Soil Biol. Biachem.Vol. 22. No. 2. 99. 251-255, 1990 0038-071790 53.00+ 0.

00
Printedin Great Britain. All rights reserved Copyright C 1990 Pcrgamon Press plc

APPLICATION OF ECO-PHYSIOLOGICAL QUOTIENTS

(qC0, AND qD) ON MICROBIAL BIOMASSES FROM
SOILS OF DIFFERENT CROPPING HISTORIES
TRAUTE-HEIDIANDERSONand K. H. DOMSCH
Department of Soil Biology, Federal Research Centre of Agriculture, 3300 Braunschweig. F.R.G.

(Accepd 1s Augusr 1989)

Summary-Metabolic quotients for CO& (~c0r-C) and microbial-C-loss (qD) were studied on soil
microbial communities under long-term monoculture (M) or continuous crop rotations (CR). Under
defined laboratory conditions the mean &0,-C (unit CO,& unit-’ IZ,,,,~h-‘) of different microbial
biomasses from t7 M systems amounted to 1.097 pg COI-C as compared to 0.645 fig CO,< of microbial
biomasses from 19 CR systems. The 1.7 times higher CO& release per unit biomass and time of microbial
biomasses from M systems was significantly different at the P = 0.001 level.
In addition, microbial C-loss in samples from M or CR plots was followed for 5 weeks. Again, mean
qD per unit microbial biomass and time was 1.6 times higher (P = 0.01) for microbial biomasses from
M systems (0.301 fig C. 14 soils) when compared with CR systems (0.188 pg C. 14 soils).
These differences were not related to soil texture, C,,, or pH of these soils. The effects of environmental
influences (soil management) on the microbial pool in terms of a changing energy demand are discussed.

lNTRODUcrlON ccosystcm dcvclopmcnt (Odum, 1969). in the

more complex CR system organisms could have
Rcccnt investigations on the relationship bctwecn cvolvcd a higher yield coctficicnt (Y). That is, as
microbial biomass-C (C,,,) and soil organic C (C,,,) less C of an organic substrate is channellcd into
of long-term agricultural plots located in the tempcr- energy metabolism more C will be fixed in microbial
ate climatic zone of Central Europe (Anderson and cells. This would affect the Cm,c-to-Ce,s ratio. In
Domsch. 1986b; Anderson and Domsch. 1989) or this context the studies on chronoscqucnces of
North America (Insam et al., 3989) have shown that “dcvcloping” soils by lnsam and Domsch (1988) and
by comparison soils under permanent monoculture lnsam and Haselwandter (1989) should bc mcn-
(M) had significantly lower amounts of microbial C tioncd. where a significantly higher metabolic quo-
per unit soil C than soils under continuous crop tient for CO2 (yC02) of microbiat communities from
rotations (CR). Percent C,,, to total Corr of soils “young” sites as compared to “matured” sites was
under permanent M averaged 2.3% as compared to reported.
2.9% for continuous CR of the simplest-the organic To test the possibility of a correlation bctwccn
matter unamended plots. the metabolic status of soil microbial biomasscs
One reason for these yuanrirarire changes observed and long-term cropping management, physiological
bctwcen microbial biomasses from different soil properties of 41 microbial communities have been
managcmcnt regimes could be a higher quantity of investigated. The basal respiration of a soil reflects
residues remaining in the plots under continuous the overall activity or energy spent by the indigenous
rotation. whereby minor shifts towards increases of microbial pool. If basal respiration rates were to
microbial-C will be detected with the measuring be related to the corresponding biomass size. the
procedure applied, while the resolutions of Cars esti- metabolic quotient qC0, so obtained would express
mates may not be equally as sensitive. A similar the CO,-C produced per unit biomass and time.
argument was discussed by Powlson et a!. (1987). Comparison of microbial communities in quan-
This would raise the microbial-C content over that of titative terms would then bc possible. The effect of
the Carl content in comparison to the simpler system, environmental influences or specifically that of dificr-
the M soils. ent cropping systems in the present study on the
On the other hand, the establishment of a microbial pool might be reflected in such &O:
higher microbial-C con~enf per unit soil C in the values. Anderson and Domsch (1986a) found the
CR might be examined from a diRerent point- yC0: a sensitive indicator of soil microbial reactions
of-view. If C availability is one of the driving vari- to differing temperature regimes.
ablcs for the Cmr-to-Gory ratio. the higher chemical In addition, death rate quotients (YD) were dcter-
diversity of organic matter input in CR plots mined. It is reasonable to assume that communities
could have favoured organisms (with time) which which differ in energy economitation should also
arc endowed with a more economic metabolism. respond with differing death rates in non-amended
Applying Odum’s theory on bioenergetics of soils.

251
252 TRAUTE-HEIDIANDERSON and K. H. Dow

MATERIALS AND .WXHODS Determinarion of microbial-C

C,, was estimated by the SIR method (substrate
Soils induced respiration) of Anderson and Domsch
(1978). except that normal outdoor air, containing
In total, 41 different long-term experimental plots 0.035% CO:, was used for aeration (Anderson and
(20 M and 21 CR plots) were analysed. Depending Domsch. 1989). The stored soils were sieved (2 mm)
on plot size. 5-20 soil samples per plot were and adjusted to room temperature for 5 days. The
taken from the upper A, horizon (0-10cm) and optimal water potential for respiratory activity of
bulked. Sampling was done in early spring sieved soils averaged -390 kPa (SD 70). CO:
(March-April), before field management operations evolution was measured at 22 ‘C using a continuously
started (Anderson and Domsch, 1986b). The soils measuring COr analyser (Wiisthoff KG, Bochum.
were stored unsieved and field moist in plastic bags F.R.G).
with cotton plugs as closures at 4C in the dark
for 5 months until analysed. Table I gives infor-
mation on soil characteristics and management. De- Determination of qCO1
tailed information on soil type. cropping practices, For estimation of the metabolic quotient (Pirt,
including crop sequences with respect to rotation 1975) for CO:, 36 soils (17 M and 19CR) were
systems. were given by Anderson and Domsch, analysed. For calculation of the qC0, (unit
(1989). Considered were (mainly) plots free of organic COr-C. unit-’ biomass-C h-‘) of the basal res-
fertilizer input except for harvest residues (stubble, piration (not glucose-activated respiration), the
leaves. chaff. roots etc). The plots were free of winter hourly mean of CO: output over IO h was taken after
cover. soils had reached a relatively constant CO, produc-
All soil analyses are expressed on an oven-dry tion rate after 20 h at 22 ‘C. The calculated quotients
basis. arc means of 3-5 rcplicatc soil samples (5Og).

01’1 CR NII 9s 6.30 O.OY? I.13 x

032 CR FYM 0.1 5.v.I 0.IX I .4? x
01 ‘3 CR NPK 93 6.1X 0. IOU I 26 *
02; I M N IO 7 06 0.165 I.78 x
O+l CR NPK II 7. I ? 0.057 O.64 *
04 ! CR NPK II 7 06 O.OM 0.76
0Ji.l CR NPK I.1 6.X4 0.064 0.77 x
04,4 CR NPK I3 7.47 0.066 0.74 *
07, I NPK I? 6.40 0.035 0.42 *
1x1 c”K s 25 7.40 0.237 2.20 x
1411 M NPK I4 6.44 0.0x7 0.90 x
IS, I M NPK 7 6.59 O.I3? 1.34 x
16,! M s I! 7.30 0. I30 1.75 *
16,J CR Nil 4s 6.6g 0.06 I 0.x0 x
16:s CR FYM .y3 7.21 0 OV5 I.20
I7 I CR NPK 11 6.3’) 0.06n 0.61 *
1712 CR N 32 6.35 0.061 0.64 *
l7,3 CR N 32 6.28 0.077 n.12 *
l7,J CR NPK 32 6 36 0.076 0.76 *
IV ? M NW IV 5.62 0.141 I.XI
IV 3 M NPK 19 5 26 0.147 1.X0
IY 6 M NPfi I9 4.9v 0.125 I.52 x
IY 7 M NPK I9 5.43 0.121 I .52 x
20 I M NPK I4 7.53 0.0x4 0.98 x
?I) ? CR NPK I4 7.99 0.117 1.29 *
xl 3 CR NPii I4 7 97 0.115 I.41 *
?I),4 CR G I.8 7.95 0 II5 I .05 x
21 I CR NPK I7 7.66 0.23 2.06 *
2: 4 CR NPK I7 7.70 020x 2.02 x
22 7 CR NPK I7 7.70 0.3 I? 2.60 x
x x CR NPK 17 7 65 0.31 I ? 60 x

23 ? M s 25 6 I7 0.09x I .OV x
23,s -2 XI NPK 34 661 0.115 I .05 *
23 7 51 NP I7 5.17 0.1 IO 0.91 x
23 I? XI NPR 25 5.x0 0.125 I.21 x
2.1 20 M Nil 33 5.40 0.690 0.75 x
24 5 hi S I6 5.92 0.101 I.31
24 6 M NPK I6 5.90 0.096 I.?? x
24.7 M NPK I6 6.10 0 092 I.?? x
2-l 9 M NPK 16 5.95 0.111 I 33 x
24 IO M NPli I6 6 2fi n.inx I.30 x

‘CR. crop rol.tlion: M. mono,cul~ure: FYM. farm yard m.anurc;NPK. nmogcn. phmphe. pofasium; S. SITBW:
G. preen m.l”“re.
“Ekforc M or CR mm;lpemcn~ commenced plots had heen under agrxultural use.
 Metabolic quotients and soil microbial biomasses 253

Determination of qD Table 3. ANOVA for the aC0.

For measuring qD, the initial microbial biomass of Signdicancc

ElkI Mean square F level
28 soils (I4 M and I4 CR) was determined. The
Cropping practicfl I.&151 31.74 .*.
soils were held for 5 weeks at 22’C in the dark.
Fertilizer O.BlQ 2.03 0. I?8
C, was recorded every week. The data are based Soil type 0.2059 2.05 0.115
on arithmetic means of five replicate soil samples Previous crop 0.3JI5 0.29 0.884
(50 g) per weekly measurement. Periodically, the Clay COnlCnlb 0.1093 0.95 0.555
moisture content was adjusted with distilled Hz0 C 0. I?07 3.03 0.109
tJl? 0.0X6 0.20 0.655
to maintain the initial water potential of
ca - 390 kPa. ‘M or CR.
bBascd on data from plots acre clay content was available (n = 29).
The C-loss quotient (unit C-loss unit-’ l**Slgniticant at 99.9% probability.
C m,c.m,duul
h-‘) was calculated in two ways,
(a) based on microbial-C-loss determined week by fertilizer. previous crop cover, soil type, % clay, C,,
week or. or pH (Table 3). However, the variances observe d
(b) based on total microbial-C-loss after 5 weeks of could be correlated with the number of years the plots
incubation using the following equation were under continuous management (Fig. I), since
(ID = I(G),, - (C,,,),:/(C,,,),,l/t: - Ii. the younger the plots were with respect to a particular
management. the higher was the observed qC0,. This
can be demonstrated by plotting qC0: against
Soil C and N &termination the age of the plots. As depicted in Fig. I, this
Percent soil Cnrg was estimated after removing interaction can be described by an exponential
inorganic C (Nelson and Sommers, 1982). Carbon- function (_v= aeha). This relationship is best shown
ate-C was removed with 10% HCI and the sample by the CR system. where a sufficient number of
subscqucntly dried on a sandbath at 70 C and % CorC years of identical plot management is covered
determined by dry combustion (inductive furnace, (Fig. la). An even better relationship between qC02
Lcco). For total soil N, the N-Kjeldahl method was and age of plot is evident if the number of years of
used (Hrcmncr and Mulvancy. 1982). C and N values continuous plot USC arc clustcrcd in I5 yr intervals
arc means of triplicate samples. and subscqucntly. a nrccm c/CO: for cvcry interval is
calculated. In pcncral. data points diffcrcd signi-
ficantly from each other at P = 0.05 and bcttcr
RWt:I,TS (Fig. lb).

To assess the size of this quotient and the postu-

latcd diHl?rsnccs as outlined in the Introduction the
amounts of microbial biomasscs of soils of long-term
M and continuous CR wcrc dctcrmincd and CO?
rates of their basal respiration rccordcd accordingly.
The n~o/r unit CO,-C which evolved per unit C,,, h-l
was signiticantly higher from microbial biomasscs of
plots undsr pcrmancnt M as compared to microbial
biomasscs under continuous CR (Table 2). Although
the obscrvcd variation within each group is large. the
groups dill&cd significantly at P = 0.001, using stan-
5 0 II 20 40 60 60 100 120

L
dard error (SE) or standard deviation (SD) as a basis E

for calculation.
(b)
In addition to the t-test of Table 2, an analysis of n-6
variance showed a highly significant effect on the T
qC0, by the cropping practice (M vs CR) while no
influcncc was seen with variables such as type of

Table ?. Comparison bc~wccn the metabolic quotient qC0: of

mxrobial biomarws rrom long-term M and CR
Mean

TYpc or YCD:
ma”;lgcmc”t (mg CO,Cmg ’ C,, h ‘1 x IO ’ Range
hl 10.97*** 6.0-17.0 ~g6 of long-term plots in years
SE O.hH’
SD 2.X n = 17 Fig. I. Relationship between age of long-term plots and
CR 6.45 1.7-12.0 amount of CO, evolved per microbial biomass and time
SE 0.45 (yC0.X). (a) The regression is based on qCO:-C to age of
SD 2.0 n - I9 indivibual plots or(b) on an average qC02-C of age of plots
l*‘Sign&xitlY ddkrcnt JI the level P = 0.001 (f-test). in ISyr intervals. Bars indicate deviations from the mean
‘SE. standard error: SD. standard deviation. (SE).
254 ANDERSON
TRAUTE-HEIDI and K. H. D~MXH

Comparison of microbial-C-loss quotients qD (qCO,-C) of those plots as described above

If the amount of available C falls below the (Table 2).
maintenance requirement of a microbial community,
the living biomass must decline. Microbial-C-loss was DISCL’SSION
followed during 5 weeks. For better comparison and
assessment of C-loss as a physiological constant. the Comparison of performances of different microbial
form of presentation of C-loss was expressed using biomasses or the study of environmental influences
Pirt’s equation for metabolic rate constants (Pirt, on any performance seems most useful when activity
1975) (see Materials and Methods). is expressed in terms of physiological constants. The
The quotients of C-losses at t2 when based on the data on qC0, or q D of soil microbial biomasses can
previous or residual biomass at t, are shown in be related to few investigations only. Earlier studies
Table 4. The calculated q values are arithmetic means showed the qC0, to be influenced by temperature
of the individual q values of n number of soils. Not (Anderson and Domsch. 1986a) and by the number
all soils which were employed at the start of the of years the plots were under management (Insam
experiments showed C-losses erery week. Indeed. and Domsch, 1988). The qC0, values obtained in
increases of the biomasses were occasionally ob- these studies were similar to those found here.
served. particularly when soils had to be remoistened in general, rate constants of the total biomass
and remixed during the week before measurements to under in situ conditions are always lower (in magni-
correct for water loss. Since these increases were tudes) than comparable values derived from soil
thought to be atypical and due to the redistribution organisms studied under optimal in vitro conditions
of nutrients by mixing the soils after watering. the (Anderson and Domsch. 1986a). Anderson and
mean qD per week is based on those soils, where a Domsch (1978) showed that the qC02 quotient
dcfinitc dccrcase was observed with respect to the changed according to the age of the organisms in
previous week. It appears that weekly C-loss (qD) culture. They reported a mean yCO,-C of 21 peg
oscillates (Table 4a). CO:--C h-’ mg-’ fungal-C of I4 soil fungi during the
The mean weekly C-loss quotient during 5 weeks early phase of exponential growth on glucose at 22 C.
was 3.0 x IO- h -’ for biomasscs from M systems vs This value decreased to 8.7 during the stationary
1.9 x IO -’ h- ’ for biomasscs from CR systems. The phase (CN I3 days). The latter value approaches
two groups differ significantly at P = 0.01 when quotients reported in our study (Table 2) for the
based on SE (Tahlc 4~1). Another way of expressing dormant microbial biomasscs (range I J-0.16). Simi-
C-loss would bc by rcfcrcncc to the initial biomass lar values of 1.1-0.7 he’ wcrc found in a study on a
only (Table 4h). f Icrc, total C-loss during 5 weeks is stcrilc soil fungus during the early stationary phase
considcrcd. The mean qD is lower when compared (100 h) or late stationary phase (I50 h) at IO C,
with the mean qD based on residual biomasscs on a rcspectivcly (Flanagan and Bunnell. 1976).
weekly basis. A death rate quotient of 4.4 x IO-‘h-’ for the
Soil tcxturc dots not seem to be rcsponsiblc for the total soil microbial biomass was calculated by Smith
diffcrcnccs of qD values seen between M and CR (1982) using data of Ladd and Paul (1973) and
plots. since silt and clay soils were rcpresentcd in both Shields et al. (1973). The quotient cited by Smith is
types of management groups. The higher death rate exactly one magnitude higher than the ones found for
observed for biomasses of M plots coincides with the soil microbial biomasscs in the present investigation
highrr CO:-C production rate per unit biomass (range 3.5-1.0 x IO-‘. Table 4). Smith had based his
calculations on populations which had developed on
glucose, by taking the exponential decline following
the initial peak of growth for estimating qD. In
4D contrast, the qD in this investigation was based on
(mg C-kwsmy ’ Cm,,,,,,h ‘) x IO ’ C-loss of biomasses under more or less continuous
----- -.--
C-deprived conditions. These two values thus stand
M CR
for the two extremes-death after maximal growth
01 3.-w ” 12
= I .9J n = I3 and death during or close to starvation periods. A
I -2 ?.I4 II =9 I.00 n =n
23 3.5.1 n = 9 2.06 n * 9
death rate quotient of 2 x IO-‘h-’ similar to those
3-l I.7d ,I =h ?.d9 n = 8 reported in our study for soil microbial biomasses can
d5 d.IS n =x I.HY n-x be calculated from data presented by Van Vccn et al.
(1985). The death rate constants of IO and
Mc.ln I 3.01.. I .xx
SE O.d6 n = 5 SE 0.25 n = 5
20 x IO-’ h-’ for soil bacteria and soil fungi, respec-
tively, (McGill et al., 1981) appear too high.
It is indeed intriguing that microbial communities
may difTcr with respect to energetic efflcicncy. The
Tahlc J(b). C-loss quol~cnln of microbial observation that microbial biomasses from plots un-
blomasws of ather .M or CR syems aftcr 5
der long-term M produce. as a mean, almost twice as
weeks ;I, 22 C (um~c as in Tahlc da)
much CO, per unit cell mass and time and show a
4D death rate which is ca twice as high as compared to
51 CR microbial biomasses from continuous CR would
2 I?...
I.JI speak for a higher energy demand in the first group.
SEO.11 n=Id SE 0.07 n = Id These yualitatire differences between microbial com-
“‘Signific;mtly different a! P = 0.001: (r-rcsr). munities can only be associated with differences in
SE. SunJ~rd error. potential of species which a certain community is
 Metabolic quotients and soil microbial biomasses 255

composed of. With respect to selective adaptation of in Weizenfeldem. Zeischrijt ftir Pflun:enermihrung und
microbial populations towards certain plant residues, Bodenkunde 119. 134-149.
no conclusive evidence can be found in the literature Flanagan P. W. and Bunnell F. L. (1976) Decomposition
models based on climatic variables. substrate variables,
(Domsch and Gams, 1968; Verstraete and Voets,
microbial respiration and production. In The Role of
1977). although bacteria or fungi will respond to
Terrestrial and Aquatic Organisms in Decomposition Pro-
specific plant residues (Domsch er al.. 1968; Mar- cesses. 17th S_vmposium of the British Ecological Sociely.
tyniuk and Wagner. 1978; Allison and Killham. 1988; pp. 437-457. Blackwell. Oxford.
Fyles ef al.. 1988). On the other hand, changes in Fyles I. H.. Juma N. G. and Robertson L. A. (1988)
enzyme activities due to cropping and fertilizer prac- Dynamics of microbial biomass and fauna1 populations in
tices are very frequently reported, especially increases long-term plots on gray luvisol. Cunodiun /ourno/ of Soil
of urease. phosphatase and dehydrogenase contents. Science 68. 91-100.
with increasing complexity of organic matter input Gostkowska K. and Furczak J. (1983) The effect of the
cultivation system used on the microbiological activity of
(Jsggi, 1974; Klein and Koths. 1980; Gostkowska
loess soil. Polish Journal of Soil Science 16, 39-45.
and Furczak. 1983; Beck. 1985: Bolton er ~1.. 1985).
lnsam H. and Domsch K. H-. (1988) Relationship between
Also, long-term studies by Storozhenko (198-I) showed soil organic carbon and microbial biomass on chrono-
a higher activity of cellulose breakdown in soils under sequences of reclamation sites. Microbial Ecology IS,
continuous CR compared with continuous M. 177-188.
That indeed energetic optimization (Odum. 1969) lnsam H. and Haselwandter K. (1989) Metabolic quotient
may occur on the microbial community level was of the soil microflora in relation to plant succession.
shown by lnsnm and Domsch (1988). The release of Orco/ogiu 79. I74- I 78.
CO: per unit biomass and time was significantly lnsam H., Parkinson D. and Domsch K. H. (1989) Influence
of macroclimate on soil microbial biomass. Soil Biology
higher for microbial biomusses from “young” sites as
& Biochmlisrrv 21 . 211-221.
compared to “matured” ones. In the present study we
Jiggi W. (1974) &)denmikrobiologische Untersuchungen in
could show a similar tcndcncy. einem Diingungsvcrsuch. Schwixrische Lundwirrsrhuft-
Applications of physiological constants may bc an lichc For.whung 13. 53 I -547.
additional way for quantifying the cffccts of cnviron- Klein T. M. and Koths J. S. (19X0) Urcase. protease and
mcntul changes on microbial communities. acid phosphatasc in soils continuously cropped to corn
by conventional or no-tillagc methods. Soil Biokogy &
~~~./~~Io~~~/~,I~~I~~~I~~~I~.v--WC
thank K. StclTcns for expert tcch- Biochcnrisfr~ 12. 2Y3 294.
nical assistance. Our thanks arc extcndcd IO Professor Ladd J. N. and Paul E. A. (1973) Changes in enzymic
T. R. G. Gray for reading and correcting parts of the activity and dislrihulion of acid-soluble. amino-acid
English. This research ws in port fun&d hy the German nitrogen in soil during nitrogen immobilization and
Rcscarch Foundation (DI’G). mincralil;~tion. Soi/ Biohwv & Birwhcw~isrrv 5. 825.-840.
.L

Martyniuk S. and Wagner G. tl. (lY7X) Quantitative and

qu:llitativc examination of soil microflora associated with
dillizrcnl managrmcnl systems. Soil Science 125. 343-350.
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SRB :::--I