Mutualism

To do: Define symbiosis and mutualism

•Chap.9 Mutualism •2
Symbiosis
Symbiosis: the intimate and prolonged
association between two or more
organisms of different species.

-Parasitism (+, -)
-Mutualism (+, +)
-Commensalism (+, o)

hackberry nipple gall

Symbiosis

•Symbiosis refers to a close and prolonged interaction between organisms of different

species. Previously, the term is restricted to a mutualistic relationship wherein both
organisms benefit from the interaction. In mutualism, the relationship between the two
organisms is interdependent. Each one gains benefits from the other. This type of
relationship between different species where both of the organisms in question benefit from
the presence of the other. An example is the hermit crab whose shell offers a niche for
anemones to exist in which the anemone can defend the crab with its stinging capabilities.
•At present, the scope of the term became broader. Now, it includes other forms of
associations such as parasitism and commensalism. Anton de Bary's original definition of
symbiosis is it is a long-term relationship between two different species. To this can be
added, mutualism, (benefit for both), commensalism (benefit for one neutral for the other)
and parasitism (benefit for one, costly for the other).
•
Word origin: Greek symbíōsis, sym- + biō, bioûn (to live) + -sis
Parasites

Parasites can have hugely varied life

forms.

Microscopic: viruses, bacteria,

fungal. Associated with “disease”

Bird flu virus

Parasites

Parasites can have hugely varied

life forms.

Macroscopic: flatworms,
roundworms, lice, ticks, smuts
and “plants.”

Mistletoe: a parasite on trees in N. American and

Europe. Can induce kissing for those standing under
during the Yuletide.
Parasites

Parasites can be ecotoparasites or

endoparasite.

They can have intermediate hosts which

may not have any negative consequences.

They often move around using “vectors.”

Relationships can be quite complex.
Ticks carry Lyme disease. Deer carry ticks
Lyme disease is a bacterial parasite that is
transmitted through tick bites. In general,
the ticks are moved around by mammals_
especially white-tailed deer. The ecology of
white-tailed deer, then is directly related to
human health because of disease
transmission.
Parasites
Many parasites/diseases of humans (and other
animals) have very complex ecological
relationships.

Example: Malaria is a disease caused by a

parasitic protozoan. The protozoan is
transferred effectively by mosquitoes.
(particularly Anopheles). Malaria infection kills
~ 1 million people/year.
Parasites

Meningeal worm infection. Snails carry the larvae. Deer

accidentally eat the snails when grazing. Larva escape the
snails, move into the blood stream, infect tissues surrounding
the brain, eventually move into the lungs.
Hosts evolve defenses,
parasites evolve new
mechanisms. Long-term
interplay of selection between
hosts and parasites..
Parasitism can, and often does,
interact with other aspects of an
organism’s ecology.

In this case, the fish is infected

with a parasite, its behavior
changes, and then more of those
individuals are eaten by birds.
Mutualism

•In this type of symbiosis, both organisms of different species rely on one another
for nutrients, protection and other life functions, hence, they are usually found living in
close proximity.
•It can be thought of as a form of "biological barter" since the species trade resources (for
example carbohydrates or inorganic compounds), or services such
as gamete or offspring dispersal, or protection from predators.
•There are two types of mutualism: obligate mutualism and facultative mutualism.
•
Word origin: L. mūtu(us): mutual, reciprocal + Gk -ismos, -isma: often directly, often
through.
Mutualism

Mutualisms are extremely varied,

influencing virtually every living
organism (and, thus) ecosystem on
Earth.

As mentioned before, humans are a

mutualistic assemblage.

Some organisms are very clearly

mutualists.

Lichens are a fascinating example

Mutualism

The mutualism is so tight for lichens

that they are sometimes called
“composite organisms!” They consist of
an intimate mingling of fungus and
algae.

Fungal piece is the “mycobiont” and

provides nutrients.

An algea- generally green algae is the

“photobiont” and provides energy
through photosynthesis.

If you think about it, that is a wicked

combination! Lichens are extremely
hardy organisms_ able to survive in
very extreme conditions. (Example- on
the shingles of your roof!).

If you grow the fungus and algae

separate from one another, they take on
a very different form from the symbiosis.
 Mutualism

Corals are another interesting mutualism. In

this case an animal (the coral “polyp”) is
associated with an algae. The algae
provides energy via photosynthesis. The
coral provides protection and substrate to the
algae.
Mutualism

Rhizobium is a genus of bacteria that forms a

mutualistic relationship with plants in the family
Fabaceae (Legumes…think “peas”). When the
bacteria are free in the soil, they act like bacteria.
When they encounter root hairs of a legume plant they
form a mutualism in which “nodules” are formed.
Within these nodules, the bacteria can “fix” gaseous
nitrogen to ammonia. This can have huge impacts on
soil nitrogen availability.
Mycorrhizae is the name for a mutualism that occurs between
plants and fungi. The fungi attach to root hairs of plants and can
vastly increase the extent of the roots and increase the plants
capacity for “foraging” for nutrient and water. The fungi, in turn,
receives access to the produces of photosynthesis (energy!).
There is a cost to forming the relationship for the plant (giving up
energy) and these relationships are often associated with poor
growing environments. Practically all plants do this!
 RoadMap

• Mutualism: an association between two organisms

that benefits both
• Seed dispersal mutualism: disperser acquires a meal
and the plant gets its seed dispersed
• Through mutualism, species are better able together
to secure resources or better able to defend
themselves
• Mutualism is difficult to model; models tend to result
in runaway densities

•Chap.9 Mutualism •19

 9.1 Plant-Pollinator Mutualism
• Most frequent type of mutualism (Figure 9.1)
– 45% of all studies of mutualism
– Coevolved systems
• Selective pressures for plants to develop intimate
relationship with pollinators (Figure 9.2)
– More than 900 species of Ficus exist and virtually all must be
pollinated by its own species of agaonid wasp
– Yucca plants and yucca moths - highly coevolved
– The distribution of each species is controlled by the
availability of the other species
• Ex. Yucca flower abortion if too many eggs are laid

•Chap.9 Mutualism •20

 0.5
Proportion of mutualism articles

0.4

0.3

0.2

0.1

0
Pollination Seed Ant-plant Mycorrhizal Ant-insect Grazing Nutritional Other
dispersal protection protection

•Fig. 9.1 Frequency of articles on different forms

of mutualism published in 675 papers.
•Chap.9 Mutualism •21
Fig. 9.2 Blastophaga psenes, a tiny fig wasp, which crawl
inside the captifig in California to lay her eggs.
•Chap.9 Mutualism •22
• Cheating in pollination
– In the Bogs of Maine
• Grass-pink orchid produces no nectar, but mimics the
nectar-producing rose pogonia
• Some Bombus species cheat by biting through the base
of the flowers, taking the nectar without entering the
plant nor assisting with pollination

•Chap.9 Mutualism •23

 9.2 Seed Dispersal
• Seed dispersal systems account for almost 30% of all
mutualisms
– In tropics some fruits are dispersed by birds that are
frugivorous
• Fruit provides balanced diet for birds
• Birds disperse seeds
• Mechanisms for attraction
– Birds and mammals - attractive colors, and odorless (Birds)
– Nocturnal bats - give off pungent odor

•Chap.9 Mutualism •24

• Seed dispersal mechanisms are not as
obligatory as plant-pollinator systems
– Performed by more generalist agents
– Wide array of adaptations
• Ex. parrot beaks to crack and peel fruits
• A berry-eating fish from the Amazon. Ex. Figure
9.3

•Chap.9 Mutualism •25

 A Variety of Mutualisms
• Mutualisms and resources
– Neotropical leaf-cutting ants and fungus (Figure 9.4)
• Typical ant colony: 9 million
• Typical biomass of colony: Equivalent to a the biomass of a cow
• Typically cuts the equivalent of a cow's daily requirement of fresh
vegetation
• Leaf-cutting ants harvest 17% of the total leaf production in the
forest
• Ants take cut vegetation underground to grow special fungal crops
(nature's farmers?)
• Fungus grows specialized structures called gongylidia, which serve
as food for the ants

•Chap.9 Mutualism •26

Fig. 9.4 Leaf-cutting ants Atta
cephalotes, in South America,
chew up leaves and cultivate
fungus gardens underground.

•Chap.9 Mutualism •27

• Leaf-cutting ants and the parasitic fungus,
Escovopsis
– Escovopsis can infest the ants' fungi gardens,
destroying the fungi inside
– A mutualisitic streptomyces
• Occurs on the bodies of ants
• Produces antibodies that keep Escovopsis in check

•Chap.9 Mutualism •28

 Nitrogen

• Vital to plant and animal growth

– Most species can not fix atmospheric nitrogen
– Fixation is carried out by soil bacteria and
archaebacteria
• Most live in the roots of plants
• Mutualistic relationship with plant
• Excess nitrogen is available to plant
• Ex. Rhizobia bacteria in legumes
– Figure 9.5

•Chap.9 Mutualism •29

Mutualism under harsh environmental
conditions
• Strong facultative mutualism
• Leguminous shrub, Retama, and an understory plant,
Marrubium vulgare
– Grow in a semiarid region of Spain
– Retama shades Marrubium, provides favorable microclimate
– Marrubium enhances the availability of water for Retama

•Chap.9 Mutualism •30

 Retama Marrubium
30 1.0

Leaf mass (g)

20
0.5
10

0 0.0

Leaf area (cm2 /g)

Leaf area (cm2 /g) 20 100

15 75

10 50

5 25

0 0
0.6 3

0.4 2

0.2 1

•Fig. 9.6 0.0 0

+M -M +R -R
•Chap.9 Mutualism •31
Environment
 Mutualism and protection from
natural enemies
• Common example: ants and aphids(Figure
9.7)
– Aphids feed on plant sap and excrete
honeydew
– Ants drink the honeydew and in return
protect the aphids

•Chap.9 Mutualism •32

 Mutualism and herbivory

• Ants protect plants from herbivores

• Common in the tropics
– 377 myrmecophytic plants per hectare in a
Brazilian Rain Forest (Fonseca and Ganade,
1996)
– 312 ant-plant associations at a single coastal
site in Mexico (Rico-Grey, 1993)

•Chap.9 Mutualism •33

 Fig. 9.8 Thorns on Acacia
collinsii in Paloverde National
Park in Costa Rica.

• Example: Ants and the Central American acacia

trees
– Acacia trees provide food and shelter for the ants inside
large thorns
– Ants protect the acacia tree from other insects and
vertebrate herbivores
– Ants also trim foliage away from competing plants and kill
neighboring plant shoots
– Figure 9.8

•Chap.9 Mutualism •34

• Example: Fungi and plants
– Fungi reduce vertebrate herbivory
– Soay Sheep of Hirta Island (in the St. Kilda
Archipelago) Figure 9.9
– Sheep overgraze native grasses
– Periodically, sheep population crashes

Fig. 9.9 Soay sheep

•Chap.9 Mutualism •35
• Mutualism is the chief culprit
– The main forage is the grass Festuca rubra
– F. rubra contains an endophyte, the fungus Acremonium,
inside its blades
– The fungus produces toxic alkaloids
– These alkaloids function as an anti-herbivory defense
– In return, the fungus obtains food from the plant
– Frequency of infection correlated to grazing pressure. Heavy
grazing causes higher infections
– Fungi are in greatest concentration in basal regions. Heavy
grazing results in sheep reaching lower blades.

•Chap.9 Mutualism •36

 Obligate mutualism
• A mutualistic relationship, in which neither participant
could survive without the other
• Ex. Lichen: a relationship between algae and fungi
– Algae provides the photosynthate
– Fungi provides a safe habitat
• Ex. Many ruminants and symbiotic bacteria
– Bacteria break down plant tissue to provide energy for their hosts
• Ex. The roots of most plants and fungi
– Mutualistic association between the fungus and root tissue -
mycorrhizae
• Fungi obtain carbohydrates from their host
• Fungi increase access to mineral nutrition and water for the plant

•Chap.9 Mutualism •37

To do: Draw a concept map of what effects the
size of a population.

•Chap.9 Mutualism •38

 9.4 Modeling Mutualism
• Uses equations similar to Lotka-Volterra competition
equations
• For facultative mutualism
– dN1 /dt = r1N1 [K1 - N1 + aN2) / K1]
– dN2 /dt = r2N2 [K2 - N2 + bN1) / K2]
• dNi /dt = change in population size of species 1 or 2
• ri = per capita growth rate for species 1 or 2
• Ni = population size of species 1 or 2
• Ki = Maximum population density of species 1 or 2
• a = Positive effect of species 2 on
species 1
• b = Positive effect of species 1 on
species 2

•Chap.9 Mutualism •39

 Facultative mutualism

a) N1 dN b) dN2=0 c) dN
N2 — 1=0 — —1 = 0
Population density of N2

dt dt dt
N2 N2 N2
dN dN
— 2=0
dt
— 2=0
dt

dN
— 1=0
X2 dt
X2 X2

X1 N1 X1 N1 X1 N1
Population density of N1

Fig. 9.10 Graphical models of facultative (a-c) and

obligate (d-f) mutualism.
•Chap.9 Mutualism •40
• For obligate mutualism
– Different equations are needed
– Models of facultative mutualism are, in general,
more stable than models of obligate mutualism
– Figure (9.10d-f)

•Chap.9 Mutualism •41

 Obligate mutualism
N1
d) dN e) dN1 = 0 f) dN
N2 —2= 0 — —1 = 0
dt
dt dt
Population density of N

N2 N2 N2
dN dN
— 1=0 — 2=0
dt
dt

dN 2=0
—
dt

N1 N1 N1
Population density of N1

•Chap.9 Mutualism •42

 9.5 Mutualisms and
Community Process

• Mutualism can have strong indirect effects

on the community
– Ex. mycorrhizal fungi and herbivory load
• Pinyon pines and mycorrhizae: Density of needle
scale insect, Matsucoccus acalyptus:
• Mycorrhizae can improve pine vigor and/or
increase plant investment in antiherbivory
defenses
• Density of needle scale insects will decrease

•Chap.9 Mutualism •43

• Ex. Endophytes (fungi that live inside leaves)
and vascular plant hosts - Defend host against
herbivory
• Mycorrhizal fungi and plant species diversity
(Figure 9.11)

•Chap.9 Mutualism •44

 d)
a) 7
0.8

Simpson’s diversity

Hyphal length
6

(m g -1 soil)
0.6
index 5
0.4
4
0.2 3
0 2
0 2 4 6 8 10 12 14 0 2 4 6 8 10 12 14
e)

-1 soil)
b) 130 20
120
Shoot biomass

15
110

Soil P (mg kg
(g m-2 )

100 10
90
5
80
70 0
0 2 4 6 8 10 12 14 0 2 4 6 8 10 12 14
f)
c) 160 2500
Root biomass (g m -2)

-2
140 2000

120 1500

100
-2 1000 Fig. 9.11
80 500
0 2 4 6 8 10 12 14 0 2 4 6 8 10 12 14
•Chap.9 Mutualism •45
Number of mycorrhizal fungal species Number of mycorrhizal fungal species
To do:
After you watch the biointeractive video, answer
the following questions using the example of the
paper you just read.

¿What are the fundamental question in

mutualism?

•Chap.9 Mutualism •46