N_1tn2 fixation and denitrification in a floodplain forest in Central Amazonia, Brazil [Elektronische Ressource] / vorgelegt von Heidi Kreibich

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N FIXATION AND DENITRIFICATION 2IN A FLOODPLAIN FOREST IN CENTRAL AMAZONIA, BRAZIL Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften (Dr. rer. nat.) dem Fachbereich Biologie der Philipps-Universität Marburg vorgelegt von HEIDI KREIBICH aus Ansbach Marburg/Lahn 2002 Vom Fachbereich Biologie der Philipps-Universität Marburg als Dissertation am 19.06.2002 angenommen. Erstgutachter Prof. Dr. D. Werner Zweitgutachter Prof. Dr. P. Frenzel Tag der mündlichen Prüfung am 25.06.2002 Parts of this thesis are published in the following papers: KREIBICH H, KERN J (2002) N fixation and denitrification in a floodplain forest near Manaus, 2Brazil. Hydrological Processes (in press) KERN J, KREIBICH H, DARWICH A (2002) Nitrogen dynamics on the Amazon flood plain in relation to the flood pulse of the Solimões River. In: McClain ME (ed.) The Ecohydrology of South American Rivers and Wetlands. IAHS Special Publication no. 6 (in press) Other results have been presented on scientific conferences (with published abstracts): KREIBICH H, KERN J, FÖRSTEL H. Studies on Nitrogen Fixation in Amazonian Floodplain thForests. 12 International Congress on Nitrogen Fixation, Foz do Iguacu, September 12–17, 1999. In: Nitrogen Fixation: From Molecules to Crop Productivity. Pedrosa FO, Hungria M, Yates G, Newton WE (eds.) Kluwer Academic Publishers, Dordrecht 2000, p.

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N FIXATION AND DENITRIFICATION 2
IN A FLOODPLAIN FOREST IN
CENTRAL AMAZONIA, BRAZIL




Dissertation
zur
Erlangung des Doktorgrades
der Naturwissenschaften
(Dr. rer. nat.)





dem
Fachbereich Biologie
der Philipps-Universität Marburg
vorgelegt von

HEIDI KREIBICH
aus Ansbach



Marburg/Lahn 2002






















Vom Fachbereich Biologie
der Philipps-Universität Marburg als Dissertation am 19.06.2002 angenommen.


Erstgutachter Prof. Dr. D. Werner
Zweitgutachter Prof. Dr. P. Frenzel
Tag der mündlichen Prüfung am 25.06.2002
Parts of this thesis are published in the following papers:

KREIBICH H, KERN J (2002) N fixation and denitrification in a floodplain forest near Manaus, 2
Brazil. Hydrological Processes (in press)

KERN J, KREIBICH H, DARWICH A (2002) Nitrogen dynamics on the Amazon flood plain in
relation to the flood pulse of the Solimões River. In: McClain ME (ed.) The
Ecohydrology of South American Rivers and Wetlands. IAHS Special Publication no. 6
(in press)

Other results have been presented on scientific conferences (with published abstracts):

KREIBICH H, KERN J, FÖRSTEL H. Studies on Nitrogen Fixation in Amazonian Floodplain
thForests. 12 International Congress on Nitrogen Fixation, Foz do Iguacu, September
12–17, 1999. In: Nitrogen Fixation: From Molecules to Crop Productivity. Pedrosa FO,
Hungria M, Yates G, Newton WE (eds.) Kluwer Academic Publishers, Dordrecht 2000,
p. 544

KREIBICH H, KERN J. Studies on the role of nitrogen fixation and denitrification in the
floodplain forest of the várzea. International Symposium Manaus’99 Hydrological and
Geochemical Processes in Large Scale River Basins, Manaus, November 15–19, 1999.
In: Ecohydrology. McClain ME, Zalewski M (eds.) UNESCO Technical Documents in
Hydrology No. 47, Paris 2001, p. 31


Contents
_____________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________

1. INTRODUCTION 1
1.1 N cycle 3
1.2 N fixation 7 2
1.3 Denitrification 13
2. STUDY AREA 17
2.1 Geography 17
2.2 Climate 19
2.3 Geology and pedology 20
2.4 Hydrology 21
2.5 Vegetation 22
3. MATERIAL AND METHODS 24
3.1 Forest structure analysis 24
3.2 Physical and chemical soil analyses 28
3.3 Microbiological methods 29
3.4 Genotypic characterisation of rhizobia 31
3.5 Acetylene reduction assay 34
3.6 Acetylene block method 38
153.7 N natural abundance method 40
3.8 Isotope dilution method 45
3.9 Statistics 47
4. RESULTS 49
4.1 Forest 49
Contents
_____________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________
4.1.1 Stand structure 49
4.1.2 Legume species 52
4.2 Soil 54
4.2.1 Physical properties 54
4.2.2 Chemical properties 58
4.2.3 Microbiological properties 61
4.3 Rhizobia 62
4.4 Non-symbiotic N fixation 66 2
4.4.1 N fixation rates and influencing factors 67 2
4.4.2 N gain via non-symbiotic N fixation 71 2
4.5 Symbiotic N fixation in the forest 73 2
154.5.1 δ N of water and soil 73
154.5.2 δ N of the plants 74
4.5.3 Reference plants 81
154.5.4. δ N of N fixing plants grown without soil N 85 2
4.5.5 %Ndfa and N gain of N fixing plants 87 2
4.6 Symbiotic N fixation under experimental conditions 90 2
154.6.1 δ N of the soil 90
154.6.2 δ N values of the plants grown under experimental conditions 91
4.6.3 %Ndfa and the amount of fixed N per nodulated legume 95
4.6.4 Comparison of experimental with forest results 97
4.7 Denitrification 99
4.7.1 Denitrification rates and influencing factors 99
4.7.2 N loss via denitrification 104
5. DISCUSSION 106
5.1 Forest, soil and rhizobia 106
5.2 Non-symbiotic N fixation 113 2Contents
_____________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________
5.3 Symbiotic N fixation 117 2
5.3.1 Symbiotic N fixation in the forest 117 2
5.3.2 Symbiotic N fixation under experimental conditions 124 2
5.4 Denitrification 126
6. N FIXATION AND DENITRIFICATION IN THE CONTEXT OF 2
THE N CYCLE 130
7. SUMMARY 134
7.1 Summary 134
7.2 Zusammenfassung 136
7.3 Resumo 140
8. REFERENCES 143
APPENDIX
ACKNOWLEDGEMENTS



List of abbreviations
_____________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________
List of abbreviations
β isotopic fractionation coefficients:
15 14 15 15 14 15β = ( N/( N+ N)) / ( N/( N+ N)) or approximate: substrate product
15 15β = 1- ( δ N - δ N )/1000 (KOHL and SHEARER, 1980) substarte product
15δ N isotopic ratio of biologically fixed nitrogen from atmosphere A
15δ N reference value (isotopic ratio of non-N fixing reference plants) R 2
%Ndfa percentage of N derived from atmosphere
Abs absorption 260 wavelength
ANOVA analyses of variance
ARA acetylene reduction assay
ARDRA amplified ribosomal DNA restriction analysis
a.s.l. above sea level
ATP Adenosin-triphosphat
bp base pair(s)
CCD charge coupled device
CENA Centro de Energia Nuclear na Agricultura
cfu colony forming unit(s)
CTAB cetyltrimethylammonium bromide
DMSO dimethyl sulfoxide
DNA deoxyribonucleic acid
dw dry weight
EDTA ethylenediamine tetraacetic acid
ECD electron capture detector
EtBr ethidium bromide
FID flame ionisation detector
FIV family important value
FZJ Forschungszentrum Jülich
fw fresh weight
GC gas chromatograph(y)
INPA Instituto Nacional de Pesquisas da Amazônia
IVI important value index
kb kilo base pair(s)
LSD least significant difference List of abbreviations
_____________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________
M molar
MPN Most Probable Number
n number of samples
ND not determined
p probability value
PCR polymerase chain reaction
r Spearman correlation coefficient
R² degree of explanation
rep-PCR repetitive extragenic palindromic sequence-based PCR
RFLP restriction fragment length polymorphism
SDS sodium dodecil sulphate
SHIFT Studies of Human Impact on Forests and Floodplains in the Tropics
sp(p). species
TAE tris-acetate-EDTA buffer
TBE tris-borate-EDTA buffer
TE tris-EDTA
tiff tagged image file format
UGPMA unpaired group method using arithmetic averages
UV ultra violet radiation
v/v volume by volume
w/v weight by volume

used chemical formulas, symbols and unit abbreviations are conform with international
standards