Lecture 9: Mutualism and symbiosis Lecture 9: Mutualism and symbiosis

• Lecture clip 1: • Lecture clip 1:

• Population dynamics of mutualism; invasional • Population dynamics of mutualism; invasional
meltdown meltdown
• Mutualism and community structure • Mutualism and community structure
• Lecture clip 2: • Lecture clip 2:
• How mutualists adapt to one another; Darwin’s • How mutualists adapt to one another; Darwin’s
orchid and coevolution orchid and coevolution
• Generalists vs. specialists; mutualistic networks • Generalists vs. specialists; mutualistic networks
• Microbiomes • Microbiomes

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• Symbiosis = “living together” Mutualisms typically involve

• Mutualism = beneficial reciprocal exchange of goods or
interaction for both species services between species
• Mutualism ≠ symbiosis
• Nutritional mutualisms:
• Legumes & rhizobia: exchange fixed C for fixed N
• Plants & mycorrhizal fungi: exchange C for P
• Defensive mutualisms:
• Ants & plants: exchange protection for food (e.g.,
extrafloral nectar) or housing
• Cleaner fish & client fish: exchange parasite removal
for food
• Dispersal mutualisms:
• Plants & animal seed dispersers: exchange seed
dispersal for food
• Plants & animal pollinators: exchange gamete
dispersal for food
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Mutualism between humans Think back to the Lotka-Volterra

and free-living wild animals model of inter-specific competition
• Yao people in Mozambique harvest wild honey, but
can’t find bees’ nests easily
• Honeyguides (Indicator indicator) eat bees wax and
How would you change these equations
know where nests are, but can’t access them easily Claire Spottiswoode to model mutualism?
Spottiswoode et al. (2016, Science)

A Yao honey-hunter and a wild, Honeyguides recognize the specific sound Remember that αij = per-capita effect on i by j
free-living honeyguide that Yao honey-hunters make to attract5them 6

© 2021 Prof. Megan Frederickson 1

Think back to the Lotka-Volterra
model of inter-specific competition Population dynamics of mutualism
How would you change these equations • “[Lotka-Volterra models of mutualism] lead to
to model mutualism? silly solutions in which both populations
undergo unbounded exponential growth, in
an orgy of mutual benefaction.”
May R. 1981. Theoretical Ecology
1 • What limits the population growth of
mutualists?
• Strong intra-specific competition
1 • A third species such as a predator or a competitor
• Diminishing returns to mutualism as the
population grows
Remember that αij = per-capita effect on i by j
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Spring ephemerals =
perennial understory
Invasional meltdown herbs that flower right
after snow melts,
• Positive feedback between mutualists tends producing a short-lived
to generate runaway population growth “carpet of flowers”
• What if two invasive species interact as
mutualists?
• Simberloff and Von Holle (1999) coined the
term “invasional meltown” for the process by
which two non-native species facilitate one
another’s spread

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With native ant, With invasive ant,

mostly native plants mostly invasive plants

Native seed-dispersing ant, Invasive seed-dispersing ant,

Aphaenogaster rudis Myrmica rubra

Experimental mesocosms
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© 2021 Prof. Megan Frederickson 2

 After 12 days

• Cleaner fish feed on

“Cleaner” fish (Labroides ectoparasites on the
dimidiatus) cleaning a ”client” fish bodies of client fish;
cleaners gain food,
clients benefit from
fewer parasites After 12 hours

Video: Vassia Atanassova \ Wikimedia Commons

• Cleaners often have
“cleaning stations”
(territories) that
clients visit
• Experimentally After 24 hours
removing cleaner
fish increases
parasite (gnathiid)
abundance on client
fish
13 Grutter (1999, Nature) 14

Presence of cleaner fish in reefs affects

species diversity of other reef fish Lecture 9: Mutualism and symbiosis
Cleaner gone New cleaner
Bshary (2003, J. Anim. Ecol.)

• Studied natural variation in
cleaner fish presence/absence
• Also added/removed cleaner fish
• Grey = short-term (2-4 weeks),
white = long-term (4-24 months)
• Lecture clip 1:
• Population dynamics of mutualism; invasional
meltdown
• Mutualism and community structure
• Lecture clip 2:
• How mutualists adapt to one another; Darwin’s
orchid and coevolution
• Generalists vs. specialists; mutualistic networks
• Microbiomes
Redouan Bshary 15 16

Left: Alfred Russel

Darwin’s orchid, Wallace’s 1867 drawing of
Angraecum sesquipedale a moth that might
pollinate A. sesquipedale.
• Epiphytic orchid Below: Xanthopan
from Madagascar morganii praedicta,
• Darwin (1862) discovered in 1903 (21
famously years after Darwin died)
predicted that it
must be
Photo: Minden Pictures/SuperStock

pollinated by an
insect with a very
long proboscis
(sucking
mouthpart) Nectar spur

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© 2021 Prof. Megan Frederickson 3

Reciprocal adaptation Bacterial
(coevolution) between endosymbionts in

Photo: quantamagazine.org
flowers and insects aphids
• Aphids feed on phloem
sap that is rich in sugars,
but poor in essential
amino acids Nancy Moran
• Aphids have intracellular
bacteria (Buchnera) that
provide their hosts with
essential amino acids
• Buchnera are vertically
transmitted; they are
passed in aphid eggs
Pauw et al. (2009, Evolution) from mothers to offspring
19 Aphids 20

Vertically transmitted endosymbionts Are mutualisms often highly

often have tiny genomes
• Buchnera has a much smaller genome than free-
living bacteria (e.g., E. coli); other endosymbiotic
bacteria also have tiny genomes
• In humans: mitochondrial genome is ~17000 base
pairs (and encodes just 37 genes); nuclear
genome is > 3 billion base pairs
• Endosymbiotic bacteria lose genes that they no
longer need
• Some functions unnecessary because bacteria are no
longer free-living; bacteria protected inside host cells
• Other functions ‘outsourced’ to host genome
specialized?
• Most aphid species have their own species of Buchnera
bacteria
• But most mutualisms are NOT tightly coevolved,
species-specific interactions
• Most mutualisms are horizontally transmitted; partners
are acquired anew each generation
• Mutualisms are rarely one-to-one interactions; usually
many-to-many interactions
• Current ‘hot’ areas of mutualism research include:
• Understanding networks of interactions among large numbers
of species
• Microbiomes: this term refers to either all the microbes living
together in a community (often, a host) or their collective
genomes

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Most plants have many pollinator

species; most pollinators visit many Microbiomes
plant species
A plant-pollinator
network in Arctic
tundra, Greenland

Orange circles = pollinators

Green dots = plants
Blue lines = interactions

Just a few recent microbiome headlines from the

Bascompte & Jordano (2007, AREES)
23 CBC and the New York Times 24

© 2021 Prof. Megan Frederickson 4

 Characterizing microbial diversity Diverse mammalian gut microbiomes
Humans
in a host (or environmental sample)
• Culture-based • Sequencing-based
methods methods
• Sequence a highly
conserved (i.e., slowly
evolving) gene, usually the
bacterial 16S rRNA gene
• Cluster sequences into
operational taxonomic units
(OTUs) based on sequence
similarity
• Frees us from having to
culture microbes in order to
study them

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Ley et al. (2008, Science)

Mammal gut
microbiomes Microbiome research is a rapidly
reflect diet, moving field
phylogeny,
• Accumulating evidence suggests a host’s microbiome
and affects its metabolism, immune system, and other traits
morphology • Researchers have compared microbiomes among
human cultures, body parts, and medical conditions
• Microbiome researchers often ask the same questions
ecologists ask of any community:
• What determines which and how many species live together
in a community?
• How do communities change across space or time or along
Photo: leylab.com

environmental gradients?
• Can we predict complex community dynamics from knowing
what happens in simpler systems involving only one or a few
species?
• How do species adapt to their (host) environment and each
other?
Ruth Ley
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Ley et al. (2008, Science)

Lecture 9– Key things to know

Topics & concepts – symbiosis vs. mutualism,

reciprocal exchange of goods or services,
population dynamics of mutualism, invasional
meltdown, how mutualism affects species
diversity, Darwin’s orchid as an example of
reciprocal adaptation, mutualistic networks,
microbiomes

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© 2021 Prof. Megan Frederickson 5