title: Niche segregation of ammonia-oxidizing archaea and anammox bacteria in the Arabian Sea oxygen minimum zone.
abstract: Ammonia-oxidizing archaea (AOA) and anaerobic ammonia-oxidizing (anammox) bacteria have emerged as significant factors in the marine nitrogen cycle and are responsible for the oxidation of ammonium to nitrite and dinitrogen gas, respectively. Potential for an interaction between these groups exists; however, their distributions are rarely determined in tandem. Here we have examined the vertical distribution of AOA and anammox bacteria through the Arabian Sea oxygen minimum zone (OMZ), one of the most intense and vertically exaggerated OMZs in the global ocean, using a unique combination of intact polar lipid (IPL) and gene-based analyses, at both DNA and RNA levels. To screen for AOA-specific IPLs, we developed a high-performance liquid chromatography/mass spectrometry/mass spectrometry method targeting hexose-phosphohexose (HPH) crenarchaeol, a common IPL of cultivated AOA. HPH-crenarchaeol showed highest abundances in the upper OMZ transition zone at oxygen concentrations of ca. 5  μM, coincident with peaks in both thaumarchaeotal 16S rDNA and amoA gene abundances and gene expression. In contrast, concentrations of anammox-specific IPLs peaked within the core of the OMZ at 600  m, where oxygen reached the lowest concentrations, and coincided with peak anammox 16S rDNA and the hydrazine oxidoreductase (hzo) gene abundances and their expression. Taken together, the data reveal a unique depth distribution of abundant AOA and anammox bacteria and the segregation of their respective niches by >400  m, suggesting no direct coupling of their metabolisms at the time and site of sampling in the Arabian Sea OMZ.
authors: Department of Marine Organic Biogeochemistry, NIOZ Royal Netherlands Institute for Sea Research, Den Burg, The Netherlands.
authors: Angela
authors: A
authors: Pitcher
authors: Laura
authors: L
authors: Villanueva
authors: Ellen C
authors: EC
authors: Hopmans
authors: Stefan
authors: S
authors: Schouten
authors: Gert-Jan
authors: GJ
authors: Reichart
authors: Jaap S
authors: JS
authors: Sinninghe Damsté
references: Trends Microbiol. 2010 Aug;18(8):331-40
references: 20598889
references: Environ Microbiol. 2010 Sep;12(9):2450-65
references: 20406296
references: Environ Microbiol. 2000 Aug;2(4):355-65
references: 11234923
references: Environ Microbiol. 2008 Mar;10(3):810-8
references: 18205821
references: FEMS Microbiol Ecol. 2007 Dec;62(3):242-57
references: 17991018
references: Nature. 2005 Sep 15;437(7057):349-55
references: 16163345
references: Environ Microbiol. 2008 Nov;10(11):2931-41
references: 18973620
references: Oecologia. 1979 Jan;40(1):51-62
references: 28309603
references: Environ Microbiol. 2010 Mar;12(3):679-88
references: 20002133
references: Environ Microbiol. 2007 May;9(5):1162-75
references: 17472632
references: Nature. 2005 Sep 22;437(7058):543-6
references: 16177789
references: Appl Environ Microbiol. 2010 Nov;76(22):7575-87
references: 20870784
references: Appl Environ Microbiol. 2007 Jul;73(14):4648-57
references: 17526789
references: Appl Microbiol Biotechnol. 2010 Mar;86(2):781-90
references: 20107988
references: Appl Environ Microbiol. 2009 Aug;75(16):5345-55
references: 19542347
references: Environ Microbiol. 2009 Apr;11(4):971-80
references: 19077007
references: Proc Natl Acad Sci U S A. 2005 Oct 11;102(41):14683-8
references: 16186488
references: Nature. 2003 Apr 10;422(6932):608-11
references: 12686999
references: Science. 2010 Dec 3;330(6009):1375-8
references: 21071631
references: ISME J. 2010 Apr;4(4):542-52
references: 20033067
references: Proc Natl Acad Sci U S A. 2010 Nov 2;107(44):18755-60
references: 20974919
references: Can J Biochem Physiol. 1959 Aug;37(8):911-7
references: 13671378
references: Appl Environ Microbiol. 2011 May;77(10):3468-77
references: 21441324
references: Appl Environ Microbiol. 2008 Apr;74(8):2433-40
references: 18296531
references: ISME J. 2008 Apr;2(4):429-41
references: 18200070
references: Nature. 2009 Sep 3;461(7260):49-50
references: 19727190
references: Proc Natl Acad Sci U S A. 2009 Mar 24;106(12):4752-7
references: 19255441
references: Chem Rev. 2007 Feb;107(2):577-89
references: 17300141
references: Nat Rev Microbiol. 2008 Mar;6(3):245-52
references: 18274537
references: FEMS Microbiol Rev. 2009 Sep;33(5):855-69
references: 19453522
references: Environ Microbiol. 2009 Sep;11(9):2228-43
references: 19453700
references: Appl Environ Microbiol. 1997 Jun;63(6):2446-8
references: 16535633
references: Proc Natl Acad Sci U S A. 2008 Feb 12;105(6):2134-9
references: 18250313
references: Nature. 2009 Sep 3;461(7260):78-81
references: 19727197
references: Nature. 2009 Oct 15;461(7266):976-9
references: 19794413
references: Environ Microbiol. 2007 Apr;9(4):1001-16
references: 17359272
references: Nature. 2001 Jan 25;409(6819):507-10
references: 11206545
references: Nature. 2003 Apr 10;422(6932):606-8
references: 12686998
references: Environ Microbiol. 2010 Jul;12(7):1989-2006
references: 20345944
references: Appl Environ Microbiol. 2002 Jun;68(6):2997-3002
references: 12039760
references: Proc Natl Acad Sci U S A. 2006 Aug 15;103(33):12317-22
references: 16894176
references: Nature. 1999 Jul 29;400(6743):446-9
references: 10440372
references: Proc Natl Acad Sci U S A. 2005 May 3;102(18):6478-83
references: 15843458
references: Proc Natl Acad Sci U S A. 2007 Apr 24;104(17):7104-9
references: 17420469