & Derocher 2012, Global Change Biology
survival, blue line)
4. Modelbackwards
Looking tests: hindcasting
in time. demographic
Do the modelsimpacts
match reality?
Seasonal Ice Ecoregion (SIE)
SH: “Females sacrificing body condition DS: “Cub recruitment lowest of all SIE
to maintain lactation, high survival” subpopulations, adult survival high”
(Obbard et al. 2018) (Peacock et al. 2013)
Fast duration in relation to impact
thresholds for cub recruitment &
adult male survival
1980 1990 2000 2010
1980 1990 2000 2010 1980 1990 2000 2010

FB “High reproduction & BB: “Decreasing offspring
survival, stable subpopulation” recruitment since mid-1990s; high
(Stapleton et al. 2016) adult survival” (Laidre et al. 2020)
Looking forwards in time. When do we expect dramatic declines in cub
recruitment and/or adult survival in each subpopulation?
Bears are affected when an impact

threshold is crossed if the
population’s body mass distribution
falls below the corresponding value.
This leads to a double whammy:
Longer fasts require better body
condition, but becoming fat is more
difficult with longer fasts.
Projections under Two Climate Scenarios: The Impacts of Sea Ice Loss and Resultant Prolonged Fasting
Combining projections for sea ice loss with the threshold estimates for how long polar bears can fast shows when we should expect
to see substantial declines in cub recruitment (pink), adult male survival (blue), and/or adult female survival (red) to occur. The
RCP8.5 scenario (rising emissions through 21st century) leads to the likely extirpation of polar bears almost everywhere.
Projections under Two Climate Scenarios: The Impacts of Sea Ice Loss and Resultant Prolonged Fasting
Combining projections for sea ice loss with the threshold estimates for how long polar bears can fast shows when we should expect
to see substantial declines in cub recruitment (pink), adult male survival (blue), and/or adult female survival (red) to occur. The
RCP8.5 scenario (rising emissions through 21st century) leads to the likely extirpation of polar bears almost everywhere.
The RCP4.5 scenario (emissions rise till 2040, then decline) still leads to the likely extirpation of the southernmost
subpopulations, but allows persistence elsewhere (albeit with reduced cub recruitment) to the end of the century.
TEMPERATURE,
Temperature, a Key Limiting Factor
Thermoregulation: Poikilotherms, Ectotherms, Endotherms
Organisms can be grouped by the degree to which they control their internal temperature.
Poikilotherms Ectotherms use external Endotherms use behavioral Some endotherms, called
do not regulate factors (e.g., sun, shade) thermoregulation, plus internal homeotherms, are able
their internal to regulate their internal processes (e.g., metabolic heat, to maintain their internal
temperature. temperature. shivering) to regulate their temperature in a very
internal temperature. narrow range.
Thermal Performance Curves
Temperature controls the rates of all metabolic reactions, and thus, the rates of all physiological and behavioral
processes (e.g., rates of movement, growth, reproduction) It is a key factor for determining the performance of
organisms.
Lower lethal limit: cellular Upper lethal limit: cellular

respiration slows and may cease, structures and DNA break down,
leading to death. leading to death
Example: Thermal Performance of Lace Monitor Lizards
Example: Lace monitor lizards regulate their internal body temperature throughout the day through a series of
behavioral choices (e.g., seeking out shade vs basking in the sun).
Example: Thermal Performance of Lace Monitor Lizards
Lecture 1
Example: The swimming speed of the cercariae of

Schistosoma mansoni (the parasite causing
schistosomiasis)
from Molnár et al. 2017, Journal of Parasitology.
TEMPERATURE,
Application: Understanding Climate Change

Impacts on Disease Spread
Application: Understanding Climate Change Impacts on Disease Spread (Temperature as a Limiting Factor)
Can we use our understanding of a parasite’s

niche and its thermal performance to predict how
climate change may shift the geographic range of
diseases?
Observed Climate Change Impacts on Disease Spread
Example: Malaria in the East African highlands
Due to generally cool temperatures, the East

African highlands used to be malaria-free. A
temperature increase in the region now allows the
malaria parasite to complete its life cycle, and the
disease established.
https://www.scientificamerican.com/article/east-africa-malaria-rises-under-climate-change/
500 km
2008
Example: Umingmakstrongylus pallikuukensis is a lungworm of muskoxen. Adult parasites form cysts
in the lungs. Larvae are expelled into the environment via muskox feces, where they need to find a
Kugluktuk taken up by grazing
slug to develop to the infective larval stage. Infective larvae are accidentally
muskoxen. Larval development rates, mortality rates, and infection rates all depend on temperature.
Before 2008, U. pallikuukensis

was restricted to the Canadian
mainland but a warmer climate
allowed its expansion and 500 km
establishment to muskox herds
on Victoria Island.
2015 2014
Hadley Bay
Ulukhaktok
2012
2008
Kugluktuk
Think – Pair – Share
In your future job as
a disease ecologist /
epidemiologist, you
are tasked with
determining the
likely future range of
malaria and various
other diseases for
different climate
change scenarios.
How would you
approach this
problem?
Approaches to Predicting the Impacts of Climate Change on Disease Spread
Two of the most commonly used strategies for predicting climate change impacts on
the range of species are climate envelope models and models based on thermal
performance curves.
A. Climate envelope models use statistical modelling to determine under what climate
conditions a species currently occurs. Subsequently, they consider climate projections to
figure out where those same conditions are likely to occur in the future to determine a likely
future range for the species.
B. Thermal performance based population models try to estimate where a species could
occur based on its thermal constraints. Combining thermal performance curves for
development rate, birth rate, death rate, and other demographic variables, can yield estimates
of population growth rates under different temperature regimes. Combined with climate
projections, this approach can yield estimates of where the species is likely to occur in the
future.
Approaches to Predicting the Impacts of Climate Change on Disease Spread: 1. Climate Envelope Models
Climate envelope models use statistical modelling to determine under what climate conditions a species
currently occurs. Subsequently, they consider climate projections to figure out where those same conditions
are likely to occur in the future to determine a likely future range for the species.
Step 1: record all geographic locations where the species is found

Step 2: record climate covariates at every point, and use statistical approaches to determine
under what conditions the species occurs
Step 3: use climate models to determine where on the map those same climatic conditions
will / will not occur in the future. The hypothesis is that this will roughly represent the new
distribution of the species.
Sample Short Answer Question
Given its importance as an intermediate host for schistosomes, your team is trying to predict
the future distribution of a certain freshwater snail species. After extensive field surveys, you
find that the snail occurs in areas of Africa where temperatures do not drop below 15C and do
not rise above 35C. After consulting the latest climate projections, you find that such
temperatures could become common in southern Europe in the near future, but not yet in
more northerly areas.
a) Give one reason why the snails might not become present in southern Europe despite your
climate envelope predictions.
b) In area A, temperatures could drop as low as 12C, but not lower.

Can you rule out, based on the data that you collected (and assuming
no evolutionary adaptation), that the snail will not establish in area A
under these conditions? Why (not)?
Approaches to Predicting the Impacts of Climate Change on Disease Spread: 2. Thermal Performance Based Population Models
Thermal performance based population models try to estimate where a species could occur based on its thermal constraints.
Combining thermal performance curves for development rate, birth rate, death rate, and other demographic variables, can yield
estimates of population growth rates under different temperature regimes. Combined with climate projections, this approach can
yield estimates of where the species is likely to occur in the future.
e.g. R0 for the muskox lungworm U.p. =
prob. that free-living larva finds snail before

dying *
prob. that larva in snail develops to infective

stage before dying *
prob. of getting ingested by a muskox and

developing to adult stage before dying *
fecundity of adult parasite
Step 1: Determine the parasite’s R0 (the average number of offspring produced by an

2008
individual over its lifetime; cf. Lectures 4, 5, and 8). It can be calculated from the probabilities of
reaching each life stage and the fecundities at that life stage.
Thermal performance based population models try to estimate where a species could occur based on its thermal constraints.
Combining thermal performance curves for development rate, birth rate, death rate, and other demographic variables, can yield
estimates of population growth rates under different temperature regimes. Combined with climate projections, this approach can
yield estimates of where the species is likely to occur in the future.
e.g. R0 for the muskox lungworm U.p. =
prob. that free-living larva finds snail before dying *
prob. that larva in snail develops to infective stage before dying *
prob. of getting ingested by a muskox and developing to adult stage before dying *
fecundity of adult parasite
Step 2: Estimate thermal performance curves for each of the terms in the equation defining R0.
This allows us to estimate how R0 varies as a function of temperature.2008
e.g., U. pallikuukensis
The colored lines mark how the

R0 = 1 boundary has shifted
northwards with a warming
climate over the past four
decades
Step 3: Combine climate maps/projections with our estimates of how R0 varies with temperature
to estimate how disease pressures are/will be varying across landscapes. Of particular interest is
2008(R0 > 1) from areas
hereby the R0 = 1 boundary, which separates areas with population growth
with population declines (R0 < 1).
Sample Short Answer Question
We observed muskoxen that tested positive for the Umingmakstrongylus pallikuukensis,
despite being north of our estimate for the R0 = 1 line. Does this mean that our calculations
have a mistake? Why (not)?
Example: The Fundamental Thermal Niche of Malaria & Its Shifting Geographic Range
Measuring thermal
performance curves of
mosquitoes and the malaria
parasites for various life
history traits around the life
Erin Mordecai,
cycle… Stanford U.
from Mordecai et al. 2013, Ecology Letters 16, 22-30.

… allowed estimating the

temperature-dependence of
the parasite’s basic
reproductive number R0, blue line: R0 as a
function of
which in turn… temperature Erin Mordecai,
(ignore red line) Stanford U.
from Mordecai et al. 2013, Ecology Letters 16, 22-30.

… allowed estimating how

the range of malaria may
change with climate
change.
from Ryan et al. 2015, Vector-Borne and Zoonotic Diseases, 16: 718-725.
The Generality of the Approach: Thermal Performance Predictions for Zika, Dengue Fever, West Nile Virus, et al.
Eastern equine encephalitis virus (EEEV), Western equine

encephalitis virus (WEEV), Sindbis virus (SINV), West Nile
virus (WNV), St. Louis encephalitis virus (SLEV), Rift Valley
fever virus (RVFV), dengue virus (DENV), Ross River virus
(RRV), Murray Valley encephalitis virus (MVEV), Zika virus
(ZIKV).
from Mordecai et al. 2019, Thermal biology of mosquito-borne disease, Ecology Letters.
ASSIGNED READINGS:
Chapter 5,
excluding 5.4 (Water Balance) and the
second half of 5.5 (Plant Thermoregulation)

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BIOB50H3F

Fall 2023, Week 3

- Understand limiting factors and how they determine an organism’s niche.

A limiting factor is a resource or environmental

Example: In Lecture 2, we have seen how the limiting

A species’ tolerances to its limiting

Formally, a (Hutchinsonian) niche

An organism’s performance, often quantified as the rate of

Variation in the preferences of a species Variation in physiological performance due

In nature, the realized niche of a species (the set of

Example: the barnacle Semibalanus balanoides only occurs

Heterotrophs obtain energy by consuming organic

Autotrophs convert the energy of sunlight or

Eintake = Erespiration + Eassimilation + Ereproduction + Ewaste

Eintake = Erespiration + Eassimilation + Ereproduction + Ewaste

P = E / C, where E = energy gained from a food item, and

P is maximized when energy

Ratio of profitability of large prey to

Application: Understanding Climate Change

Can we use our understanding of energy budgets

Molnár et al. 2020. Nature Climate Change 10: 732-738 https://www.nytimes.com/2020/07/20/climate/polar-bear-extinction.html

reduced sea ice

less food, longer fasting

reduced energy stores fewer & lighter cubs

increased cub mortality

higher adult & population declines

Eintake = Erespiration + Eassimilation + Ereproduction + Ewaste

Storage Cubs Reproduction

Eintake = Erespiration + Eassimilation + Ereproduction + Ewaste

Storage Cubs Reproduction

Green: Seasonal Ice Ecoregion (SIE)

Example: increasingly long ice-free seasons in the

Sea ice extent in Western Hudson Bay subpopulation (SIE)

Longer ice-free seasons →

Feeding On-shore fast

ice melts freeze-up

Past fast durations:

Projected fast durations (based on Community

- RCP4.5 scenario (emissions peak around

Survival: almost certain

Fast duration in relation to impact

1980 1990 2000 2010

1980 1990 2000 2010 1980 1990 2000 2010

Bears are affected when an impact

Lower lethal limit: cellular Upper lethal limit: cellular

Example: The swimming speed of the cercariae of

Application: Understanding Climate Change

Can we use our understanding of a parasite’s

Example: Malaria in the East African highlands

Due to generally cool temperatures, the East

Before 2008, U. pallikuukensis

Step 1: record all geographic locations where the species is found

b) In area A, temperatures could drop as low as 12C, but not lower.

e.g. R0 for the muskox lungworm U.p. =

prob. that free-living larva finds snail before

prob. that larva in snail develops to infective

prob. of getting ingested by a muskox and

fecundity of adult parasite

Step 1: Determine the parasite’s R0 (the average number of offspring produced by an

e.g. R0 for the muskox lungworm U.p. =

prob. that free-living larva finds snail before dying *

prob. that larva in snail develops to infective stage before dying *

fecundity of adult parasite