Plant-Microbe Interactions

 Plant-microbe interactions diverse – from the plant perspective:

• Negative – e.g. parasitic/pathogenic
• Neutral
• Positive – symbiotic

lecture  important positive interactions with respect to plant

 This
abundance and distribution – related to plant nutrient and water
supply:

Decomposition
Mycorrhizae
N2 fixation
Rhizosphere

 the role of this interaction in the N cycle

I. Decomposition
Primary supplier of plant nutrients – particularly N & P

A. Raw material
Soil organic matter derived primarily from plants –
• Mainly leaves and fine roots
• Wood can be important component in old growth forests

Input rates –
• Generally follow
rates of production
• Deciduous =
evergreen
B. Processes
1. Fragmentation –
• Breakdown of organic matter (OM) into smaller bits = humus
• By soil ‘critters’ – including nematodes, earthworms, springtails,
termites
• consume and excrete OM  incomplete digestion

nematode

springtail (Isotoma viridis) termites

2. Mineralization
• Breakdown OM  inorganic compounds
• Microbial process: accomplished by enzymes excreted into the soil

For Nitrogen
energy for heterotrophic bacteria

proteins amino Mineralization Ammonium

(insoluble) proteases acids NH4+

Nitrification
Nitrite
energy for
NO2-
nitrifying
Microbial uptake bacteria*

Nitrate
Plant uptake NO3-

* In 2 steps by 2 different kinds of bacteria – (1)

Nitrosomonas oxidize NH3 to nitrites + (2)
C. N uptake by plants – Chemical form taken up can vary

- mineralization
1) Nitrate (NO 3)
proteins NH4+
• Preferred by most plants, easier to take up
• Even though requires conversion to NH4+
before be used  lots of energy

• vs. taking up & storing NH4+ problematic

• More strongly bound to soil particles
• Acidifies the soil
• Not easily stored NO3-
plant uptake
2) Ammonium (NH4+ ) –
• Used directly by plants in soils with low nitrification rates (e.g. wet soils)
3) Some plants can take up small amino acids (e.g. glycine)
• Circumvents the need for N mineralization
• Facilitated by mycorrhizae

mineralization
proteins amino NH4+
acids

immobilization
nitrification

microbial uptake

NO3-
Direct uptake
plant uptake
D. Controls on rates of decomposition
1) Temperature –
• Warmer is better
• <45°C Soil Microbial Respiration

2) Moisture – intermediate is best

• Too little  desiccation
• Too much  limits O2 diffusion
T

Soil Moisture %
3) Plant factors – Litter quality
a) Litter C:N ratio (= N concentration)
• If C relative to N high  N limits microbial growth
• Immobilization favored
• N to plants 

Decomposition rate
as fn(lignin, N)

Deciduous forest spp

b) Plant structural material
• Lignin – complex polymer, cell walls
• Confers strength with flexibility
– e.g. oak leaves
• Relatively recalcitrant
• High conc.  lowers decomposition
c) Plant secondary compounds
• Anti-herbivore/microbial OH
• Common are phenolics – e.g. tannins
– Aromatic ring + hydroxyl group, other compounds R

• Control decomposition by:

Bind to enzymes, blocking active sites  lower mineralization

N compounds bind to phenolics  greater immobilization by soil
Phenolics C source for microbes  greater immobilization by microbes

Consequence of controlling
soil OM chemistry and
microclimate …
Plants important factor
controlling spatial
variation in nutrient
cycling
II. Mycorrhizae
A. Symbiotic relationship between plants (roots) & soil fungi
• Plant provides fungus with energy (C)
• Fungus enhances soil resource uptake

Widespread –
• Occurs ~80% angiosperm spp
• All gymnosperms
• Sometimes an obligate relationship
B. Major groups of mycorrhizae:
1) Ectomycorrhizae –
• Fungus forms “sheath” around the root (mantle)
• Grows in between cortical cells = Hartig net – apoplastic
connection

• Occur most often

in woody spp
2) Endomycorrhizae –
• Fungus penetrates cells of root
• Common example is arbuscular mycorrhizae
(AM)
• Found in both herbaceous & woody plants
• Arbuscule = exchange site
Arbuscule in plant cell
C. Function of mycorrhizae:
1) Roles in plant-soil interface –
a) Increase surface area & reach for absorption of soil water & nutrients

b) Increase mobility and uptake of soil P

c) Provides plant with access to organic N
d) Protect roots from toxic heavy metals
e) Protect roots from pathogens

2) Effect of soil nutrient levels on mycorrhizae

• Intermediate soil P concentrations favorable
• Extremely low P – poor fungal infection
• Hi P – plants suppress fungal growth
– taking up P directly

• N saturation
III. N2 Fixation

N2 abundant – chemically inert

N2 must be fixed = converted into chemically usable

form
• Lightning
• High temperature or pressure (humans)
• Biologically fixed

 Nitrogenase – enzyme catalyzes N2  NH3

 Expensive process – ATP, Molybdenum

 Anaerobic – requires special structures

A. Occurs only in prokaryotes:
• Bacteria (e.g. Rhizobium, Frankia)
• Cyanobacteria (e.g. Nostoc, Anabaena)

 Free-living in soil/water – heterocysts

 Symbiotic with plants – root nodules
 Loose association with plants

Anabaena with heterocysts

Symbiosis with plants – Mutualism
• Prokaryote receives carbohydrates
• Plant may allocate up to 30% of its C to the symbiont

• Plant provides anaerobic site – nodules

• Plant receives N
Examples of plant–N2-fixing symbiotic systems –
1) Legumes (Fabaceae)
• Widespread
• bacteria = e.g., Rhizobium spp.
• Those with N2-fixing symbionts form root “nodules”
– anaerobic sites that “house” bacteria
alpine clover

soybean
root
Problem of O2 toxicity –
• Symbionts regulate O2 in the nodule with leghemoglobin
• Different part synthesized by the bacteria and legume

Cross-section of nodules of soybean nodules

 2) Non-legume symbiotic plants –
• “Actinorhizal”= associated with actinomycetes (N2-fixing bacteria)
• genus Frankia
• Usually woody species – e.g. Alders, Ceanothus

• Bacteria in root or small vesicles

Ceanothus velutinus - snowbrush

Ceanothus roots, with

Frankia vesicles

Buffaloberry (Shepherdia argentea)

- actinorhizal shrub (Arizona)
B. Ecological importance of N2 fixation
1) Important in “young” ecosystems –
• Young soils low in organic matter, N
2) Plant-level responses to increased soil N conc:

Some plants (facultative N-fixers) respond to soil N

concentration 

• Plant shifts to direct N uptake

• N fixation 
• Number of nodules decreases
3) Competition: N fixers-plant community interactions
N2-fixing plants higher P, light, Mo, and Fe requirements
 Poor competitors
• Competitive exclusion less earlier in succession
• Though - N2 fixers in “mature” ecosystems

Example N-fixing plants important in early stages of succession:

• Lupines, alders, clovers, Dryas
 Natural N cycle
N 2O
PLANT

IV. N losses from ecosystem

• Leaching  to aquatic systems

• Fire  Volatization REMAINS
• Denitrification  N2, N2O to
atmosphere
– Closes the N cycle! PLANT

• Bacteria mediated
• Anaerobic
 Annual release
NATURAL SOURCES (1012 g N/yr)
Soil bacteria, algae, lightning, etc. 140

ANTHROPOGENIC Annual release

SOURCES (1012 g N/yr)
Altered N cycle
Fertilizer 80
Legumes, other plants 40
Fossil fuels 20
Biomass burning 40
Wetland draining 10
Land clearing 20

Total from human sources 210

From - Peter M. Vitousek et al., "Human Alteration of the Global Nitrogen Cycle - Causes and
Consequences," Issues in Ecology, No. 1 (1997), pp. 4-6.
 V. Rhizosphere interactions
– the belowground foodweb
Fine root

Zone within 2 mm of roots – hotspot of biological activity

• Roots exude C & cells slough off = lots of goodies for soil microbes  lots of microbes for their
consumers (protozoans, arthropods)
• “Free living” N2-fixers thrive in the rhizosphere of some grass species
 Summary
• Plant–microbial interactions play key roles in plant nutrient
dynamics
 Decomposition –
 mineralization, nitrification …
 immobilization, denitrification …

 Rhizosphere – soil foodweb

 Mycorrhizae – plant-fungi symbiosis

 N fixation – plant-bacteria symbiosis

• Highly adapted root morphology and physiology to accommodate

these interactions

• N cycle, for example, significantly altered by human activities