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Geoderma, 56 (1993) 527-537 527

Elsevier Science Publishers B.V., Amsterdam

Phosphorus mineralization during laboratory


incubation in soils derived from different
textured parent materials

D. L6pez-Hermindezand M. Nifio
Laboratorio de Estudios Ambientales, IZT. Facultad de Ciencias UCV, Apdo. 47058, Caracas 1041-
A, Venezuela
(Received November 28, 1991; accepted after revision June 10, 1992)

ABSTRACT

L6pez-Hern~mdez, D. and Nifio, M., 1993. Phosphorus mineralization during laboratory incubation
in soils derived from different textured parent materials. In: L. Brussaard and M.J. Kooistra (Edi-
tors), Int. Workshop on Methods of Research on Soil Structure/Soil Biota Interrelationships. Geo-
derma, 56: 527-537.

The dynamics of phosphorus in soils derived from different textured parent materials were studied.
Soils weathered from the three parent materials were Haploborolls but different in texture and solum
depth. Microbial phosphorus by hexanol fumigation, and mineralizationrates of organic P (Po) were
assessed with a 32p dilution technique and P was fractionated using a modification of the method of
Hedley et al.
Microbial P and Po mineralization rates were significantly affected by parent materials. The high
rates of mineralization for the fine-textured soils correspond well with their higher content in sub-
strates able to mineralize e.g. total carbon, Po and microbial Pi. The residual P (e.g. more stable Pi
and Po forms) is the principal P fraction in the analyzed Mollisols.

INTRODUCTION

The amounts and forms of soil phosphorus (P) vary with soil-forming fac-
tors (Walker, 1965; Smeck, 1973). Moreover, Walker (1965) emphasized
the role of P as perhaps the key element in pedogenesis because of its great
ecological significance. More recently Cole and Heil (1981 ) have discussed
the influence of P concentration on the tendency of C and N to accumulate
during pedogenesis.
Precipitation inputs of P are generally below 1 kg ha-1 yr-1 in both tem-
perate (Likens et al., 1977) and tropical (L6pez-Hern,'lndez, 1991 ) ecosys-

Correspondence to: D. L6pez-Hern~indez, Laboratorio de Estudios Ambientales, IZT. Facultad


de Ciencias UCV, Apdo 47058, Caracas 104 l-A, Venezuela.

0016-7061/93/$06.00 © 1993 Elsevier Science Publishers B.V. All rights reserved.


528 D. LOPEZ-HERN,~qDEZ AND M. NII~O

tems. Therefore P, one of the macronutrients contained in soil organic mat-


ter, must be supplied almost entirely by the parent materials (Walker, 1965 ).
Phosphorus released in soils from weathering of primary P-bearing min-
erals and additions of plant residues and fertilizers combines primarily with
the clay fraction (Black, 1968; L6pez-Hern~indez and Burnham, 1974). As a
result, the P percentage of the clay fraction usually exceeds that of the coarser
particle size fraction. Moreover, if other condition are similar, the P percent-
age of the soil as a whole usually increases as the texture becomes finer (Walker
and Brown, 1936; O'Halloran et al., 1985, 1987).
The inorganic fractionation of P by Chang and Jackson (1957 ) and later
modifications (Williams et al., 1967 ) has been used to characterize inorganic
P (Pi) and has served as the basis for concepts of P transformation with pe-
dogenesis (Westin and De Brito, 1969; Smeck, 1973 ). The method, however,
does not differentiate organic phosphorus (Po) fractions which are recog-
nized to be very important in quantity and which play a significant role in
plant nutrition. An alternative P fractionation method was developed by
Hedley et al. ( 1982 ) to characterize not only Pi but also labile and more sta-
ble Po in soils under different incubation conditions and management (Hed-
ley et al., 1982; Tiessen et al., 1984; O'Halloranet al., 1987).
The availability and distribution of P have been studied across enviromen-
tal gradients and as a function of topography (Roberts et al., 1985; L6pez-
Hern~indez, 1987). However, little detailed information exists on the distri-
bution of organic and inorganic soil phosphorus as affected by different tex-
tured parent materials. Therefore, the primary purpose of this study was to
determine the influence of different textured parent material on the nature
and distribution of soil P by using a modified Hedley et al. ( 1982 ) sequential
technique. A secondary objetive was to relate Po mineralization rates with the
textural characteristics of the soils.

METHODS

Site description

Soils from the summit position of three toposequences were used in this
study. The sites are located in southwestern North Dakota, and have been the
subject of intensive studies on erosion, organic carbon and nitrogen changes
resulting from cultivation (Aguilar, 1984; Schimel et al., 1985; Schimel, 1986 ).
The toposequences were on three different parent materials with varying tex-
tures (sandstone, siltstone and shale) with grassland as the native plant com-
ponent. The siltstone site has never been fertilized; the other sites have re-
ceived about 10 kg N ha -t yr -t (Schimel et al., 1985) The soil of all three
sites were classified as Typic Haploborolls (Soil Survey Staff, 1975 ). Selected
soil properties are shown in Table 1.
PHOSPHORUS MINERALIZATIONDURING LABORATORYINCUBATION 529

TABLE 1

Selected soil chemical and textural properties of the analyzed soils

Parent % Sand % Silt % Clay % Carbon % Nitrogen Total phosphorus Organic P


material (rag kg- 1) (mg kg- l )

Sandstone 71 16 13 1.58 0.16 554 204


Siltstone 20 56 24 3.23 0.31 685 325
Shale 13 49 38 2.44 0.26 662 290

FieM sampling

Soil samples for incubations were collected in August 1984. Three 7.5 cm
diameter, l 0 cm deep cores were taken from each site. The samples were kept
in a cooler with ice and returned to Fort Collins, Colorado for analysis.

Incubations

Fifty grams < 2 m m samples of dry soil were placed into Nalgene PMP
wide mouth 125 ml jars, wetted to 50% field capacity with distilled water and
incubated with 1 ml of garden soil solution (E.T. Elliott, pers. commun., 1987)
to restore biological activity. The wetter samples were left to incubate for 4
days. After that period, 2 ml 32p-labelled 10-5M KH2PO4 carrier with an ac-
tivity of approximately 40/zCi m l - l was added to the soil in order to obtain
80% of field capacity and the system was mixed thoroughly with a glass rod
to give a distribution of 32p in the soil as uniform as possible. The labelled
soils were maintained in an incubation chamber designed to measure CO2-C
evolution and at a constant room temperature (25 °C). Three aliquots from
each sample were destructively sampled at days 0, 2, 7, 14, 21 and 51. On
each sampling day the individual systems were thoroughly mixed to distrib-
ute the 32p evenly and sub-samples were taken for subsequent phosphorus,
activity and water contents analysis.

Isotope analysis

On each sampling date, triplicate samples from each site were taken in 100
ml centrifuge tubes and shaken for 1 h in a reciprocal shaker with 40 ml of de-
ionized water plus 0.5 g of bagged anion exchange resin (Dowex 1 * 80-50
> 30 mesh) in a bicarbonate form.
After the shaking period the resin bag was removed and Pi was recovered
by exchanging with 30 ml ofa 0.5M HC1 solution for 1 h. Aliquots of the resin
recovered were analyzed for Pi by Murphy and Riley (1962) reagent and for
32pi by Cerenkov liquid scintillation counting. Isotopic exchangeable phos-
530 D. LOPEZ-HERN,~iJqDEZ AND M. NIIT/O

sol.dE1 (a)

•-e,~ SAMPLE A 2 . 0 g TRIPLICATE SAMPLES


ADD Imt HEXANOL. EVAPORATE HEXANOL OVERNIGHT
SHAKE Ih IN 40rn~. DEIONIZED WATER WITH O.4g
BAGGED DOWEX ANION RESIN IN BICARBONATE FORM"-~'RESIN Pi HEXANOL
REMOVE RESIN BAG, RECOVER AND ANALYZE Pi

•-=-'SAMPLE B 2,0,g TRIPLICATE SAMPLES


SHAKE Ih IN 4 0 m [ DEIONIZED WATER WITH O.4g
BAGGED OOWEX ANION RESIN IN BICARBONATE FORM----~RESIN Pi
REMOVE RESIN BAG, RECOVER AND ANALYZE Pi

TO BE CONTINUED WITH P FRACTIONATION (SAMPLE A) ,

[ SAMPLE AJ
(b)

SHAKE 15h IN 4OraL DEIONIZED WATER WITH 0.4g


BAGGED DOWEX ANION RESIN IN BICARBONATE FORM
REMOVE RESIN BAG, RECOVER AND ANALYZE Pi

RESIN Pi 16h

RESIDUE + S U P E R N A T A N T

ADD t.6g NoCL CENTRIFUGE AND DECANT CLEAR SUPERNATANT

FOR Pi AND Po
WATER SOLUBLE

Pi AND Po

RESIDUE

SHAKE 16h IN 4OraL 0.1N NaOH


CENTRIFUGE AND REMOVE SUPERNATANT FOR Pi AND Po
ANALYSIS
NaOH

Pi AND Po

RESIDUE

SHAKE 16h tN 4OraL 0.1N HCL

CENTRIFUGE AND REMOVE SUPERNATANT FOR Pi ANALYSIS


_"-'. HCL Pi

Fig. 1. (a) Microbial Pi determination. Microbial Pi corresponds to Hexanol-Resin Pi minus


Resin P~. (b) Modified Hedley et al. (1982) P fractionation method.
PHOSPHORUS MINERALIZATION DURING LABORATORY INCUBATION 531

phorus (IEP) was determined after decay and humidity corrections at each
sampling date by using the isotope dilution technique (White, 1976) accord-
ing to the formula:
total activity added (cpm g-1 soil)
IEP -
specific activity of resin extract (cpm #g-~ P)
Resins have been shown to adsorb practically only inorganic phosphorus
(Hedley et al., 1982), therefore during the incubation period any increase in
IEP is a direct consequence of the release of Pi by microbial activity, since
exchange of 3zp with inorganic mineral forms appears to contribute little to
IEP in the studied soils (Sweet, 1981 ).
Rates of Po mineralization were calculated from the difference between IEP
values at the start (0 day) and IEP values at a given incubation period.

Phosphorusfractionation
At the last sampling date (day 51 ) a modified version of the Hedley et al.
(1982) fractionation of soil P was performed (Fig. 1). In the procedure, a
sequential extraction removed labile Pi and Po first, followed by the more
stable forms. The most biologically labile Pi was extracted first with anion
exchange resin (Amer et al., 1955 ). A duplicate sample of the soil was treated
with hexanol (McLaughlin et al., 1986) and then extracted with the resin.
The extra Pi and Po extracted (Hexanol-Resin P minus Resin P) corresponds
to part of the microbial P present in the soil (Fig. la). The suspension was
flocculated with 1.6 g of NaCI, the clear solution contains mainly Po desorbed
from the soil surface.
The resin extraction was followed by sodium hydroxide to remove Po as-
sociated with humic compounds (Batsula and Krivonosova, 1973; Fares et
al., 1974; Bowman and Cole, 1978 ) and Pi more strongly bound to Fe and A1
compounds (L6pez-Hern~lndez and Burnham, 1974, Burnham and L6pez-
Hern~indez, 1982). An acid extractant (1M HCI) removed mainly apatite-
type minerals (Williams et al., 1971 ). Residual P was not analyzed, a sepa-
rate total P analysis by NaOH fusion was performed, however.
Total P (Pt) in the neutralized extracts were determined by acidified so-
dium persulfate oxidation (Environmental Protection Agency, 1971 ) and Po
calculated as the difference between Pt and Pi-

RESULTS

Rates of mineralization

During the experimental incubation period, the amount of isotopic-ex-


changeable phosphorus (IEP) increased (Fig. 2 ). At the beginning of the ex-
532 D. LOPEZ-HERN~tNDEZ AND M. NllqO

NORTH DAKOTA SOILS

"--" SANDSTONE
IO0- -- -- SHALE
SfLTSTONE

T
50-
f ~ y
O.

2r I 511

Doys

Fig. 2. Changes in isotope exchangeablephosphorus (IEP) during incubation.

periment (days 0-7 ), the mineralization increased rapidly, particularly in the


case of the shale and siltstone sites, and in a minor extent for the sandstone.
Then the slopes of the curves decrease (days 7-21 ), and finally values be-
come constant until the end of the incubation study (days 21-51 ). A 14 day
period seems to be enough for an important part of the mineralization process
to occur for the sandstone and siltstone, whereas the shale site still showed a
significant increase in IEP values after 14 days of incubation. Assuming an
incubation period of 51 days a low rate of mineralization was found for the
sandstone (0.48 mg P kg-~ day-~, Table 3 ) when compared with the shale
and siltstone with 1.25 and 1.03 mg P kg- i day- ~ mineralized, respectively.
Decay corrected specific activities (SA, data not presented) showed a de-
crease during the incubation period that is a consequence of a flush of 3]pi
released by mineralization.

P fractionation

The shale parent material site gave the highest value in resin Pi ( 7.1 #g g- ] )
as compared with the other two sites (3.9 and 4.5 for sandstone and siltstone,
respectively) (Table 2 ). Microbial Pi was also present in appreciable amounts
in the fine materials ( 14.3 and 10.3 #g g - ~for the siltstone and shale, respec-
tively), whereas the coarse-textured sandstone site yielded a lower value ( 7.3
/zg g- ] ). As expected after resin extraction the amount of P~ remaining in
solution was low (0.63-0.80 gg g - ] ) in all the three sites. On the other hand,
Po remaining in solution was, however, important although the values did not
differ significantly between sites (Table 2 ). Inorganic P extracted with NaOH
was also significantly higher in the siltstone than in the sandstone and shale.
PHOSPHORUSMINERALIZATIONDURINGLABORATORYINCUBATION 533

TABLE 2

Phosphorus fractions (mg kg- t ) from the sequential extraction. Standard deviation in parentheses

Parent Resin Pi Microbial Pi Po


material 16 h Pi water water

Sandstone 3.9 (0.31) 7.6 (1.38) 0.63 (0.05) 9.3 (1.10)


Siltstone 4.5 (1.90) 14.3 (1.6) 0.80 (0.19) 10.8 (0.95)
Shale 7.1 (3.15) 10.3 (6.8) 0.66 (0.19) 9.7 (0.54)

Parent NaOH-Pi NaOH-Po HC1-Pi Sum fraction


material

Sandstone 15.5 (2.87) 44.0 (7.50) 159.1 (12.60) 240.03


Siltstone 20.2 ( 1.80 ) 56.0 ( 1.80 ) 139.6 ( 15.98 ) 246.20
Shale 12.1 (0.69) 43.4 ( 13.33 ) 164.9 (44.00) 248.16

A similar result was found for NaOH extracted Po. HCl-extracted Pi w a s high
for the sandstone and shale sites, and significantly differs from the siltstone
site.
Finally, total P as determined by NaOH fusion as well organic P for the
fine-textured parent material (shale and siltstone) exceeded the value of the
coarse particle-size sandstone (Table 1 ). Substracting the sum of the frac-
tions from total P, residual P is calculated. It appears that residual P which
corresponds to the more stable Pi and Po forms, is the principal P fraction in
all the analyzed Mollisols; residual P fraction was higher in the shale (414
#g-~) when compared with the sandstone and siltstone with 314 and 359/zg
g- 1, respectively.

DISCUSSION

The productivity of many natural and agricultural ecosystems is deter-


mined by the quantity of Pi made available through mineralization during the
growing season. In some agricultural ecosystems, P fertilizer additions are
non-existent or negligible, particularly in the case of many MoUisols in the
Great Plains, where Haas et al. ( 1961 ) measured a significant average reduc-
tion in Po of 35% after 31 to 48 years of cropping, compared with virgin sod.
In this situation, the quantities of Pi coming from mineralization are of par-
amount importance. Most plant species have low P requeriments (Fox, 1981;
Lrpez-Hern~ndez et al., 1987 ). Consequently, only a small proportion of the
Po must be mineralized to provide Pi requeriments.
Higher rates of mineralization were measured in the case of the fine-tex-
tured parent material soils ( 1.25 and 1.03 mg P kg- 1 day- 1, Table 3 ) com-
pared with the sandstone derived soil (0.48 mg P kg- 1 day- ~). These values
534 D. LOPEZ-HERN,/d~DEZAND M. NII~IO

TABLE 3

IEP values at the beginning (To) and 51 days of the incubation period. % Po mineralized at 14 days
as compared with 51 days

Parent IEP at To IEP at 51 days mg P kg- l Rate of % IEP


material (mg P kg- ~) (mg P kg- 1) mineralized mineralization 14 days/51 days
(mg p kg -1 day - l )

Sandstone 5.9 30.3 24.4 0.48 93


Siltstone 9.9 62.2 52.3 1.03 79
Shale 6.7 70.3 63.6 1.25 66

are higher than the theoretical value (0.13 mg kg- 1 day- 1 ) found by Harri-
son (1982a) for a woodland in the Lake District in England. The data, how-
ever, cannot be compared directly because the method used by Harrison
( 1982b ) included a shorter incubation period. Harrison's experiments were
performed using an incubation temperature of 13°C, the experiments re-
ported here were done at a higher temperature (25°C) and perhaps a more
optimal moisture content. Temperature is very well known to affect the rate
of Po mineralization (Thompson and Black, 1947; Ruiz and Lrpez-Hermin-
dez, 1977).
The high rate of mineralization for the fine-textured soils corresponds well
with their higher content in substrates able to mineralize e.g. total carbon,
organic P and microbial Pi (Tables 1 and 2 ).
O'Halloran et al. (1985, 1987) studied P fractionation in a spatial varia-
bility design study. They reported a decrease in resin Pi and NaOH Pi with
increasing sand content. The results presented here are not entirely compa-
rable with O'Halloran's results because of the reduced number of samples
used. The information confirms, however, the results in the case of resin Pi,
but not for NaOH extractable Pi. When comparing the information obtained
for HC1-Pi non significant differences were found amoung the different tex-
tured parent materials. O'Halloran et al. ( 1985 ), on the contrary, reported a
positive correlation for HC1-P~ and sand content.
Schimel (1986) measured net gross N mineralization and CO2 evolution
during the first 4 days of incubation in soils derived from the same parent
materials. He concluded that net N mineralization was similar in all sites;
however, the gross N mineralization, and particularly the CO2 evolution, was
higher in the fine-textured materials than in the coarse-textured sandstone.
The results presented by Schimel (1986) for N and C coincide with the infor-
mation presented here for Po mineralization rates. CO2 evolution curves (to
be discussed in another paper) during a 28 d incubation period resembles
those obtained for isotopic exchangeable P, indicating that the increase in the
exchangeable P fraction (IEP) is a direct consequence of the release of Pi by
microbial activity.
PHOSPHORUS MINERALIZATION DURING LABORATORY INCUBATION 535

CONCLUSIONS

Our results for soils with the same taxonomic clasification (Typic Haplo-
borolls) but differing in textural characteristics show that higher rates of mi-
neralization occurred in the fine-textured parent material soils compared with
the sandstone derived soil. The high rate of mineralization for the fine-tex-
tured soils corresponds well with their higher content in organic P and micro-
bial Pi. The residual P (e.g. stable Pi and Po forms) is the principal P fraction
in the analyzed Haploborolls.

ACKNOWLEDGEMENTS

C.V. Cole and E.T. Elliott contributed to the genesis of this study and pro-
vided the soil samples and laboratory facilities. This research was supported
by NSF-BSR-810528 l, the Great Plains Agroecosystems Project. Work re-
ported in this manuscript was done while the senior author was at NREL,
Fort Collins, CO, on sabbatical leave from Universidad Central de Venezuela.

REFERENCES

Aguilar, R., 1984. Parent material - - Topographic-management control on organic and inor-
ganic nutrients in semiarid soils. PhD. Diss. Colorado State Univ., Fort Collins, CO.
Amer, F., Bouldin, D.R., Black, C.A. and Duke, F.R., 1955. Characterization of soil phosphorus
by anion exchange resin adsorption and 32p equilibration. Plant Soil, 6:391-408.
Batsula, A.A. and Krivonosova, G.M., 1973. Phosphorus in the humic and acid fulvic acids of
some Ukrainian soils. Soviet Soil Sci., 5: 347-350.
Black, C.A. 1968. Soil-Plant Relationships. Wiley, New York, 792 pp.
Bowman, R.A. and Cole, C.V., 1978. Transformation of organic phosphorus substrates in soils
as evaluated by NaHCO3. Soil Sci., 125: 49-54.
Burnham, C.P. and L6pez-Hern~indez, D., 1982, Phosphate retention in different soil taxon-
omic classes. Soil Sci., 134: 376-380.
Chang, S.C. and Jackson, M.L., 1957. Fractionation of soil phosphorus. Soil Sci., 84:133-144.
Cole, C.V. and Heil, R.D., 1981. Phosphorus effects on terrestrial nitrogen cycling. In: F.E.
Clarke and T. Rosswall (Editors), Terrestrial Nitrogen Cycle, Processes, Ecosystems and
Management Impact. Ecol. Bull. (Stockholm), 33: 363-374.
Environmental Protection Agency, 1971. Methods of chemical analysis of water and wastes.
Environmental Protection Agency, Cincinnati, OH.
Fares, F., Fardeau, J.C. and Jacquin, F., 1974. Etude quantitative du phosphore organique dans
different type du sol. Phosphore Agric., 63: 25-41.
Fox, R.L., 1981. External phosphorus requirements of crop. In" Chemistry in the Soil Environ-
ment. Am. Soc. Agron., pp. 223-239.
Haas, H.J., Grunes, D.L. and Riechman, G.A., 1961. Phosphorus changes in Great Plains soils
as influenced by cropping and manure application. Soil Sci. Am. Proc., 25:214-218.
Harrison, A.F., 1982a. 32p method to compare rates of mineralization of labile organic phos-
phorus in woodland soils. Soil Biol. Biochem., 14: 337-341.
536 D. LOPEZ-HERNANDEZ AND M. NllqO

Harrison, A.F., 1982b. Labile organic phosphorus mineralization in relationships to soil prop-
erties. Soil Biol. Biochem., 14:343-351.
Hedley, M.J., Stewart, J.W.B. and Chauhan, B.S., 1982. Changes in inorganic soil phosphorus
fractions induced by cultivation practices and by laboratory incubations. Soil Sci. Soc. Am.
J., 46: 1970-1976.
Likens, G.E., Bormann, F.H., Pierce, R.S., Eaton, J.S. and Johnson, N.M., 1977. Biochemistry
of a Forested Ecosystem. Springer, New York.
L6pez-Hern~ndez, D., 1987. Phosphate adsorption variability within soil series and in diverse
soil populations. Soil. Sci., 144:408-411.
L6pez-Hernandez, D., 1991. Phosphorus dynamic in a flooded savanna. In: H. Tiessen, D. Lo-
pez-Hern~ndez and I. Salcedo (Editors), Phosphorus Cycles in Terrestrial and Aquatic Eco-
systems. Regional Workshop 3: South and Central America. Sask. Inst. of Pedology. Univ.
of Saskatchewan, Saskatoon, Sask., pp. 95-105.
L6pez-Hern~indez, D. and Burnham, C.P., 1974. The covariance of phosphate sorption with
other soil properties in some British and tropical soils. J. Soil Sci., 25: 196-206.
Lopez-Hernandez, D., Coronel, I. and Alvarez, L., 1987. The external phosphate requirement
of cowpea on five dissimilar soils. Soil Sci., 144: 339-343.
McLaughlin, M.J., Alston, A.M. and Martin, J.K., 1986. Measurement of phosphorus in the soil
microbial biomass: a modified procedure for field soils. Soil Biol. Biochem.. 18: 437-443.
Murphy, J. and Riley, J.P., 1962. A modified single solution method for the determination of
phosphate in natural waters. Anal. Chim. Acta, 27:31-36.
O'Halloran, I.P., Kachanoski, R.G. and Stewart, J.W.B., 1985. Spatial variability of soil phos-
phorus as influenced by soil texture and management. Can. J. Soil Sci., 65: 475-487.
O'Halloran, I.P., Kachanoski, R.G. and Stewart, J.W.B., 1987. Influences of texture and man-
agement practices on the forms and distribution of soil phosphorus. Can. J. Soil Sci., 67:
147-163.
Roberts, T.L., Stewart, J.W.B. and Bettany, J.R., 1985. The influence of topography on the
distribution of organic and inorganic soil phosphorus across a narrow environmental gra-
dient. Can. J. Soil Sci., 65:651-665.
Ruiz, J. and L6pez-Hern~indez, D., 1977. Tasa de mineralizacion de f6sforo org~inico en suelos
tropicales en relaci6n con el contenido de humedad y la temperatura. In: Syrup. on Soil Or-
ganic Matter Studies. IAEA and FAO, Braunschweich IAEA-SM-211/68 1, pp. 255-261.
Schimel, D.C,, 1986. Carbon and nitrogen turnover in adjacent grassland and cropland ecosys-
tems. Biogeochemistry, 2: 324-357.
Schimel, D.S., Coleman, D.C. and Horton, K.A., 1985. Soil organic matter dynamic in paired
rangeland and cropland toposequences in North Dakota. Geoderma, 36: 201-214.
Smeck, N.E., 1973. Phosphorus: an indicator of pedogenic weathering processes. Soil Sci., 115:
199-206.
Soil Survey Staff, 1975. Soil Taxonomy. A Basic System of Soil Classification for Making and
Interpreting Soil Surveys. Agric. Handbook No. 436. USDA Soil Conservation Service.
Sweet, J.J., 1981. The mineralization of organic phosphorus in soils as determined by isotope
dilution. M.Sc. Thesis, Colorado State Univ., Fort Collins, CO.
Thompson, L.M. and Black, C.A., 1947. The effect of temperature on the mineralization of soil
organic phosphorus. Soil Sci. Soc. Am. Proc., 12: 323-326.
Tiessen, H., Stewart, J.W.B. and Cole, C.V., 1984. Pathways of phosphorus transformation in
soils of differing pedogenesis. Soil Sci. Soc. Am. J,, 48: 853-858.
Walker, R.H. and Brown, P.E., 1936. The phosphorus, nitrogen and carbon content of Iowa
soils. In: P.E. Brown (Editor), Soils of Iowa. Iowa Agric. Exp. Sta. Spec. Rep. 3.
Walker, T.W., 1965. The significance of phosphorus in pedogenesis. In: D.S. Hallsworth and
D.V. Crawford (Editors), Experimental Pedology. Butterworths, London, pp. 295-315.
PHOSPHORUS MINERALIZATION DURING LABORATORY INCUBATION 537

Westin, F.C. and De Brito, J., 1969. Phosphorus fractions of some Venezuelan soils as related
to their stages of weathering. Soil Sci., 107:194-202.
White, R.E., 1976. Concepts and methods in the measurement of isotopically exchangeable
phosphate in soil. Phosph. Agric., 67: 9-16.
Williams, J.D.H., Syers, J.K. and Walker, T.W., 1967. Fractionation of soil inorganic phosphate
by a modification of Chang and Jackson's procedure. Soil Sci. Soc. Am. Proc., 31: 736-739.
Williams, J.D.H., Syers, J.K., Harris, R.F. and Armstrong, D.E., 1971. Fractionation of inor-
ganic phosphate in calcareous lake sediments. Soil Sci. Soc. Am. Proc., 35: 250-255.

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