Microbial Processes and Features of the Microbiota in Histosols From a Black Alder (Alnus glutinosa (L.) Gaertn.) Forest

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  • This article was downloaded by: [141.214.17.222]On: 20 October 2014, At: 20:51Publisher: Taylor & FrancisInforma Ltd Registered in England and Wales Registered Number:1072954 Registered office: Mortimer House, 37-41 Mortimer Street,London W1T 3JH, UK

    Geomicrobiology JournalPublication details, including instructions forauthors and subscription information:http://www.tandfonline.com/loi/ugmb20

    Microbial Processes andFeatures of the Microbiotain Histosols From a BlackAlder (Alnus glutinosa (L.)Gaertn.) ForestOliver Dilly, Hans-Peter Blume, LudgerKappen, Werner L. Kutsch, Ulrike Middelhoff,Jorg Wotzel, Francois Buscot, Klaus Dittert,Hans-Jurgen Bach, Bernhard Mogge, KarinPritsch, Jean Charles MunchPublished online: 29 Oct 2010.

    To cite this article: Oliver Dilly, Hans-Peter Blume, Ludger Kappen, WernerL. Kutsch, Ulrike Middelhoff, Jorg Wotzel, Francois Buscot, Klaus Dittert,Hans-Jurgen Bach, Bernhard Mogge, Karin Pritsch, Jean Charles Munch (1999)Microbial Processes and Features of the Microbiota in Histosols From a BlackAlder (Alnus glutinosa (L.) Gaertn.) Forest, Geomicrobiology Journal, 16:1,65-78, DOI: 10.1080/014904599270758

    To link to this article: http://dx.doi.org/10.1080/014904599270758

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  • Microbial Processes and Features of theMicrobiota in Histosols From a Black Alder

    (Alnus glutinosa (L.) Gaertn.) Forest

    OLIVER DILLYHANS-PETER BLUMELUDGER KAPPENWERNER L. KUTSCHULRIKE MIDDELHOFFJ ORG W OTZEL

    Okologie-Zentrum

    Universit at KielKiel, Germany

    FRANC OIS BUSCOT

    Institut f ur Ern ahrung und UmweltUniversit at Jena

    Jena, Germany

    KLAUS DITTERT

    Institut f ur P anzenern ahrung und Bodenkunde

    Universit at Kiel

    Kiel, Germany

    HANS-J URGEN BACHBERNHARD MOGGEKARIN PRITSCHJEAN CHARLES MUNCH

    Institut f ur Boden okologie

    GSFForschungszentrum f ur Umwelt und GesundheitNeuherberg, Germany

    Microbiological features and in situ microbial activities were analyzed in soils at ablack alder forest adjacent to the eutrophic Lake Belau during the course of the in-terdisciplinary program, Ecosystem Research in the Bornh oved Lake District. The

    Received 13 January 1998; accepted 24 September 1998.

    We thank Elke Erlebach, Friederike Sch utze, J orn Sprenger, Mirsad Haskovic, Cathrin Schmidt, Anke

    Buckenauer, Birgit Vogt for their excellent technical assistance; Drs. P. Weppen, O. Heinemeyer, E.-A. Kaiser, and

    T.-H. Anderson (FAL, Braunschweig) for the use of laboratory facilities; Dr. U. Schleu for helpful discussion,

    and Nancy A. Weider-Zehrbach for the improvement of the English. These studies were supported by the German

    Ministry of Education, Science, Research and Technology (BMBF), project no. 0339077E, and the state of

    Schleswig-Holstein.

    Address correspondence to Oliver Dilly, Okologie-Zentrum, Universit at Kiel, Schauenburgerstra e 112,

    24118 Kiel, Germany. E-mail: oliver@pz-oekosys.uni-kiel.de

    Geomicrobiology Journal, 16:6578, 1999

    Copyright C 1999 Taylor & Francis0149-0451/99 $12.00 + .00 65

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  • 66 O. Dilly et al.

    microbiological data were combined to evaluate the functional status of the Histosols.It was hypothesized that carbon accumulation typical for Histosols would mainly takeplace at the wet part ( wet site ) close to the lake shore and not at the drier part( dry site ) of the forest. Rates of leaf litter decomposition, in situ soil C mineraliza-tion, and in situ N2- xation were higher at the wet site. Furthermore, the compositionof the bacterial communities and the presence of ectomycorrhizas indicated suf cientO2 availability and high microbial vitality in the soil at the wet site. An anthropogeniclowering of the lake water table during the 1930s seems still to control the actual soilconditions, resulting in humus degradation in the two Histosols of the forest. The twosoils clearly differed in productivity and C and N cycling, being controlled either byupland, acid runoff or by eutrophic lake water. Lake water seems to buffer but also tointensifymicrobial transformationsat the wet site and to supply nutrients, althoughhu-mus decaymay possiblybe deceleratedby a temporarilyhighwater table and refractoryhumic substances.

    Keywords blackalder forest,carboncycling,Histosol,microbialcommunity,nitrogencycling

    Black alder (Alnus glutinosa (L.) Gaertn.) forests are widespread throughout NorthernGermany, occurring naturally near rivers and lakesmostly on soils with high organic carbon

    contents. This type of ecosystem was selected for study because human impact regulating

    thewater table severely controls the structure ofwetland ecosystems.Thus, this system lled

    a signi cant and distinct role during the interdisciplinary program, Ecosystem Research

    in the Bornh oved Lake District, which aims to analyze and model structures, dynamics,and functions of terrestrial and limnic ecosystems. Nineteen subprojects have been carried

    out at the alder forests by 11 German research groups.

    Histosols are soil types with net C accumulation over long periods attributable to

    retarded biological degradation of assimilated C caused by high water levels. However, the

    lakewater tablemay not be the only indicator for estimatingnet C accumulationin soil sincemicrobiologicalprocesses are also affected by proton concentration,mineral nutrient status,

    and vegetation type. Because whole soils, and their constituentparts, should re ect general

    properties of ecosystems (Elliott 1994), the broad spectra of microbiological data obtained

    during the main research period from 1988 to 1995 were combined to elucidate the actual

    functionalstatewith reference to netC accumulationof theHistosols at the alder forest. Bothsoil biochemistry and the structure of the soil microbiota were considered and completed

    by system theory because the data of every component represent only particular aspects,

    may be restricted by the methodology, and, therefore, should not be applied separately for

    drawing general conclusions.

    Materials and Methods

    Site and Soils

    The research site is located 30 km south of Kiel in Schleswig-Holstein, Northern Germany

    (54060 N, 1014 0 E; Figure 1). The landscape, formed during the Pleistocene, consists ofmorainic hills and lakes. The climate is in uenced by both the North Sea and the BalticSea. Long-term (1951 to 1980) mean annual total precipitation was 697 mm, and average

    annual air temperature was 8.1C, according to the local meteorological stations.Along a transect from a kames hill to Lake Belau, a catena running west to east was

    established with a sequence of forests: a beech forest (Fagus sylvatica), a sloping mixedforest, and the black alder forest at the bottom of the catena (Figure 1). The soils of thebeech and the mixed forest were predominantly acidic and sandy; the soils from the black

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  • FIG

    URE1Locationandproperties

    oftheresearch

    site(guregenerouslyprovided

    byDr.W.Kluge,Kiel);H,F,

    S,andUindicatepeat,mud,sand,andsilt,respectively.

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  • 68 O. Dilly et al.

    TABLE 1 Properties of the topsoils of the black alder forest along Lake Belau in theBornh oved Lake district of Northern Germany

    Horizon Depth pHa Corga Cks

    b C/Na C/Nksb C/NRootc(cm) (H2O) [mg g

    1 dry soil] (w/w)

    Dry site H 020 4.1 275 0.35 15 4 36

    Wet site H 020 6.0 230 0.18 15 16 32

    aFrom Dilly and Munch (1995).bks, potassium sulfate-soluble C compounds of soil (control values of fumigation-extraction

    method; sampling October 1994).cFrom Wachendorf et al. (1997).

    alder had high organic matter content. The black alder forest contained single specimens of

    beechandoak (Quercus roburL.) and some shrubs,mainlyhazel (CorylusavellanaL.). Twosites were separated for the investigations in the alder forest: The dry site was situated at

    the bottomof the slope, and the wet site was connectedby a reed belt (PhragmitesaustralisTrin. ex Steud.) to Lake Belau. Soil properties of the two Histosols are presented in Table 1.

    According to the FAO (1988), soils were classi ed as Fibric Histosols with dystrophic and

    eutrophic conditions at the dry and wet site, respectively. The organic horizons, L and Of,

    could easily be differentiated, whereas underlying horizons (topsoil, up to 20 cm deep)could be separated extensively (Wachendorf 1996) and showed a high spatial heterogeneity

    (Figure 1).

    Sampling and Methods

    In situ CO2 emission rates were determined between 15 May and 15 October 1992 witha continuous- ow inverted-box system and an infrared gas analyzer (Kutsch 1996). Four

    boxes were used in parallel, covering an area of 16 12.5 cm = 200 cm2 and a volumeof 2800 cm3. Each box was equipped with 2 Pt100 temperature sensors. Concentrationsof CO2 and mean Pt100 signals at each box were recorded at least 3 times per hour. Data

    were integrated for the measurement period with a model that is dependent on temperature

    (Kutsch and Kappen 1997). After nishing CO2 measurements in October 1992, the mass

    of roots with diameter

  • Microbiota in Histosols From a Black Alder Forest 69

    of all sampling between May and October 1992 (6 samplings) were taken and summarized

    for the period.Microbiological features of the litter were derived from the experiment with leaf litter

    described in detail by Dilly and Munch (1996), in which microbial biomass contentwas es-

    timated by using substrate-induced respiration, and basal respiration rates were determined

    at 22C bothwithoutmodifying the water content ( ) and after adding deionizedwater (+ )to a maximum of 2.5 g of H2O per gram of dry litter.

    For analysis of the topsoils,multiple cores were taken at each site with a drill and mixedtogether,gentlysieved, stored at 4C, and analyzedwithin4 weeks. The fraction

  • 70 O. Dilly et al.

    for the root mass directly below the CO2 measuring system (dry weight including root

    nodules, to a depth of 20 cm), which was 425 and 976 g of dry roots per meter2 at thedry and wet sites, respectively (Figure 3). In agreement to this nding, more detailed

    investigations showed higher amounts and growth rates of roots in the soil at the wet site

    (Table 2). Root growth rates were higher at the dry site only in soil depths between 10 and

    70 cm.

    During the period from May to October, the microbial C content was signi cantly

    higher in leaf litter and topsoil at the dry site than at the wet site (Figures 4 and 5). Incontrast, basal respiration rates and qCO2 did not differ signi cantly between the two sites.However, respiration rates and qCO2 of the leaf litter that had eld moisture content weresigni cantly lower than basal respiration rates at the two sites. The basal respiration rates

    were slightly higher in the leaf litter at the dry site than at the wet site, whereas the opposite

    was observed for data under eld moisture content. The qCO2 was slightly higher in thetopsoil at the wet site.

    N Cycling

    N2 xation by the AlnusFrankia symbiosis was lower at the dry site than at the wet site(Figure 2). The respective amounts, as determined from 15N natural abundance,were 4045

    and 7085 kg N ha 1 a 1 (Dittert 1992).In situN2O emission rates from soil in the alder forest were higher than those from soil

    in the nearby beech forest (Mogge et al. 1998). Within the alder forest, N2O emission rates

    varied signi cantly, being highest at that site reported byMogge et al. (1998). The emissionrates at that site (near the slopingmixed forest), at the dry site, and at the wet site were 1.03,

    0.66, and 0.09 mg N2O-N m 2 d 1, respectively. This estimation was made only once and

    emission rates may change during the year. However, a similar pattern was also determined

    at Lake Belau by Rusch (1996). The modeled denitri cation losses of 60 kg (N) ha 1 a 1

    were similar at the two sites (Wetzel et al. 1996).

    The Cmic/Nmic ratio differed slightly between the dry and wet sites (Figure 5). Arginineammoni cation rates were slightly higher in the soil at the wet site during the observa-

    tion period, whereas protease activity was signi cantly higher in the topsoil at this site

    (Figure 6).

    Structure of the Microbiota

    The dry site was not investigated for bacteria. The dominant culturable bacteria isolated

    on complex heterotrophic media from the topsoil at the wet site were Pseudomonas,Flavobacterium, Cytophaga, Alcaligenes, Arthrobacter, Promicromonospora, and otherunidenti ed organisms. With use of selective media, Pseudomonas uorescens biotypeswere found on gelatin medium (proteolysis); actinomycetes,Bacillus, and Favobacterium /Cytophaga were found on xylan-containing medium; and Cellulomonas, Pseudomonas,and nocardioformes were found on cellulose-containingmedium (Bach 1996).

    The roots of the alder trees were extensively colonized by ectomycorrhizal but not by

    vesciculararbuscularmycorrhizal fungi. The vitality of mycorrhizal rootletswas generallylower at the dry site. In addition, the potential for root regeneration was obviously higher

    at the wet site and was particularly evident during the rapid root growth in spring. The total

    length of the mycorrhizal roots was threefold longer at the wet site than at the dry site. The

    diversity of mycorrhizal types was reduced at the dry site.

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  • FIG

    URE2N2-xationbytheAlnus-Frankia-symbiosis(activevolumeofFrankia-nodules)intheblack

    alderforestalongLakeBelau.

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