Plant and Soil 256: 115–130, 2003.

© 2003 Kluwer Academic Publishers. Printed in the Netherlands.

115

Effects of plant species on phosphorus availability in a range of grassland

soils

C. R. Chen1,4,5 , L. M. Condron1 , S. Sinaj2 , M. R. Davis3 , R. R. Sherlock1 & E. Frossard2

1 Soil, Plant and Ecological Sciences Division, PO Box 84, Lincoln University, Canterbury, New Zealand. 2 Institute

of Plant Sciences, Swiss Federal Institute of Technology Zürich (ETHZ), Postfach 185, CH-8315, Eschikon-Lindau,
Switzerland. 3 New Zealand Forest Research Institute, PO Box 29237, Fendalton, Christchurch, New Zealand.
4 Co-operative Research Centre for Sustainable Production Forestry, Faculty of Environmental Sciences, Griffith

University, Nathan, Qld 4111, Australia. 5 Corresponding author∗

Received 22 November 2001. Accepted in revised form 23 April 2003

Key words: isotopic exchange kinetics, Lolium perenne, 33 P, phosphorus uptake, Pinus radiata, root phosphatase
activity, soil phosphorus availability

Abstract
Vegetative conversion from grass to forest may influence soil nutrient dynamics and availability. A short-term (40
weeks) glasshouse experiment was carried out to investigate the impacts of ryegrass (Lolium perenne) and radiata
pine (Pinus radiata) on soil phosphorus (P) availability in 15 grassland soils collected across New Zealand using
33 P isotopic exchange kinetics (IEK) and chemical extraction methods. Results from this study showed that radiata

pine took up more P (4.5–33.5 mg P pot−1 ) than ryegrass (1.1–15.6 mg pot−1 ) from the soil except in the Temuka
soil in which the level of available P (e.g., E1min Pi , bicarbonate extractable Pi ) was very high. Radiata pine tended
to be better able to access different forms of soil P, compared with ryegrass. There were no significant differences
in the level of water soluble P (Cp, intensity factor) between soils under ryegrass and radiata pine, but the levels
of Cp were generally lower compared with original soils due to plant uptake. The growth of both ryegrass and
radiata pine resulted in the redistribution of soil P from the slowly exchangeable Pi pool (E>10m Pi , reduced by
31.8% on the average) to the rapidly exchangeable Pi (E1min−1d Pi , E1d−10m Pi ) pools in most soils. The values
of R/r1 (the capacity factor) were also generally greater in most soils under radiata pine compared with ryegrass.
Specific P mineralisation rates were significantly greater for soils under radiata pine (8.4–21.9%) compared with
ryegrass (0.5–10.8%), indicating that the growth of radiata pine enhanced mineralisation of soil organic P. This
may partly be ascribed to greater root phosphatase activity for radiata pine than for ryegrass. Plant species × soil
type interactions for most soil variables measured indicate that the impacts of plant species on soil P dynamics was
strongly influenced by soil properties.

Introduction factors including soil properties and parent material,

It is widely acknowledged that continued improve-
ment in our understanding of soil P dynamics and
availability is critical for sound agronomic and en-
vironmental management of agricultural and forest
ecosystems (Frossard et al., 2000). Soil P dynamics
and availability is determined by a combination of
∗ FAX No: +61-7-38757459.
E-mail: chengrong.chen@mailbox.gu.edu.au
type of vegetation, environmental conditions and land
management practices (e.g., Krämer and Green, 1999;
Oberson et al., 1996).
Plant species differ in their ability to obtain P
from soils, and adopt different physiological mech-
anisms and strategies for P acquisition (Lajtha and
Harrison, 1995). Plants also exert significant effects on
soil P availability and dynamics through litterfall, root
turnover and exudation, and specific interactions with
microbes in the rhizosphere (e.g., Attiwill and Adams,
116
1993; Lajtha and Harrison, 1995). This is important
when considering the effects of land-use change on
soil P properties and processes. In New Zealand the
impacts of recent widespread land-use change from
grassland to plantation forestry (predominately radiata
and other pine species) on soil fertility and nutrient
dynamics have been the subject of extensive invest-
igation. Results of these investigations indicated that
afforestation of grassland increased concentrations of
available P, while causing concomitant decreases in
concentrations of soil organic P (e.g., Chen et al.,
2000; Davis and Lang, 1991). However, there is little
information available about how plant species affect P
availability in different soils.
In addition, chemical extraction techniques were
used in most of above studies for determination of soil
P of different availability. However, chemical reagents
such as sodium bicarbonate (NaHCO3 ) and sodium
hydroxide (NaOH) used in these methods remove only
a fraction of available P in soil together with signific-
ant amounts of unavailable P and can therefore only
provide an approximate measure of potentially avail-
able P (Fardeau, 1996; Kato et al., 1995). It should
be noted that interpretation of results is limited by
uncertainty associated with the relationship between
the chemical solubility of soil P and its bioavailability
(Frossard et al., 2000). Moreover, chemical extraction
methods can only provide the quantity factor of avail-
able soil P. According to Beckett and White (1964),
soil P availability is governed by a combination of
intensity, quantity and capacity factors. Isotopic ex-
change kinetics (IEK) utilizing 32 P or 33 P as a tracer
can be used to describe these three factors, and thus
provides an alternative means of characterizing soil P
availability with minimum chemical modification of
the forms of P present, and has been used extensively
to assess inorganic P availability and associated bio-
logical P dynamics in a variety of soils (e.g., Chen et
al., 2003; Oehl et al., 2001a, b; Sinaj et al., 2001).
The main objective of the present study was to invest-
igate the effect of two contrasting plant species on the
nature and availability of P in a range of New Zealand
grassland soils as characterized by chemical extraction
and IEK methods.
Materials and Methods
Soils
Fifteen surface soil samples (0–7.5 cm) that were
originally developed under native vegetation (mainly
evergreen forest) prior to European settlement of New
Zealand and were subsequently converted into grass-
land were collected from around New Zealand. These
included soils from the North Island (Te Kauwhata,
Oruanui, Taupo, Stratford, Egmont, Patoka, Hi-
matangi, Mangamahu) and the South Island (Mapua,
Richmond, Hurunui, Okarito, Temuka, Pukaki, Fork),
and encompassed a range of soil types (Table 1). The
pH values of the soils were generally lower than 6,
except for the Himatangi (pH 7.0) and Temuka (pH
6.5) soils. Total organic C concentrations ranged from
19.5 to 130.4 g kg−1 , total N from 1.0 to 8.5 g kg−1 ,
and total P from 375 to 2607 mg kg−1. Organic P (Po)
comprised 35–81% of total P in the soils. Dithionite
extractable Fe and Al ranged from 1.0 to 19.3 and 1.2
to 12.2 g kg−1 , while oxalate extractable Fe and Al
ranged from 0.5 to 7.4 and 1.0 to 15.8 g kg−1 , re-
spectively. Clay content ranged from 40 to 310 g kg−1 ,
while P sorption index (PSI) ranged from 3.1 to 59.4
(mg 100 g−1 )/(µmol L−1 ).
Soil samples were air-dried and passed through a
4-mm sieve prior to the glasshouse experiment. Separ-
ate air-dried subsamples of each soil were ground and
passed through a 2-mm and a 150-µm sieve prior to
chemical and physical analyses as described below.
Glasshouse experiment
Each of the soil samples was weighed into six small
pots (80 × 80 × 120 mm). Amounts of each soil
used are shown in the Table 1. Two plant species,
radiata pine (Pinus radiata – breed ‘GF12’ and per-
ennial ryegrass (Lolium perenne - cultivar ‘Grasslands
Nui’), were used in this glasshouse experiment. Seeds
were directly sown in the pot and plants of each spe-
cies were thinned to five per pot after germination.
There were three replicates for each soil and each plant
species (total 90 pots). All pots were placed on a capil-
lary mat in the glasshouse in a completely randomized
design, and soil moisture contents were maintained
at ca. 70% field capacity using an autowater sys-
tem (GARDENA 1030). Radiata pine seedlings were
inoculated with mycorrhizae (Rhizopogon rubescens
Tul.) at a rate of 1 × 107 spores per pot, applied in a
water suspension, 2 weeks after sowing. No nutrients
Table 1. Chemical and physical properties determined for the selected grassland soils before planting

Soil series USDA soil pH Organic C Total N Total P Organic P C/P Dithionite Oxalate Sand Silt Clay P sorption Amounts of
classification (g kg−1 ) (g kg−1 ) (mg kg−1 ) (mg kg−1 )a (g kg−1 ) (g kg−1 ) (g kg−1 ) (g kg−1 ) (g kg−1 ) index soil used
Fe Al Fe Al (PSI) (g pot−1 )b
(mg 100 g−1 ) /
(µmol L−1 )

1 Mapua Dystrochrept 5.2 19.5 1.0 116 82 (71) 168 7.5 1.2 1.1 1.0 410 380 210 17.5 278.8
2 Mangamahu Dystrochrept 5.3 39.3 3.8 401 315 (78) 98 9.8 3.4 6.3 2.5 420 400 180 32.0 244.3
3 Okarito Orthod 5.4 130.4 7.3 597 479 (80) 218 1.0 1.7 0.5 1.3 520 380 100 7.2 168.0
4 Egmont Udand 5.7 53.0 4.7 629 485 (77) 84 17.3 6.2 4.4 6.3 400 400 200 34.5 226.2
5 Pukaki Dystrochrept 5.2 49.1 3.1 663 534 (81) 74 5.4 2.4 2.6 2.8 600 280 120 20.8 202.8
6 Fork Dystrochrept 6.0 25.2 2.0 690 242 (35) 37 3.8 2.4 1.6 2.1 640 240 120 11.7 303.4
7 Richmond Dystrochrept 5.7 36.5 3.3 813 568 (70) 45 11.8 3.2 3.3 2.7 290 400 310 15.6 274.1
8 Himatangi Ustochrept 7.0 29.0 3.0 862 331 (38) 34 3.6 1.6 2.4 1.0 840 110 50 3.1 289.5
9 Hurunui Dystrochrept 5.6 78.7 6.7 904 670 (74) 87 12.1 2.7 2.5 1.9 350 420 230 14.2 211.9
10 Te Kauwhata Humult 5.7 50.0 4.8 938 606 (66) 53 19.3 4.1 5.9 2.9 360 380 260 18.8 212.6
11 Temuka Aquept 6.5 39.6 3.8 1056 509 (48) 38 4.1 2.0 2.8 1.2 390 400 210 3.7 240.0
12 Oruanui Vitrand 5.3 88.9 6.2 1127 701 (62) 79 4.2 5.5 2.8 6.7 710 240 50 44.0 161.6
13 Taupo Vitrand 5.1 55.4 4.4 1311 479 (37) 42 3.3 3.6 2.4 4.9 800 160 40 24.1 187.4
14 Patoka Udand 5.7 93.4 8.5 1585 1051 (66) 59 8.5 12.2 4.7 15.8 590 290 120 59.4 170.0
15 Stratford Udand 5.4 67.8 7.0 2746 1166 (43) 25 9.0 7.9 7.4 14.0 620 270 110 48.2 216.7
Mean 5.7 57.1 4.6 963 548 (62) 76 8.0 4.0 3.4 4.4 530 320 150 23.6 225.8

a Data in the brackets are percentage of soil organic P over total P. b The amounts of each soil used in the experiment.
117
118
were added to the pots during the experiment. The av-
erage daily temperature during the experiment ranged
from 12 to 25 ◦ C.
Plants were harvested after 40 weeks and separated
into root and shoot components. Approximately 1 g of
fresh fine roots was sub-sampled from three replicates
of each plant species for determination of root phos-
phatase activity. The shoot material and the remainder
of the root material were oven-dried at 65 ◦ C for 72
h and weighed. Plant samples were finely ground (<
150 µm) prior to analysis.
Fresh soil samples from each pot were thoroughly
mixed and a subsample was taken and stored at
4 ◦ C until analysis of resin extractable P and micro-
bial properties. The remaining soil was air-dried and
ground (< 2 mm and < 150 µm) prior to other chem-
ical analysis. All soil analyses were carried out in
duplicate. The results were expressed on an oven-dry
basis.
Soil analysis
Selected chemical properties of the original soils (pH,
dithionite and oxalate extractable iron (Fe) and alu-
minium (Al), particle size distribution, phosphate
sorption index) and all soils (total organic carbon
(TOC), total nitrogen (N)) were determined according
to the method described by Chen et al. (2003). The
total P, inorganic P (TPi) and organic P (TPo) contents
of soil were determined according to the Saunders and
Williams (1955) ignition method. Sodium bicarbonate
extractable Pi and Po (BPi and BPo ) were determined
according to the method outlined by Tiessen and Moir
(1993), while macroporous resin (Lewatit MP500A)
extractable Pi and Po were determined according to the
method described by Rubæk and Sibbesen (1993).
Isotopic exchange kinetics was carried out on <
2-mm sieved soil (both before and after planting) ac-
cording to the methodology described by Frossard and
Sinaj (1997). This involved shaking 10 g of soil with
99 mL of deionised water for 16 h prior to the addi-
tion of 1 mL of solution containing about 0.1 MBq
of carrier-free 33 P (Product BF 1003, Amersham–
Pharmacia Biotech UK Limited) with continuous agit-
ation on a magnetic stirrer. Solution samples were
removed from the soil/water suspension after 1, 10,
30, and 60 min and filtered immediately < 0.20 µm
prior to determination of 31 P by malachite green col-
orimetry (Sinaj et al., 1998) and 33 P by scintillation
counting (1 mL + 5 mL scintillation cocktail (Packard
Ultima Gold)). When 33 PO4 ions are added to a soil
solution system at a steady-state equilibrium, the ra-
dioactivity in solution decreases with time according
to the following equation:
r(t ) /R = {r(1) /R} ∗ {t + [r(1)/R]1/n }−n + r(∞) /R,
(1)
∼ 0.1
where R is the total introduced radioactivity (=
MBq); r(1) and r(∞) are the radioactivity (MBq) re-
maining in the solution after 1 minute and infinite
time, respectively, and n is a parameter describing the
rate of disappearance of the radioactive tracer from the
solution after 1 min. The parameter n is calculated
as the factor (slope) of the linear regression between
log[r(t )/R] and log(t). The ratio r(∞) /R, which is the
maximum possible dilution of the isotope, is opera-
tionally approximated by the ratio of the water soluble
P to the total soil inorganic P (PT , expressed in mg P
kg−1 soil). Thus:
r(∞) /R = 10 ∗ CP /PT , (2)
where CP is the water soluble P (mg P L−1 ). The
factor 10 arises from the fact that, during an isotopic
exchange experiment, the soil:solution ratio is 1:10 so
that 10*CP is equivalent to the water soluble P quantity
in the soil expressed in mg kg−1.
The quantity, E(t ) (mg P kg−1 soil), of isotopically
exchangeable P at a time t can be calculated assuming
that (i) 31 PO4 and 33 PO4 ions have the same fate in
the system, and (ii) whatever the time, t, the specific
activity of the phosphate ions in soil solution is similar
to that of isotopically exchanged phosphate ions in the
whole system:
r(t ) /(10 ∗ CP ) = R/E(t ) (3)
Therefore,
E(t ) = 10 ∗ CP ∗ R/r(t ) (4)
For t = 1 min,
E1min = 10 ∗ CP ∗ (R/r1 ) (5)
R/r1 is the ratio of the total introduced radioactivity
to the radioactivity remaining in the solution after 1
min and is considered as an estimation of the P-ions
buffering capacity of soils (Frossard et al., 1993).
Isotopic exchange kinetics provided data for the
intensity factor (CP , water soluble P), capacity factors
(R/r1 and n) and the quantity factor (E values, iso-
topically exchangeable soil P). The quantity factor is

presented by a pluricompartmental model (Frossard
and Sinaj, 1997) and thus soil Pi pool of differing
mobility, as calculated according to Equations (1)–(5)
described previously, includes: (a) Pi isotopically ex-
changeable within 1 min (E1min), (b) Pi isotopically
exchangeable between 1 min and 1 day (E1min−1d),
(c) Pi isotopically exchangeable between 1 day and
10 months (E1d−10m), and (d) Pi which cannot be
exchanged within 10 months (E>10m ).
Root phosphatase activity
Root phosphatase activity was measured by the modi-
fied method of Alexander and Hardy (1981) and Dodd
et al. (1987). A preliminary experiment was carried
out using a series of buffers ranging from pH 3 to
12 (3–7 citrate buffer; 7–12 Tris buffer) to determine
the optimum pH for root phosphatase activity of each
plant species. Results showed that optimum pH for
root surface phosphatase activity was 4.5 for ryegrass
and 3.5 for radiata pine (data not presented). All sub-
sequent measurements of root phosphatase activities
were made at the determined optima. In brief, 200 mg
of fine roots (< 0.5 mm diameter) were incubated for 1
h at 25 ◦ C in 1 mL of 50 mM p-nitrophenyl phosphate
and 4 mL of 1.0 mM sodium citrate buffer. Roots were
removed and 5 mL of 0.5 M NaOH were added to the
solution and the absorbance read at 400 nm.
Statistical analysis
A two-way ANOVA was carried out using Genstat
4.2 (Lawes Agricultural Trust, Rothamsted, UK) on
the data (root and shoot biomass and P uptake, and
soil chemical variables) to test significant effects of
plant species and soil type. The least significant dif-
ference (LSD) test was used to separate differences
between species for individual soils when the species
× soils interaction was significant. Multiple regres-
sion analysis was also carried out using Genstat 4.2;
the stepwise estimation procedure was used and only
the independent variables with significant correlation
with the dependent variable were fitted into the final
regression equation (Hair et al., 1995).
Results
Plant growth and P uptake
Main effects of plant species and soil type, and the
interactions between plant species and soil type for
119
dry mass of root, shoot and whole plant and shoot:
root (S:R) ratio were all significant. However, effects
of the soil type and plant × soil interactions were small
compared with main effects for plant species (Table 2).
The dry matter (root, shoot, total) of radiata pine har-
vested was 1.4–5.5 times higher than that of ryegrass
from the 15 soils (Table 2). Shoot:root ratio of radiata
pine was also generally higher than that of ryegrass,
although significant differences were observed only
in Mangamahu, Pukaki, Fork, Himatangi, Hurunui,
Oruanui soils (Table 2).
Phosphorus concentrations in the roots of radiata
pine were significantly higher than those of ryegrass
except for the Himatangi soil, while P concentrations
in shoots of radiata pine were significantly lower than
ryegrass except in the Egmont, Taupo and Pukaki soils
(Table 3). Radiata pine generally took up more P (4.5–
33.5 mg P pot−1 ) than ryegrass (1.1–15.6 mg P pot−1 )
(Table 3). It should be noted that the P concentration in
the shoots of ryegrass in the Temuka soil was 3.5 times
higher than that of the radiata pine and so P uptake
by the ryegrass was greater than radiata pine in the
Temuka soil (Table 3). These differences contributed
to significant plant × soil interactions.
Soil pH, total organic C and total N
Soil pH, TOC and total N concentrations were gener-
ally lower after 40 weeks under radiata pine compared
with ryegrass (Table 4).
Chemically extractable soil P
Significant plant species and soil type and plant ×
soil interactions were apparent for all the extractable P
fractions determined (Table 5). Concentrations of BPi
were significantly higher under radiata pine than under
ryegrass for Okarito, Pukaki, Richmond, Himatangi,
Hurunui, Temuka, Oruanui, Taupo, and Stratford soils.
Concentrations of BPo were significantly higher in
Mangamahu, Egmont, Hurunui, Patoka and Strat-
ford soils under radiata pine compared with ryegrass
(Table 5). Similarly, concentrations of resin extract-
able Pi and Po were also generally higher in most soils
under radiata pine compared with ryegrass (Table 5).
After 40 weeks, total P was significantly lower in
most soils under radiata pine than under ryegrass, but
significantly higher in Temuka soil under radiata pine
(Table 6). Compared with the original soils, total P
decreased by 1.7–12.6% in soils under ryegrass and
by 4.8–16.9% in soils under radiata pine. Concentra-
120
Table 2. Root, shoot and total biomass (dry weight, g pot−1 ), and shoot: root ratios (S:R) of ryegrass and
radiata pine determined after a 40-week period of growth a

Soil Root Shoot Total S:R ratio

Grass Pine Grass Pine Grass Pine Grass Pine

Mapua 0.28 1.52‡ 0.72 3.95‡ 1.00 5.47‡ 2.71 2.60

Mangamahu 0.90 2.31‡ 1.39 6.73‡ 2.30 9.03‡ 1.57 2.93‡
Okarito 1.61 4.03‡ 4.22 7.90‡ 5.83 11.92‡ 2.63 2.02
Egmont 0.94 3.79‡ 1.80 8.31‡ 2.74 12.10‡ 1.94 2.27
Pukaki 0.70 1.67‡ 0.90 5.95‡ 1.60 7.62‡ 1.33 3.58‡
Fork 1.06 2.31‡ 1.64 5.52‡ 2.71 7.83‡ 1.55 2.40‡
Richmond 0.74 2.36‡ 1.88 7.70‡ 2.62 10.06‡ 2.64 3.27
Himatangi 1.62 3.62‡ 3.55 11.89‡ 5.16 15.51‡ 2.21 3.31‡
Hurunui 2.06 2.57‡ 3.62 7.40‡ 5.69 9.97‡ 1.76 2.85‡
Te Kauwhata 2.38 3.28‡ 3.54 7.60‡ 5.92 10.88‡ 2.63 2.33
Temuka 1.14 2.10‡ 2.92 6.70‡ 4.05 8.80‡ 2.58 3.19
Oruanui 1.74 3.04‡ 2.91 7.84‡ 4.64 10.88‡ 1.67 2.57‡
Taupo 0.96 2.67‡ 2.78 7.41‡ 3.74 10.08‡ 3.03 2.78
Patoka 0.98 3.25‡ 1.73 8.31‡ 2.71 11.56‡ 1.79 2.62
Stratford 2.32 2.59 4.01 6.56‡ 6.33 9.16‡ 1.91 2.50

Mean 1.29 2.74 2.51 7.32 3.77 10.06 2.13 2.75

Standard error 0.16 0.19 0.29 0.44 0.44 0.60 0.14 0.11
LSD0.05 0.73 1.55 2.15 0.85
Two-way ANOVA (F ratios):
Plant species 233.7∗∗∗ 577.7∗∗∗ 514.3∗∗∗ 31.4∗∗∗
Soil type 10.32∗∗∗ 10.3∗∗∗ 10.8∗∗∗ 2.3∗
Plant × soil 3.79∗∗∗ 3.8∗∗∗ 3.59∗∗∗ 2.9∗∗
a Grass – ryegrass; pine – radiata pine. ‡ indicates values for ryegrass are significantly lower than those for
radiata pine in the same row (P < 0.05, separated by LSD0.05 ). ∗ , ∗∗ , ∗∗∗ indicate significant differences at
0.05, 0.01 and 0.001 level (P < 0.05, 0.01 and 0.001), respectively.

tions of TPi were significantly higher in only two soils
(Stratford and Temuka) under radiata pine compared
with ryegrass (Table 6). However, concentrations of
TPo were consistently lower in soils under radiata pine
than under ryegrass (Table 6).
Specific P mineralisation rate (SMR) represents
the amount of organic P mineralised as a percent of
the total organic P present in the original soils prior
to planting (Grierson et al., 1999). Results showed
that the SMR was significantly higher in all soils un-
der radiata pine (8.4–21.9%) compared with ryegrass
(0.5–10.8%) (Figure 1).
Isotopically exchangeable soil P
Isotopic exchange kinetic parameters and exchange-
able P pools in soils under ryegrass and radiata pine
are shown in Tables 7 and 8. There were no significant
plant species effects on the levels of water soluble P
(CP ) in most soils. Concentrations of CP were signific-
antly higher in the Stratford, Himatangi and Hurunui
soils, but significantly lower in the Temuka soil un-
der ryegrass compared with radiata pine (Table 7).
Compared with the original soils, levels of CP in
soils under ryegrass and radiata pine were generally
lower due to plant uptake (Table 7). Data for the ca-
pacity factors (R/r1 , n) showed that values of R/r1
were significantly higher under radiata pine in Mapua,
Mangamahu, Pukaki, Te Kauwhata, Patoka and Strat-
ford soils, while there were no significant effects of
plant species on the values of R/r1 found for the
remaining soils. The values of R/r1 for most soils
increased under radiata pine and ryegrass compared
with the original soils except for the Patoka soil un-
der ryegrass (Tables 7). Plant species also affected the
value of n, which was higher in Mangamahu, Okarito
and Pukaki soils and lower in Mapua, Fork, Himatangi
and Te Kauwhata soils under ryegrass compared with
radiata pine (Table 7). The n value tended to be higher
 121
Table 3. Phosphorus uptake by ryegrass and radiata pine determined after a 40-week period of growth a

Soil P concentration (mg kg−1 ) P uptake (mg pot−1 )

Root Shoot Root Shoot Total
Grass Pine Grass Pine Grass Pine Grass Pine Grass Pine

Mapua 745 934‡ 1274 800† 0.21 1.44‡ 0.91 3.09 1.12 4.52
Mangamahu 881 1151‡ 1350 997† 0.79 2.66‡ 1.88 6.74‡ 2.68 9.39‡
Okarito 644 1118‡ 2053 1286† 1.04 4.48‡ 8.60 10.21 9.64 14.69‡
Egmont 663 1096‡ 868 1031 0.62 4.08‡ 1.56 8.58‡ 2.18 12.66‡
Pukaki 846 1228‡ 1260 1330 0.58 2.05‡ 1.13 7.90‡ 1.72 9.96‡
Fork 1018 1500‡ 1785 1374† 1.08 3.49‡ 2.93 7.58‡ 4.01 11.07‡
Richmond 908 1789‡ 2017 1328† 0.67 4.22‡ 3.79 10.23‡ 4.46 14.45‡
Himatangi 2162 1965† 2895 2215† 3.52 7.14‡ 10.19 26.32‡ 13.70 33.46‡
Hurunui 1056 1510‡ 2397 1478† 2.18 3.89‡ 8.69 10.97‡ 10.87 14.86‡
Te Kauwhata 798 1464‡ 1868 1315† 1.89 4.83‡ 6.54 9.93‡ 8.43 14.77‡
Temuka 1185 1524‡ 4875 1400† 1.35 3.20‡ 14.23 9.39† 15.58 12.58
Oruanui 970 1269‡ 1942 1496† 1.68 3.86‡ 5.60 11.62‡ 7.29 15.48‡
Taupo 1062 1524‡ 2111 1818 1.02 4.09‡ 5.79 13.41‡ 6.81 17.49‡
Patoka 909 1201‡ 1798 1153† 0.90 3.87‡ 3.12 9.53‡ 4.02 13.41‡
Stratford 1247 1635‡ 2126 1530† 2.87 4.27‡ 8.54 10.08 11.40 14.36

Mean 1006 1394 2041 1370 1.36 3.84 5.57 10.37 6.93 14.21
Standard error 88 68 224 82 0.22 0.31 0.94 1.21 1.10 1.49
LSD0.05 175 333 1.11 2.85 3.65
Two-way ANOVA
(F ratios):
Plant species 292.9∗∗∗ 243.1∗∗∗ 296.9∗∗∗ 170.0∗∗∗ 238.8∗∗∗
Soil type 48.1∗∗∗ 45.9∗∗∗ 13.9∗∗∗ 29.6∗∗∗ 27.9∗∗∗
Plant × soil 7.09∗∗∗ 24.7∗∗∗ 2.4∗∗ 9.8∗∗∗ 7.4∗∗∗
a Grass – ryegrass, pine – radiata pine. The symbol †indicates values for ryegrass are significantly higher than those
for radiata pine at the same row while ‡ indicates values for ryegrass are significantly lower than those for radiata pine
(P < 0.05, separated by LSD0.05 ). ∗ , ∗∗ , ∗∗∗ indicate significant differences at 0.05, 0.01 and 0.001 levels (P < 0.05,
0.01 and 0.001), respectively. P uptake was calculated as P concentration multiplied by plant biomass.

for most soils after 40 weeks growth compared with
the original soils (Table 7).
There were no significant effects of plant species
on the concentrations of E1min Pi for most soils, al-
though concentrations of E1min Pi were significantly
greater in the Egmont and Stratford soils and lower in
the Temuka soil under ryegrass compared with radi-
ata pine (Table 8). Concentrations of E1min−1d Pi were
higher in most soils (except for Mapua, Mangamahu,
Okarito, Egmont and Oruanui) under radiata pine
compared with ryegrass (Table 8). The values of
E1min−1d Pi for most soils under radiata pine (except
for Mangamahu, Okarito, Egmont and Oruanui) and
ryegrass (except for Okarito, Egmont, Fork, Hurunui,
Temuka, and Oruanui) were higher than those in the
original soils (Table 8). Concentrations of E1d−10m Pi
were significantly lower in Fork, Himatangi and Strat-
ford soils under ryegrass compared with radiata pine,
while there were no significant effects of plant species
on E1d−10m Pi in the remaining soils (Table 8). Plant
species had no different effects on the concentrations
of E>10m Pi in soils, while the concentrations of E>10m
Pi for most soils under ryegrass and radiata pine were
generally lower than those in the original soils (Tables
8). Concentrations of E1min−10m Pi (the sum of E1min ,
E1m−1d and E1d−10m Pi ) were significantly lower in
Fork, Himatangi, Te Kauwhata, Patoka and Stratford
soils under ryegrass compared with radiata pine while
there were no significant differences in the remaining
soils (Table 8). The values E1min−10m Pi determined
for most soils under radiata pine were also greater than
those for the original soils (Table 8).
122
Table 4. Selected soil chemical properties determined after a 40-week period of
growth under ryegrass and radiata pine a

Soil pH TOC (g kg−1 ) Total N (g kg−1 )

Grass Pine Grass Pine Grass Pine

Mapua 5.29 4.98† 19.7 18.9 0.84 0.91

Mangamahu 5.36 5.00† 39.6 35.5 2.78 2.46†
Okarito 5.48 4.71† 140.8 137.4 7.16 6.72†
Egmont 5.86 5.36† 51.1 46.2 4.17 3.91†
Pukaki 5.66 5.26† 47.4 42.4 2.99 2.71
Fork 5.89 5.46† 24.5 18.9 1.75 1.37†
Richmond 5.65 5.39† 37.5 36.1 3.27 3.01†
Himatangi 6.42 6.04† 31.3 25.1 2.88 2.26†
Hurunui 5.73 5.09† 75.5 75.9 5.98 5.99
Te Kauwhata 5.85 5.26† 49.8 45.5 4.56 4.21†
Temuka 6.09 5.47† 38.7 36.7 3.63 3.32†
Oruanui 5.40 5.21† 86.7 80.9 5.46 5.08†
Taupo 5.66 5.17† 52.7 46.3 3.87 3.39†
Patoka 5.68 5.42† 94.3 89.0 8.31 7.83†
Stratford 5.49 5.09† 60.8 58.8 5.80 5.46†

Mean 5.70 5.26 56.7 52.9 4.23 3.91

Standard error 0.08 0.08 0.82 0.82 0.05 0.05
LSD0.05 0.11 N/A 0.26
Two-way ANOVA
(F ratios):
Plant species 935.9∗∗∗ 61.2∗∗∗ 91.8∗∗∗
Soil type 107.2∗∗∗ 1153.4∗∗∗ 951.2∗∗∗
Plant × soil 8.6∗∗∗ 1.3 1.9∗
a See Table 3 for explanation of symbols. TOC = total organic C. N/A = not
applicable (due to insignificance in plant species × soil type interactions).

Figure 1. Specific phosphorus mineralisation rate (SMR) determined in soil after a 40-week period of growth under ryegrass and radiata pine
(standard errors are shown by vertical bars).
 123
Table 5. Bicarbonate extractable P (BPi and BPo ) (mg kg−1 ) and resin extractable P (resin Pi , Po )
(mg kg−1 ) determined in soils after a 40-week period of growth under ryegrass and radiata pine a

Soil BPi BPo Resin Pi Resin Po

Grass Pine Grass Pine Grass Pine Grass Pine

Mapua 1.0 0.7 7.4 3.1 3.0 3.1 7.0 12.3‡

Mangamahu 1.0 4.4 11.7 21.7‡ 3.0 6.7 12.6 16.5‡
Okarito 1.9 6.3‡ 11.3 14.4 10.8 18.3‡ 12.0 14.0
Egmont 1.3 3.4 4.9 19.3‡ 5.1 7.2 12.8 16.0
Pukaki 1.4 5.6‡ 13.1 17.7 7.5 13.0‡ 12.7 18.0‡
Fork 6.1 9.1 8.4 15.3 19.5 19.0 12.8 18.1‡
Richmond 1.3 6.8‡ 8.9 12.8 6.7 15.6‡ 14.5 16.5
Himatangi 13.7 22.1‡ 3.8 9.7 32.6 37.2‡ 6.4 10.2‡
Hurunui 2.7 9.7‡ 12.5 20.1‡ 16.0 22.9‡ 20.2 22.6
Te Kauwhata 4.5 7.0 17.0 15.2 13.0 17.5 16.3 16.0
Temuka 8.4 21.4‡ 10.4 11.8 29.6 52.5‡ 17.6 18.1
Oruanui 21.1 27.0‡ 26.6 29.9 29.2 37.4‡ 25.8 22.6
Taupo 39.5 47.4‡ 19.9 20.1 63.8 64.1 22.3 24.4
Patoka 1.8 5.3 11.8 27.6‡ 11.8 11.5 19.3 21.6
Stratford 24.4 35.1‡ 26.9 39.2‡ 29.8 39.9‡ 23.6 26.1

Mean 8.7 14.1 13.0 18.5 18.8 24.4 15.7 18.2

Standard error 2.9 3.5 1.8 2.3 4.2 4.6 1.5 1.2
LSD0.05 4.1 7.2 4.8 3.8
Two-way ANOVA
(F ratios):
Plant species 75.9∗∗∗ 35.6∗∗∗ 159.7∗∗∗ 20.6∗∗∗
Soil type 144.9∗∗∗ 16.9∗∗∗ 378.6∗∗∗ 44.7∗∗∗
Plant × soil 4.9∗∗∗ 2.6∗∗∗ 11.7∗∗∗ 3.7∗∗∗

a See Table 3 for explanation of symbols.

Root phosphatase activity
Root surface phosphatase activities measured at the re-
spective optimum buffer pH were consistently higher
in radiata pine (average 1450 µg p-NP g−1 h−1 ) than
ryegrass (average 755 µg p-NP g−1 h−1 ) for all soils
(Figure 2).
Discussion
During this 40-week experiment, no apparent symp-
toms of nutrient deficiency were observed across the
15 soils although no nutrients were added over the
experimental period. Differences in growth (biomass)
between radiata pine and ryegrass under this exper-
imental condition reflected the differences in plant
genetics and also the plant species × soil environment
interactions. There were significant plant species ×
soil type interactions for most soil variables measured
in this study (Tables 4–8). These interactions presum-
ably arose from the variations in the properties of the
original soils (Table 1).
Plant P uptake in different soils
It is well known that plant genus, species or even
genotypes of the same species may vary in their abil-
ity to take up nutrients from soil (e.g., Gahoonia et
al., 1999). This has been related to a combination of
factors including root size and distribution (including
root hair length and root surface), root exudation (H+ ,
HCO3 − , reducing agents, chelates, organic anions,
enzymes), mycorrhizal infection, and transpiration
rate (e.g., Foehse et al., 1988). The overall interac-
tion of soil and plant properties (nutrient availability
and acquisition by plants) determines the rate and the
quantity of P transfer into plants (Jungk, 1996).
In this study, radiata pine and ryegrass showed
differences in plant growth and nutritional charac-
124

Table 6. Concentrations of soil inorganic, organic and total P (mg kg−1 ) de-
termined after a 40-week period of growth under ryegrass and radiata pine
a

Soil TPi TPo Total P

Grass Pine Grass Pine Grass Pine

Mapua 30 24 81 75 111 99
Mangamahu 76 105 301 252† 377 356
Okarito 64 80 472 423† 536 503†
Egmont 117 134 469 408† 586 542†
Pukaki 121 167 531 429† 652 596†
Fork 415 403 240 220 655 623†
Richmond 228 243 547 479† 775 722†
Himatangi 445 440 309 286 755 725†
Hurunui 197 214 617 581 815 795
Te Kauwhata 295 334 598 519† 894 853†
Temuka 465 540‡ 454 418 919 958‡
Oruanui 345 384 694 618† 1039 1002†
Taupo 712 668 461 422 1173 1089†
Patoka 451 495 1031 927† 1482 1422†
Stratford 1471 1705‡ 1136 911† 2607 2615

Mean 362 396 529 464 891 860

Standard error 93 105 71 60 148 149
LSD0.05 50 40 30
Two-way ANOVA
(F ratios):
Plant species 27.4∗∗∗ 153.4∗∗∗ 65.8∗∗∗
Soil type 953.7∗∗∗ 623.1∗∗∗ 5979.6∗∗∗
Plant × soil 6.3∗∗∗ 6.8∗∗∗ 3.8∗∗∗

a See Table 3 for explanation of symbols. TP – total P ; TP – total Po.

i i o

Figure 2. Root phosphatase activities of ryegrass and radiata pine determined after a 40-week period of growth (standard errors are shown by
vertical bars).
 125
Table 7. Isotopic exchange kinetics parameters (CP , R/r1 , n) determined in soils after a 40-week period of growth
under ryegrass and radiata pine a

Soil CP (mg L−1 ) R/r1 amc3n

Original Grass Pine Original Grass Pine Original Grass Pine

Mapua 0.10 0.05 0.06 2.9 4.1 5.7‡ 0.31 0.42 0.49‡
Mangamahu 0.02 0.03 0.03 7.4 9.0 12.1‡ 0.46 0.46 0.35†
Okarito 1.90 1.33 1.33 1.0 1.0 1.0 0.10 0.14 0.09†
Egmont 0.08 0.07 0.04 9.5 13.2 13.8 0.43 0.43 0.43
Pukaki 0.05 0.04 0.04 2.7 3.4 5.0‡ 0.50 0.50 0.45†
Fork 0.11 0.11 0.09 1.8 1.7 2.0 0.43 0.41 0.45‡
Richmond 0.11 0.11 0.12 3.0 4.3 4.5 0.42 0.45 0.43
Himatangi 0.97 1.12 0.88† 1.1 1.0 1.1 0.12 0.13 0.21‡
Hurunui 0.81 0.45 0.39† 1.4 1.6 1.7 0.25 0.31 0.34
Te Kauwhata 0.19 0.13 0.11 2.6 4.2 5.7‡ 0.40 0.45 0.48‡
Temuka 1.65 0.84 1.26‡ 1.3 1.6 1.4 0.16 0.21 0.22
Oruanui 0.10 0.09 0.08 3.1 5.2 5.2 0.50 0.49 0.50
Taupo 0.26 0.17 0.15 2.4 2.9 3.8 0.45 0.50 0.50
Patoka 0.04 0.04 0.04 17.7 13.0 19.7‡ 0.44 0.50 0.50
Stratford 0.16 0.20 0.14† 8.9 10.5 13.6‡ 0.46 0.46 0.49

Mean 0.44 0.317 0.32 4.5 5.1 6.4 0.37 0.39 0.40
Standard error 0.16 0.11 0.12 1.2 1.1 1.5 0.04 0.03 0.03
LSD0.05 0.05 1.2 0.04
Two-way ANOVA
(F ratios):
Plant species 0.34 69.6∗∗ 0.78
Soil type 1099.4∗∗∗ 272.5∗∗∗ 200.3
Plant × soil 32.1∗∗∗ 9.2∗∗∗ 6.8∗∗∗
a Data for original soils are not included in ANOVA, but as reference. See Table 3 for explanation of abbreviations
and symbols.

teristics. Radiata pine produced 1.4–5.5 times more
biomass than ryegrass, while the S:R ratios of ra-
diata pine were also generally higher than those of
ryegrass. Moreover, radiata pine generally took up
more P in most soils (except for the Temuka soil)
compared with ryegrass (Table 3). Levels of E1min Pi
in the original soils were negatively correlated with
the corresponding P uptake ratios by radiata pine to
ryegrass (Figure 3). This indicated that radiata pine
was able to take up more P than ryegrass when the
level of available P was low, which in turn showed
that radiata pine was better able to access sparingly
soluble sources of soil P. On the other hand, differ-
ences in P uptake between the two species declined
as soil available P (E1min Pi ) increased (especially for
E1min Pi > 15 mg kg−1 ), while ryegrass took up more
P than radiata pine when the levels of available P were
high (Figure 3). For example, the Temuka soil had a
high level of available P (E1min Pi 29 mg kg−1 before
planting), and ryegrass took up more P than radiata
pine (Tables 3 and 8), which may have resulted in
lower levels of available Pi and total P in the soil un-
der ryegrass (Tables 5 and 6). The obvious differences
in P uptake between radiata pine and ryegrass may
be attributed to the differences in root morphology
and associated chemical, biochemical and biological
properties.
Vesicular–arbuscular mycorrhizae (VAM) are as-
sociated with ryegrass (Powell, 1977), while ectomy-
corrhizae (ECM) are associated with roots of radiata
pine (Chu-Chou and Grace, 1990). Detailed examin-
ation of mycorrhizae associated with ryegrass and ra-
diata pine was not undertaken in this study. However,
ectomycorrhizal hyphae were clearly visible in soils
under radiata pine, and abundant mycorrhizae as char-
acterized by dense mycelial sheaths were observed on
roots of radiata pine at the end of experiment. There
is a lack of direct comparisons of root-VAM and root-
 126

Table 8. Isotopically exchangeable Pi pools (mg kg−1 ) determined in soils after a 40-week period of growth under ryegrass and radiata pine a

Soil E1min E1min−1d E1d−10m E>10m E1min−10m

Original Grass Pine Original Grass Pine Original Grass Pine Original Grass Pine Original Grass Pine

Mapua 3.3 2.4 3.4 12.0 15.6 16.3 12.9 10.2 4.0 5.9 1.8 0.3 28.2 28.2 23.7
Mangamahu 2.1 2.4 3.3 31.1 32.3 26.4 42.3 36.2 50.6 11.3 5.1 24.7 75.5 70.9 80.3
Okarito 21.4 14.0 14.2 10.4 9.6 6.0 12.0 12.3 7.7 74.3 28.1 52.1 43.8 35.9 27.9
Egmont 8.4 8.8 6.2† 73.1 64.2 62.2 52.6 38.7 54.4 9.6 5.3 11.2 134.1 111.7 122.8
Pukaki 1.7 1.5 2.4 35.4 36.7 38.4 75.7 68.4 93.5 16.4 14.4 32.7 112.8 106.6 134.3
Fork 2.9 2.8 2.5 37.8 31.9 39.3 201.1 164.3 202.2‡ 206.7 216.0 159.0 241.8 199.0 244.0‡
Richmond 4.3 5.3 6.4 53.5 71.8 77.2 127.8 122.8 124.1 59.1 28.1 35.3 185.6 199.9 207.7
Himatangi 13.2 14.9 13.3 10.0 14.0 26.1 23.0 24.4 71.2‡ 484.9 391.7 329.4 46.2 53.3 110.6‡
Hurunui 15.9 9.4 8.6 41.1 40.6 46.8 72.5 82.2 98.4 104.0 64.8 60.2 129.5 132.2 153.8
Te Kauwhata 6.9 6.3 7.0 71.2 86.5 120.5 163.6 162.1 174.7 90.5 40.1 31.8 241.7 254.9 302.2‡
Temuka 29.1 16.1 23.5‡ 35.8 31.2 53.9 65.4 95.0 118.1 417.1 322.7 344.5 130.3 142.3 195.5
Oruanui 4.3 5.2 4.7 142.3 20.6 119.3 205.7 180.1 212.7 73.7 39.1 47.3 352.3 305.9 336.7
Taupo 8.9 6.4 7.0 137.1 142.8 160.3 453.6 434.5 399.6 232.9 128.3 101.1 599.6 583.7 566.9
Patoka 7.6 5.7 7.2 130.3 156.9 189.9 289.6 235.8 254.7 107.0 52.6 43.2 427.5 398.4 451.8‡
Stratford 16.3 22.2 19.2† 321.0 391.7 451.3 893.5 833.7 1005.5‡ 349.1 223.4 229.0 1230.8 1247.6 1476.0‡

Mean 9.8 8.2 8.6 76.1 83.1 95.6 179.4 166.7 191.4 149.5 104.1 100.1 265.3 258.0 295.6
Standard error 2.1 1.5 1.6 21.0 25.1 29.0 56.0 55.5 64.0 37.5 32.4 29.2 75.2 80.7 93.0
LSD0.05 1.7 N/A 36.1 N/A 35.4
Two-way ANOVA
(F ratios):
Plant species 2.8 8.7∗∗ 26.3∗∗∗ 0.45 67.8∗∗∗
Soil type 199.5∗∗∗ 222.7∗∗∗ 577.4∗∗∗ 119.8∗∗∗ 1430.3∗∗∗
Plant × soil 6.9∗∗∗ 1.6 5.25∗∗∗ 1.3 10.4∗∗∗

a See Table 3 and 4 for explanation of symbols. Data for original soils are not included in ANOVA, but as reference. E
1min−10m = E1min + E1min−1d + E1d−10m .

Figure 3. Relationship between the levels of E1min Pi in the original
soils and ratios of P uptake by radiata pine to ryegrass determined
after a 40-week period of growth (the dash line indicates equality in
P uptake by ryegrass and radiata pine).
ECM associations in the literature. In general, ECM
hyphae are thought to be more efficient in P uptake
and transportation to the host plants than VAM hyphae
(Marschner and Dell, 1994). This has been ascribed
to a combination of factors including greater exten-
sion of ECM hyphae into soils compared with VAM
hyphae, low internal Pi concentration in ECM due to
formation of polyphosphates, and higher soil volume
explored by ECM due to smaller hyphae diameter and
the potential role of acid phosphatase and phytase en-
zymes produced by ECM hyphae (e.g., Jones et al.,
1998). Moreover, ECM hyphae have been found to
release significant quantities of low molecular weight
organic acids such as oxalic acid which may mobilize
P from sparingly soluble Ca-P, Fe-P or Al-P in soil
(Lapeyrie et al., 1991; Vogt et al., 1991), while lower
pH in soils under conifers such as radiata pine may
also enhance the solubility and consequent availabil-
ity of soil inorganic P and organic P (Gahoonia and
Nielsen, 1992).
In the present study, multiple regression analysis
showed that E1min Pi and R/r1 were the most import-
ant factors which influenced P uptake by ryegrass.
Accordingly, approximately 80% of the variation in
P uptake by ryegrass was explained by the following
equation:
P uptake (mg pot−1 ) = 3.674 + 0.631E1min Pi
(mg kg−1 ) − 0.437R/r1 (n = 15, P < 0.001).
However, the relationship between P uptake and soil
P for radiata pine could not be described by the same
multiple regression approach. The CP , E1min Pi and
127
E>10m Pi accounted for only 11.6–24.4% of the vari-
ation in P uptake by radiata pine, while other factors
(R/r1 , n, other exchangeable P pools and TPo ) were
not significantly correlated with P uptake by radiata
pine. These findings strongly indicate that radiata pine
was better able to utilize different forms of inorganic
and organic P in the soil, compared with ryegrass. In
addition, these results also imply that levels of labile
P did not appear to limit P uptake by radiata pine
possibly due to enhanced mineralisation of organic
P. Moreover, variation in P uptake by radiata pine
across soil types may also have been influenced by the
availability of other nutrients such as N.
In the present study, it was found that SMR varied
greatly with soil type and plant species, but the SMR in
soils under radiata pine were consistently higher than
in soils under ryegrass (Figure 1). This indicated that
radiata pine could better access forms of soil organic P
than ryegrass, and that the growth of radiata pine resul-
ted in increased mineralisation of organic P compared
with ryegrass.
Since P in soils is relatively immobile most spe-
cies take up their P from near the root surface, and
so root surface phosphatase activity may play a key
role in the P uptake by plants, especially when P
availability is low. The presence of ectomycorrhizae
increases root acid phosphatase activity (Pasqualini
et al., 1992), while Tarafdar and Claassen (1988)
reported that root-derived phosphatase activity was
significantly correlated with the amount of soluble or-
ganic P hydrolyzed. Antibus et al. (1992) also found
that increased 32 P uptake from inositol polyphosphate
was related to acid phosphatase activity of ectomycor-
rhizae. In the present study, root phosphatase activity
was 1.2–13.2 times higher for radiata pine compared
with ryegrass, which may contribute to the higher pro-
portion of P obtained from soil organic P sources by
radiata pine.
Effects of plant species on soil P forms and
availability
The effects of plant species on soil nutrient availability
have been studied in many ecosystems. For example,
Nielsen and Dalsgaard (1987) investigated the effect
of invasion of oak (Quercus ronur) on a Calluna heath
(Calluna vulgaris) in Denmark and found the rate of
organic matter decomposition increased under oak.
Magid (1993) found that under beech (Fagus sylvat-
ica) a large part of the extractable P was present in
labile inorganic fractions (resin Pi , bicarbonate Pi )
128
while under adjacent grass a large part of the extract-
able P was in labile organic P forms (bicarbonate and
fulvic acid Po ). This indicated that vegetation affected
the nature and distribution of soil P.
Despite the fact that P uptake was greater by ra-
diata pine compared with ryegrass in most soils, con-
centrations of labile P (BPi , BPo , resin Pi and resin Po ,
E1min−1d Pi , E1d−10m , Pi , E1min−10m Pi ) were gener-
ally higher in many soils under radiata pine compared
with ryegrass (Tables 5, 6 and 8). Furthermore, the
higher concentrations of labile P under radiata pine
were accompanied by enhanced mineralisation of or-
ganic P compared with ryegrass (Table 6; Figure 1).
These results are consistent with a similar study using
a restricted range of soils (Davis, 1995) and with nu-
merous ‘paired-site’ comparison studies carried out in
New Zealand (e.g., Chen et al., 2000; Condron et al.,
1996; Davis and Lang, 1991). Soil P transformations
are microbially mediated (Frossard et al., 2000). How-
ever, the extract mechanisms involved in enhanced
mineralization of soil organic P under radiata pine
compared with ryegrass is still unknown. Soil phos-
phatase activities are considered to play an important
role in the mineralization of organic P (Jungk, 1996),
but phosphatase activities were lower in most soils un-
der radiata pine compared with ryegrass in the present
study (data not shown). However, the soluble organic
C measured in this study was significantly higher in
all soils under radiata pine compared with ryegrass
(data not shown). It is postulated that increased ex-
udation of organic acids under radiata pine may have
increased solubility of organic P and then availabil-
ity of organic P for microbial attack. This may also
be partly responsible for enhanced mineralization of
organic P under radiata pine.
Both plant species were found to have lowered the
level of slowly exchangeable Pi (E>10m ) by approxim-
ately 31.8% compared with original levels (Table 8).
This is consistent with other studies that showed that
much of the P taken up by crop plants was derived
from the slowly exchangeable Pi pool when no P was
applied (Frossard and Sinaj, 1997). In addition, differ-
ences in the levels of E1min−10m between soils under
ryegrass and radiata pine were correlated with levels
of E>10m (r = 0.522∗, P < 0.05) and TPo (r =
0.661∗, P < 0.05) in original soils before planting.
Moreover, differences in Po depletion between soils
under ryegrass and radiata pine also correlated with Po
levels in original soil (Figure 4). These results indic-
ated that the extent of P redistribution also depended
on the original P levels.
Figure 4. Relationships determined between organic P in original
soils and differences in organic P depletion in soils under radiata
pine and ryegrass after a 40-week period of growth.
The above findings demonstrate that in many soils
the growth of ryegrass and radiata pine resulted in the
redistribution of P from the slowly exchangeable Pi
pool to rapidly exchangeable Pi pools. Radiata pine
caused a greater P redistribution from the organic P
pools to the inorganic P pools compared with ryegrass.
These plant species effects also varied from soil to soil.
Soil buffering capacity describes the ability of nu-
trients to transfer from the solid phase to the soil
solution. It has been shown that capacity factors (R/r1
and n) as measured by IEK can be affected by a com-
bination of soil properties and fertilization (Frossard
and Sinaj, 1997). In the present study, it was also
demonstrated that the values R/r1 and n were generally
higher in most soils after 40 weeks under ryegrass and
radiata pine compared with the original soils (Table 7)
and that the values of R/r1 in many soils under radiata
pine were higher compared with ryegrass (Table 7).
Plant species × soil interactions indicated that effects
of plant species on the values of n varied greatly with
soil type (Table 7). The higher values of R/r1 in soils
under radiata pine could be attributed to higher P up-
take by radiata pine. Depletion of P from the soil
solution is accompanied by an increase in R/r1 and
n values (Frossard and Sinaj, 1997). Morel and Hin-
singer (1999) also suggested that enhanced P uptake
in rhizosphere soil compared with bulk soil resulted
in higher R/r1 , while root activity can modify the
soil physiochemical properties and thereby affect P
exchangeability. Greater root exudation (H+ efflux,
organic acids) by radiata pine compared with ryegrass
(data not shown) may also enhance dissolution of
iron (Fe), aluminum (Al) and calcium (Ca) minerals

and consequently expose new P exchangeable sites
(Bar-Yosef, 1996).
Conclusions
Results from this glasshouse experiment clearly
showed that compared with ryegrass, radiata pine took
up more P from the soil particularly when the level
of available P (e.g., E1min Pi ) was low. The growth of
ryegrass and radiata pine resulted in the redistribution
of P from the slowly exchangeable Pi pool (E>10m Pi )
to the rapidly exchangeable Pi (E1min−1d Pi , E1d−10m
Pi ) pools in most soils, while radiata pine caused
greater P redistribution from organic P to inorganic P
compared with ryegrass. The impacts of plant species
on soil P redistribution vary greatly with soil type.
Acknowledgements
Funding for this study was provided by the New Zea-
land Forest Research Institute, Lincoln University,
the Center for Soil & Environmental Quality at Lin-
coln University, and the Swiss Federal Institute of
Technology Zürich (ETHZ). The authors would like
to thank Dr Colin Gray, Dr Ed Gregorich, Frank
O’Meara, Roger McLenaghen, Sjef Lamers, Graeme
Oliver and Wim Rijkse for their invaluable assistance
with collection of the various soil samples.
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