How

do soil microbes respond to

drought and rainfall?
Sarah Evans
Assistant Professor
Kellogg Biological Sta>on

Major ques>ons addressed in this

lecture
How does rainfall/drought alter the soil
environment?
What kind of stress does this impose for a
microbe?
What traits do microbes use to tolerate these
stresses?

But rst: why should we care?

Rainfall drives biology
Biology drives process
Rainfall changing in future
Process changing in future?

Background:
Climate - carbon - microbe interac5ons

CO2
Soil carbon
Climate
change

Climate: predicted shiLs in rainfall

More intense rainfall regimes IPCC 2007
= Higher propor>on of rain from large events
= Fluctua>on in soil moisture (dry then rewet)

More intense rainfall

 Microbes decompose soil C and produce CO2

Sudden changes in moisture are stressful to
microbes

CO2

Ques5ons:
1. How do changes in rainfall aect microbes?
2. How do these changes aect carbon ux?

CO2

How do changes in rainfall aect plants?

How does this aect carbon ux?

Mean annual temperature

Rain aects plant physiology and species

composi>on
Dierent plants will have dierent eects on
carbon ux

Mean annual precipita>on

How soil water aects microbes

Various ways that soil water affects microbial activity:

Physical (solute movement

Chemical (nutrient limited)
Biological (osmotic stress)
Physical (microbe
movement)

Chemical (O2 limitation)

Biological (desiccation
stress)

rate of solute diffusion through soil

nutrient availability / uptake into cells
solute concentration; hyperosmotic stress in saline soils
microbial movement in soil (cells swim in the fluid phase).
Bacterial movement ceases when soil dries sufficiently to
produce discontinuous water-filled pores, or when the water
film is smaller than whats needed for the microbe to swim.
For microbial motility, water films must be > 0.5 m for
bacteria; 4 m for protozoa and flagellated fungal
zoospores.
Impacts directly on status of soil aeration [especially O2
diffusion] within pore spaces, hence regulates aerobic vs.
anaerobic metabolism
drying imposes desiccation stress, common in surface soils

The soil environment

Flight through soil
Physical (solute movement
1. Physical (microbe movement)


Chemical (nutrient limited)
2. Chemical (O2 limitation)

Biological (osmotic stress)
3.
Biological (desiccation stress)

Physical constraints
Physical (solute movement
Physical (microbe movement)

 Chemical (nutrient limited)

Chemical (O2 limitation)

Chemical

 Biological (osmotic stress)

Biological (desiccation stress)

Drought

Microbial cell

Biological
Large rainfall
event

Rest of lecture: Two studies

I.

Do micro-scale mechanisms help explain soil

CO2 ux aLer a large rain event?

--Fill in traits table--

II. Can we use life history strategies to
understand microbial responses to moisture
stress?

I. Do micro-scale mechanisms help explain soil

CO2 ux aIer a large rain event?
Dicult to predict
Moisture
pulse

Observed
Predicted
Are there other m
echanisms

we havent accounted for

that could be causing this
large pulse of CO2?

Li et al. 2005

First, how is CO2 produced in soil?

1.Need microbes
2.Need movement

Enzymes
Organic
maPer

Dissolved
substrate

Biogeochemical - microbial model

Individual-based, theore>cal model

1 cm
Micro-scale
Microbial func5onal groups
3 func>onal groups
Quan>es output
Enzyme ac5vity
Respira5on

DOC:DON

Implemented water dynamics

180
Water level (mm3)

160

Rainfall

140
120
100

Water level linked to:

Diusion
Leaching
Microbial growth

80
60

Evapora>on

40
20
0

Time (days)

Mechanisms: processes responding to water

1. Water-dependent growth
2. Microbial diversity
3. Substrate diusion

Simula>ons
Growth of dierent
microbial groups

Producers
Cheaters

Patches

Diusion of substrates

Cheaters (red) respond

Rainfall
pahern:

Reproducing CO2 pulse with model

1. Water-dependent growth
With m
2. Microbial
dechanisms,
iversity
One water simulated
pulse pulse
3. Substrate diusion

CO2

What is the rela>ve inuence of

dierent mechanisms on CO2 pulse?

CO2

Mechanisms, in dierent combina>ons:

1. Water-dependent growth
2. Microbial diversity
3. Substrate diusion

Time

Rela>ve inuence of dierent mechanisms

Exclud

e Diu
s

22%

ion

e Grow
th

Exclud

CO2 respired (fmol C/hour)

86 %

Proportion of maximum pulse (%)

Produced highest:
Diusion + Water- Add
dependent growth microbial
diversity
100 %

Diusion contributed
more
Eect of diversity
depends on:
Compe>>ve
interac>ons
Specic func>onal
groups present

Conclusions: study 1
Reproduced biogeochemical pulse with a model
Once separated, physical factors (diusion)
might inuence CO2 ux more than biological
stress
CO2 ux did depend on microbial func>onal
groups that were present, but needs more work

Outline of talk: Two studies

I.

Do micro-scale mechanisms help explain soil

CO2 ux aLer a large rain event?

II. Can we use life history strategies to

understand microbial responses to moisture
stress?

Exercise
What traits would be useful to survive under
drought?

Trait
Can move (mo5lity)

Does it contribute to drought What are the disadvantages

tolerance? How?
and costs?

Cheat o of others enzyme

produc5on
Ability to store resources
Synthesizes intra-cellular
solutes
Can change metabolism
(plas5c)
Large cell size
Responds quickly to
environment
Can form spores
Other?

How can we understand microbial diversity

in an ecologically meaningful way?
Want to categorize microbes based on
their ecology
Cant measure all species traits directly

Ecological strategies
Way of organizing organisms based on growth
or stress response
e.g. r- and k- strategists (MacArthur and Wilson 1967)
life history strategy in plants (Grime 1979)
opportunist, ruderal, tolerant

Do we see ecological strategies in microbes?

Strategy based on historical condi>ons

Certain disturbance regime
results in certain life strategy
Evidence accumula>ng, history
mahers for microbes too
Disturbance regime (history)

Specic ques>ons: study 2

1. Do microbial species express certain
strategies when responding to a stress?
Are these strategies related to phylogeny?

2. Does the distribu>on of these strategies

change with disturbance history?
Is this due to shi4s in species or in strategy?

Disturbance regime: uctua>on in

moisture condi>ons

Methods: describing life strategies

Used 454 pyrosequencing
to describe rela>ve
abundance of each species
MODERATE HIGH

Visualize on heatmap
Hierarchical clustering

LOW

N=5

Results:

Q1: Do species cluster into strategies?

Strategy
Tolerant


Opportunis>c



Sensi>ve

Related to phylogeny?


Tolerant

Opportunis>c

Sensi>ve

Plot strategy on
phylogene>c tree

Does strategy
relate to
phylogeny?

Tolerant
Opportunis>c
Sensi>ve

Ques>ons
1. Do microbial species express certain
strategies when responding to a stress?

Yes

Are these strategies related to phylogeny? Varies

2. Does the distribu@on of these strategies

change with disturbance history?
Is this due to shiLs in species or shiLs in
organism strategy?

Rainfall Manipula>on Plot Study (RaMPS)

Manipulated plot: altered rainfall >ming
more >me between rainfall events, larger events
Konza biological sta@on

Ambient rainfall

Lab drying-rewesng

More drying rewesng

KONZA Biological Research Sta>on

Evans and Wallenstein 2012 Biogeochemistry

Results:

Q2: Does history aect distribu>on of strategies?

Ambient rainfall in the eld

Exposed to moisture uctua>on

Tolerant

Opportunis>c



Sensi>ve

Specula>on: related to carbon ux?

Tolerance to drying-rewesng could mean more
carbon for growth, less carbon respired.
Communi>es from Intense Rainfall plots did respire less
when exposed to moisture pulse1
Ambient rainfall in the eld
Tolerant strategy

Exposed to moisture uctua>on

Lower soil
respira@on
because more
tolerant
species?
1Evans and Wallenstein

2012 Biogeochemistry

Change species or change strategy?

Tolerant

Opportunis>c

Sensi>ve

Change species or change strategy?

Previously
exposed to
moisture pulses

75% were Control

25%
dierent spp
overlap

Do the species that responded from both

historical treatments respond in the same
way? Is life strategy conserved?
Tolerant

Opportunis>c

Sensi>ve

Change strategy?
Outside:
Control eld
treatment

Inside: drying-
reweMng
disturbance history

82% of
species
changed
strategy
Tolerant
Opportunis>c
Sensi>ve

Colors of tree = Phyla

Results summary
1. Do microbial species express certain
strategies when responding to a stress? Yes
Are these strategies related to phylogeny? Varies

2. Does the distribu@on of these strategies

change with disturbance history? Yes
Is this due to shiLs in species or shiLs in
organism strategy?

Both occur

Review: What makes a Tolerant,

Opportunis>c, or Sensi>ve taxa?
Strategy
Tolerant


Opportunis>c



Sensi>ve

Tolerant
Opportunis>c
Sensi>ve

Review: Drought in soils imposes physical,

chemical, and biological stress on microbes
Physical (solute movement
1.
Physical (microbe movement)


Chemical (nutrient limited)
2.
Chemical (O2 limitation)


Biological (osmotic stress)
3.
Biological (desiccation stress)

Trait

Like any disturbance,

need specic traits to
survive.

Traits come at a cost

Impacts carbon cycling

Can move (mo5lity)

Cheat o of others enzyme

produc5on
Ability to store resources
Synthesizes intra-cellular
solutes
Can change metabolism
(plas5c)
Large cell size
Responds quickly to
environment
Can form spores
Other?

Climate - carbon - microbe interac5ons

CO2
Soil carbon
Climate
change