global rates of atmospheric nitrogen deposition
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ELEVATED ATMOSPHERIC NITRATE DEPOSITION IN NORTHERN HARDWOOD FORESTS: IMPACTS ON MICROBIAL MECHANISMS OF PLANT LITTER DECOMPOSITION Jared L. DeForest Earth, Ecological, & Environmental Sciences University of Toledo. Global rates of atmospheric nitrogen deposition. 50.0 20.0 10.0 7.5 5.0 - PowerPoint PPT PresentationTRANSCRIPT
ELEVATED ATMOSPHERIC NITRATE DEPOSITION IN NORTHERN
HARDWOOD FORESTS: IMPACTS ON MICROBIAL MECHANISMS OF PLANT
LITTER DECOMPOSITION
Jared L. DeForest
Earth, Ecological, & Environmental Sciences
University of Toledo
Global rates of atmospheric nitrogen deposition
Galloway & Cowling, (2002)
50.020.010.07.55.02.51.00.50.30.1
kg N ha-1
Total Nitrogen Deposition(2002)
Values in 1012 g; From Schlesinger (1997)
Global Nitrogen Cycle
150
Human activities have doubled the amount of available nitrogen
Values in 1012 g; From Schlesinger (1997)
Global Nitrogen Cycle
150
Human activities have doubled the amount of available nitrogen
The deposition of nitrogen can be in two forms:
Nitrate (NO3-) or Ammonium (NH4
+)
Nitrate represents the majority of total nitrogen deposition in the Midwest
Nitrate is rapidly assimilated by the microbial community and through the process of cell
death, that nitrogen is released as ammonium
Ammonium can represent 75% of extractable total inorganic nitrogen in soil
HumanNitrate
Deposition
GPP
Respiration
Decomposition
Soils1500 Gt C
Land Plants560 Gt C
The doubling of available nitrogen can be a potent modifier of the carbon cycle
Atmosphere
750 Gt C
Adapted from Schlesinger (1997)
120 Gt C yr-1
60 Gt C yr-1
60 Gt C yr-1
Increases in nitrogen deposition can inhibit decomposition because high levels of soil
nitrogen can suppress the activity of enzymes that degrade plant litter
Lignin degrading enzymes are the most likely to be suppressed by increases in soil
ammonium availability
0
0.5
1
1.5
2
2.5
0 1 2 3 4 5 6
0
25
50
75
100
Ligninolytic Activity
Total Extracellular
Nitrogen
Ligninolytic activity is often inhibited byammonium (NH4
+)
Culture Age (days)Adapted from Keyser et al., 1978
Lignin
oly
tic Activ
ity
Am
moniu
m (
mM
)
ExtracellularAmmonium
LigninolyticActivity
Basidiomycetes are the primarydecomposers of lignin
White-rot fungi are considered the primary decomposers of lignin because they produce an array
of enzymes that can fully degrade lignin.
Degrading lignin is a specialized function giving lignin-degrading microorganism access to lignified
carbohydrates.
A relatively small population of soil bacteria, actinomycete, and fungi have the ability to
depolymerize lignin by non-enzymatic and enzymatic means.
Evidence of White-RotDecomposition
White-rot fungi are a physiological, rather
than a taxonomic, grouping of fungi.
At least 21 genera are considered
white-rot fungi.
The decomposition of lignin is important because:
Lignin is the second most abundant
organic molecule
Lignin protectsplant tissue from decomposition
Lignin
Time
Rem
ain
ing M
ass
Lignin
Lignified Cellulose
Non Lignified Cellulose
Labile Compounds
The Decomposition of Plant Litter
Adapted from Berg (1986)
Time
Rem
ain
ing M
ass
Lignin
Non Lignified Cellulose
Labile Compounds
Phase regulated by lignindecomposition rate
Phase regulatedby nutrient
level and readilyavailable carbon
Adapted from Berg (1986)
Lignified Cellulose
Time
Mass
Loss
AmbientNitrogen
ElevatedNitrogen
Phase regulatedby nutrient
level and readilyavailable carbon
Phase regulatedby lignin
decomposition rate
Adapted from Fog (1988)
LessLignolyticEnzymeActivity
Human NitrateDeposition
More Available
NH4+
Less LigninDecay
MicrobialNitrate Assimilation
And Turnover
LessLitter
Decomposition
ReducedCarbonFlow
Hypothesis
Chronic nitrate additions can suppress the lignin-degrading
activity of soil microbial communities
PredictionsPredictions
Nitrate amended soils will have:
A microbial community composition with less fungi
Lower activity of enzymes that degrade lignin and cellulose
StudySites7
9
12(kg N ha-1 y-1)
12
PLOTS
AmbientNitrogen
Deposition
Ambient NDeposition Plus
30 kg N-NO3-
ha-1 y-1A
B
D
C
Cell membranes can be used to determine Cell membranes can be used to determine microbial community compositionmicrobial community composition
Microbialcell
Cell membraneLipid bilayer
PhospholipidPhospholipid
Phospholipid Fatty AcidsPhospholipid Fatty Acids
Common to manysoil microorganisms
Unique tofungi
The length of fatty acid tails and position of double bonds on the tails can be unique to broad
taxonomic groups
FattyAcidsTails
Cellulose
Lignin
Plant Litter Compound
-glucosidase
Peroxidase
Extracellular Enzymes
Cellobiohydrolase
Phenol oxidase
Cellulose
Lignin
Enzyme AnalysisEnzyme Analysis
% m
ol f
ract
ion
Nitrate additions had no noticeable effect Nitrate additions had no noticeable effect on microbial community compositionon microbial community composition
Nitrate additions decreased microbial Nitrate additions decreased microbial biomassbiomass
0
2
4
6
8
10
p = 0.012
Control N Amended
Tot
al P
LF
A (
nm
ol P
LF
A
g-1 C
)
Nitrate addition suppressed activity of soil Nitrate addition suppressed activity of soil lignin & cellulose degrading enzymes lignin & cellulose degrading enzymes
-40% -30% -20% -10% 0% 10%
Change in Enzyme Activity
Cellobiohydrolase
-glucosidase
Peroxidase
Phenol Oxidase
*
*
* p < 0.05
Nitrate addition suppressed activity of Nitrate addition suppressed activity of lignin degrading enzymes in litterlignin degrading enzymes in litter
-40% -30% -20% -10% 0% 10%
Change in Enzyme Activity
Cellobiohydrolase
-glucosidase
Peroxidase
Phenol Oxidase*
* p < 0.05
NitrateAdditions
MicrobialCommunityComposition
Total PLFA(Microbial Biomass)
NoApparentChange
Decrease
LignolyticActivity
Decrease
Decreases in -glucosidase activity can help explain lower microbial
biomass in nitrate amended soils.
Reductions in -glucosidase activity can diminish the physiological
capacity of the microbial community to metabolize cellulose.
This reduction could reduce the energy enzymatically derived from
cellulose degradation.
Conclusions
Anthropogenic nitrate Anthropogenic nitrate deposition may diminish the deposition may diminish the physiological capacity of soil physiological capacity of soil
microbial communities to microbial communities to degrade plant litter.degrade plant litter.
Does a suppression of lignin & Does a suppression of lignin & cellulose degrading enzymes cellulose degrading enzymes
indicate a reduction in the flow indicate a reduction in the flow of carbon from these of carbon from these
compounds?compounds?
HypothesisHypothesis
Nitrate additions will inhibitthe ability of soil microorganismsto metabolize and assimilate theproducts of lignin and cellulose
degradation
13C Vanillin
MicrobialAssimilation
LigninCHO
OCH3
OHH
13C Cellobiose MicrobialAssimilation
Cellulose
13C Sequential Extractions:
Soil was incubated for 48 hours and 13C was traced into respiration, dissolved organic carbon (DOC), microbial carbon, and soil carbon.
13C PLFA Analysis:
Traced the flow of labeled 13C vanillin and cellobiose into cell membranes.
1313C PLFA AnalysisC PLFA AnalysisMicrobial
MembraneExtraction &Separation
Analysis
CHO
OCH3
OHH
13C13C
13C
13C
13C13C
13C 13C
13C13C
N additions increased the incorporation N additions increased the incorporation of vanillin into PLFAsof vanillin into PLFAs
**
*
Cellobiose
Vanillin
N additions did not alter the flow of N additions did not alter the flow of 1313C C vanillin into carbon poolsvanillin into carbon pools
0
20
40
60
80
Microbial
Respiration
DOC Microbial
Biomass
Soil Organic
Matter
% R
ecov
ery
13C
Control
N Amended
N additions did not alter the flow of N additions did not alter the flow of 1313C C cellobiose into carbon poolscellobiose into carbon pools
0
20
40
60
80
Microbial
Respiration
DOC Microbial
Biomass
Soil Organic
Matter
% R
ecov
ery
13C
Control
N Amended
0
10
20
30
40
50
60
p < 0.001
Control N Amended
N additions increased soil organic N additions increased soil organic carboncarbon
Soil O
rgan
ic C
arb
on
(g
C g
-1)
Chronicnitrate additions
UNCHANGEDVanillin or
Cellobiose intoCarbon Pools
INCREASED Soil Organic
Carbon
Excess nitrogen likely inhibits lignocellulose degradation more than vanillin or cellobiose
degradation
ConclusionsConclusions
Nitrate additions have apparently stemmed the
flow of carbon through the soil food web evident by increasing soil organic
matter formation through a reduction in lignolytic
activity.
ImplicationsImplications
Northern Hardwood Forests
AtmosphericCO2
Pools
SlowerDecomposition
HumanNitrogen
Deposition
Global ImplicationGlobal Implication
The same mechanism that decreases lignin
decomposition could be used to understand the
impact nitrogen deposition may have on broad global patterns of decomposition
Plant litter decay
LitterBiochemistry
Environmental Conditions
Global Controls of Decomposition
Actual Evapotranspirat
ion(AET)
EnvironmentalConditions
High > 1000 mm Low < 300 mm
Faster Decomposition Slower Decomposition
350-550 mm
EnvironmentalConditions
Wet Tropical Boreal
Years Required to Decompose95% of Leaf Litter
~0.5 years ~14 years
TemperateDeciduous
~4 years
LitterBiochemistry
Lignin and Nitrogen
Concentrations
Lower LigninHigher Nitrogen
Faster Decomposition Slower Decomposition
Higher LigninLower Nitrogen
HumanNitrogen
Deposition
Increase LitterNitrogen Concentrations
Decrease theDecomposition of Lignin
Increase DecompositionRates
Decrease DecompositionRates
Time
Mass
Loss
AmbientNitrogen
ElevatedNitrogen
Phase regulatedby nutrient
level and readilyavailable carbon
Phase regulatedby lignin
decomposition rate
Adapted from Fog (1988)
“Low”Lignin
“High”Lignin
Increased Decay
Nitrogen deposition impact on decomposition may depend on lignin concentrations
Decreased Decay
0
25
50
75
100
0 5 10 15 20 25 30
AET
1000 (mm)
500 (mm)
250 (mm)
Lignin Control of Decay is Greater at Higher AET A
nn
ual D
ecom
posit
ion
Rate
(%
)
Lignin Concentration (%)
Slope = -1.50
Adapted from Meentemeyer (1978)
Slope = -0.75
Slope = -0.40
As slope decreases, higher lignin
concentrations require more energy and
moisture to cause decay.
LessImpact
MoreImpact
Anthropogenic nitrogen deposition may Anthropogenic nitrogen deposition may have a larger impact on decomposition have a larger impact on decomposition
in wet-tropical environmentsin wet-tropical environments
SummarySummary
Nitrogen deposition has the potential to diminish the physiological capacity of lignin-degrading microorganisms to depolymerize lignin.
Reductions in lignocellulose-degrading enzymes and microbial biomass suggests a reduction in energy
available for microbial metabolism
Nitrogen deposition may have a greater impact on decomposition in wet tropical regions than arid or
cold regions