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The Norwegian Crop Research Institute, Apelsvoll Research Centre, N-2849 Kapp, Norway; 2The Norwegian
Crop Research Institute, Apelsvoll Research Centre, division Kise, N-2350 Nes pa Hedmark, Norway;
3
Department of plant and environmental sciences, University of Life Sciences P.O. Box 5003, N-1432 As,
Norway; *Author for correspondence: e-mail: annbjorg.overli@planteforsk.no; fax: +47-61-166901
Received 1 February 2005; accepted in revised form 14 June 2005
Key words: Hordeum vulgare L., Plant available phosphorus, Root growth, Root hair length, Starter
fertilizer
Abstract
Seed placed phosphorus (P) is hypothesized to increase P utilization in plants, particularly in cool climate.
A greenhouse experiment was conducted to evaluate the eects of seed placed P at two temperature regimes
(7 C night/9 C day or 13 C night/15 C day) within the rst weeks of the growing season on root hair
formation, shoot growth and P content of spring barley (Hordeum vulgare L.). Three soils (loam, clay loam
and sandy silt texture) were used, each with two levels of plant available P (low and high P-AL). In half of
the pots, all P (10 mg P kg1 soil) was applied 5 cm below the seed. In the other pots, half of the P was
applied in the same way, and half was given together with the seeds as starter fertilizer. When plants
reached growth stage (GS) 21, two replicates were harvested and temperature adjusted to 15 C night/17 C
day for the remaining three replicates, which were harvested at GS 49. Phosphorus fertilizer placement did
not interact with temperature, soil P level or soil type, and it had also only few and slight eects on shoot
dry matter (DM) and P content. The lower temperature regime delayed growth rate by nine days until GS
21, but shoot DM and P content was not signicantly aected. At GS 49, both shoot DM and P content
increased by 20% from low to high temperature. The soil P level inuenced shoot P content to a high
degree, with an increase by 190% at high P-AL compared to low P-AL at GS 21, and by 170% at GS 49.
The length of root hairs within treatments was very variable, and no signicant dierences were found
between the treatments. So the ability to adapt morphologically to suboptimal conditions was not great
enough to avoid reduced growth because of P deciency. Placing some P together with the seeds could
either inhibit limited P uptake at suboptimal growth conditions in this trial. Seed placement of P is
recommended in Norway, especially on silty soils. However, the results from this study support what are
observed in eld, that the positive eects of seed placed fertilizer are variable. The subject indeed needs
further investigation before the eect of seed placed P are fully understood.
Introduction
Cool weather in spring and early summer is commonly experienced in Norwegian agriculture. As
148
and uptake. Cool conditions aect uptake due to
reduced mineralization rates, less rapid diusion
towards roots (Barber 1995), lower uptake kinetics
(Bravo-F and Uribe 1981) and reduced root
growth. Cool conditions lead to slower root growth
than under more optimal conditions (Engels
and Marschner 1992; Engels 1993; Pettersson
1995).The uptake of nutrients with low mobility,
such as P, is aected largely by root density and by
the distribution of both roots and plant available P,
and less by P transport within the soil (Schenk and
Barber 1979). These factors together increase the
likelihood of P deciency occurring during early
growth, before the root system is fully developed.
In Norway, a large proportion of cereals are sown
in spring, as autumn sown cereals are limited by the
severe winters. Thus all nutrient uptake must take
place within one short season.
Optimum placement of fertilizer has been of
interest over many years. In Norway bandplacement of fertilizer is common, at about 5 cm
below the seed and spaced at 25 cm between
alternate seed rows (Lyngstad 1977; Ekeberg
1982). For the last 6 years, there has been growing
interest in placing some fertilizer even closer to
seeds, as so-called starter fertilizer. The fertilizer
is meant to ensure an optimal P availability at the
start of the growth period (Randall and Hoeft
1988). Some P in the immediate vicinity of the seed
may enhance early growth of spring barley and in
turn increase yield (Marschner 1995). Starter fertilizer is given as a part of the total nutrient supply
to the plants. Field trials by Lauzon and Miller
(1997) showed that the yield of maize increased
with starter fertilizer, regardless of soil available P
levels. The yield response of starter fertilizer was
attributed to increased shoot P concentration early
in the growth period. For barley, P banded with or
near the seed gave a higher yield increase than
when P was incorporated into the soil, but the soil
P level at which there was no yield response to
fertilizer P, varied form soil to soil and from year
to year (Malhi et al. 1993). Ketcheson (1968)
found the best eect of starter fertilizer in maize
production at high air and low soil temperatures,
while it had little eect at low air and high soil
temperatures. A factor to be kept in mind is that
excessive fertilizer applied in direct contact with
seeds, especially if other nutrients are given
together with P, may actually decrease yields by
delaying emergence (Miller et al. 1971).
149
Norway. Each soil was divided in two groups,
which were similar in most respects except for their
content of plant available P. The soils chosen
represented typical soil types used for cereal crop
production in Norway.
The loam and the clay loam were collected from
long-term fertilization eld trials, the rst from
Mystad in the county of Hedmark (6047 N,
1110 E) and the second from As in the county of
Akershus (5939 N, 1045 E). The sandy silt soil
was collected from a farm at Romerike, also in the
county of Akershus (6014 N, 1131 E).
For the loam, samples were collected from two
treatments, which had received either potassium
fertilizer only (P low) or a medium rate of farmyard manure (P high), both since 1922 (Ekeberg
and Riley 1995). For the clay loam, samples were
collected from plots that had received since 1966
either 0 (P low) or 48 kg P ha1 year1 (P high) as
mineral fertilizer (Uhlen and Rd 1983). For the
sandy silt, samples with two soil P levels (P low
and P high), due to dierences in previous
management, were collected.
The soils were passed through a 1-cm sieve and
stored moist and dark at 4 C until they were
potted. Sub-samples were air-dried at 35 C, passed through a 2-mm sieve and analysed in order to
make an initial characterization of the soils (Tables 1 and 2). For measurement of the content of
plant available P in soil, the P-AL-method was
used. The soils were extracted with a mixture of 0.1
M ammonium lactate and 0.4 M acetic acid, at pH
3.75 (Egner et al. 1960). This method is commonly
used in several Nordic countries, and is the recommended soil P-test in Norway for determining
fertilizer applications. P-AL values below
Pot experiment
The pot experiment was conducted in a phytotron
under two contrasting temperature regimes. The
experimental design was a factorial splitsplit plot
with ve replications and two harvest times. The
treatments were temperature regime, soil types,
soil P-AL levels, and various placements of P fer-
Loam, P low
Loam, P high
Clay loam, P low
Clay loam, P high
Sandy silt, P low
Sandy silt, P high
P-ALa
P-H2Ob
Olsen-Pc
org. Pd
Total P
25
133
23
139
27
116
4.8
3.0
2.0
2.2
0.4
2.9
8
42
18
78
16
24
454
529
404
532
246
669
970
1281
840
1447
498
1110
P-AL, P extracted with ammonium lactate and acetic acid as described by Egner et al. (1960).
P-H2O, P extracted with water by shaking 1.5 g soil and 30 ml distilled water for 22 h.
c
Olsen-P, P extracted with sodium bicarbonate as described by Olsen et al. (1954).
d
Organic P, the dierence between total P and inorganic P, both measured by the method of Mberg and Petersen (1982).
a
150
Table 2. Selected physical and chemical characteristics of the soils used in the pot experiment.
Ignition
Loam, P low
Loam, P high
Clay loam, P low
Clay loam, P high
Sandy silt, P low
Sandy silt, P high
Siltd
Clayd
Lossc
35
37
41
41
72
79
14
17
21
29
6
5
6.8
10.5
7.0
9.3
5.0
5.2
SOMa
pH
(H2O)
K-ALb
(mg kg1)
K-HNOc3
(mg kg1)
Mg-ALb
(mg kg1)
Ca-ALb
(mg kg1)
4.2
6.9
4.1
5.6
3.2
3.3
5.8
6.4
5.6
5.8
5.9
6.8
23
24
9
11
13
11
83
85
62
63
30
35
58
211
68
62
122
163
1920
2820
1390
1920
427
1440
April
May
June
July
August
Follo
(5940 N,
1051 E)
stre Toten
(6042 N,
1053 E)
Stjrdal
(6328 N,
1054 E)
4.1
10.3
14.8
16.1
14.9
2.3
9.0
13.7
14.8
13.5
5.1
9.5
12.5
13.9
14.1
151
at a resolution of 400 dpi (Epson Twain Pro 2.10)
in order to compute the length and mean diameter
of each measured root segment using the Win
RHIZO Pro software (Regent Instrumets Inc.).
The scanner images were analyzed using a lter
that excluded objects smaller than 0.02 mm2.
The following root diameter classes were separated: 00.5, 0.51, 11.5, 1.52 and 22.5 mm.
After scanning, root samples were collected on
lters and their dry weights were determined by
drying at 60 C. The roots from the remainder
three quarters were also washed out of the soil,
dried at 60 C and weighed. The dried roots were
ignited at 550 C for 4 h, and then reweighed. The
dierence between the root dry weight and the
weight of the ash residue is termed ash-free organic
root matter. Total root length per pot (RL, m) was
calculated using equation [1.1].
RL RL1=4 RW3=4 =RW1=4 RL1=4
1:1
Statistical analyses
Analyses of variance (General Linear Model) were
performed on shoot P content, shoot DM and root
hair length, using a splitsplitplot model with
temperature as main factor, soil type as the intermediate factor and soil P-AL level, and fertilizer P
placement as the second splitplot factor. For root
length and root mass there were no complete
replications and the assumptions of variance
analysis were thus not met. All analyses were
performed with Minitab. Results with P 0.05
were considered signicant. Dierences between
treatments were determined using least signicant
dierence (LSD) values.
Table 4. Probability levels of F-tests of the main eects and interactions of starter fertilizer, temperature regimes, P-AL levels and soil
types on shoot dry matter and shoot P content at rst and second harvests.
Treatment
First harvest, GS 21
Second harvest, GS 49
Shoot dry matter g pot1 Shoot P content mg pot1 Shoot dry matter g pot1 Shoot P content mg pot1
Starter fertilizer (SF)
Temperature (T)
P-AL (P)
Soil (S)
SFT
SF P
SF S
TP
TS
PS
SF T P
SF T S
SF P S
TPS
n.s.
n.s.
<0.001
0.004
n.s.
n.s.
n.s.
n.s.
n.s.
0.02
n.s.
n.s.
n.s.
n.s.
0.05
0.1
<0.001
0.001
n.s.
n.s.
n.s.
n.s.
0.08
<0.001
n.s.
n.s.
n.s.
n.s.
0.04
0.05
<0.001
<0.001
n.s.
0.07
n.s.
n.s.
0.01
<0.001
n.s.
n.s.
n.s.
n.s.
Signicance was assumed at P 0.05 and values approaching P 0.05 are also indicate.
n.s.
0.07
<0.001
<0.001
n.s.
n.s.
n.s.
0.02
0.07
<0.001
n.s.
n.s.
n.s.
n.s.
152
tent and shoot DM. Further, P fertilizer placement
did not interact with temperature, soil type or PAL levels. However, the main eect of P placement
was signicant (P = 0.05) for shoot P content at
rst harvest and for shoot DM at second harvest
(P = 0.04).
There were signicant interactions between soils
and soil P-AL level (P < 0.001) at both harvests.
The loam soil with highest P-AL level gave the
highest levels of shoot P content and shoot DM,
while the sandy silt soil with lowest P-AL level
gave the lowest. No other two-factor interactions
were signicant at rst harvest. At second harvest,
the interaction between soil type and temperature
was signicant (P = 0.01) for shoot DM. Plants
responded signicantly to temperature increase,
with the loam soil having the highest DM production. The interaction between temperature and
soil P-AL levels for shoot P content was also signicant (P = 0.02) at the second harvest. Higher
temperature thus seemed to improve plant use of
the soil P-AL.
153
The temperature eects on root length, root
weight and root to shoot ratio at GS 21 are given
in Table 6. The results did not show any great
dierences in root length and dry weight at different temperatures. This means that even though
the root system was almost similar at both temperatures, the roots of plants grown at low temperature seemed unable to take up and translocate
the same amount of P as plants grown at high
temperature. There may be several reasons for
this. Moorby and Nye (1984) found that inow of
phosphate in rape was independent of temperature
within the range 1023 C, but was halved at 5 C.
The lowest temperature regime in our experiment
was 7/9 C, so the decreased shoot P content was
probably a consequence of reduced inow of P or
a decrease in the mobility of membrane phospholipids (Bravo-F and Uribe 1981). It may also have
been due to a decrease in the rate of mineralization
of organic P with decreased soil temperature.
In our experiment, root to shoot ratio increased
at low temperature (Table 6). This is a well-known
response to low temperature in many species
(Anderson and Kemper 1964; Equiza et al. 2001),
and has been considered as a response to overcome
water limitation. This is because low soil temperatures reduce root hydraulic conductivity, which
can induce water decit in shoots (Equiza et al.
2001). A high root to shoot ratio is a result of
higher amounts of carbohydrates and other compounds being transported to the roots, which
reduces shoot growth.
Table 5. Shoot dry matter (g DM pot1) and shoot P content (mg P pot1) of barley grown at two temperature regimes and three soil
types.
First harvest, GS 21
Second harvest, GS 49
Temperature
Temperature
7/9 C
13/15 C
7/9 C
13/15 C
1.39
0.84
0.66
15.8
1.00
0.73
19.1
12.8
10.6
14.6
12.8
n.s
2.6
1
3.87
3.15
1.54
5.57
3.80
1.86
n.s.
28.5
23.5
13.8
35.1
26.1
16.3
n.s.
154
Table 6. Mean root dry matter (g DM pot1), root lengths
(m pot1) and root to shoot ratios of barley with and without
starter fertilizer (SF), two temperature regimes, two P-AL levels
and three soil types.
Without SF
With SF
7/9 C
13/15 C
P low
P high
Loam
Clay loam
Sandy silt
Root length
(m pot1)
Root to
shoot ratio
0.31
0.35
0.38
0.26
0.36
0.29
0.36
0.34
0.27
57
60
66
52
67
50
70
62
43
0.38
0.39
0.48
0.30
0.51
0.26
0.32
0.40
0.44
Table 7. Shoot dry matter (g DM pot1) and shoot P content (mg P pot1) of barley grown at two P-AL levels and three soil types.
First harvest, GS 21
Second harvest, GS 49
P-AL
P-AL
Low
High
Low
High
0.97
0.70
0.53
1.54
1.15
0.85
15.4
10.6
7.4
19.5
16.7
15.9
2.51
1.65
0.98
0.11
0.6
6.93
5.30
2.42
0.47
17.8
11.5
9.1
45.8
38.1
21.1
1.8
155
Figure 1. Mean root hair length (mm) of barley as aected by P-AL level, two dierent P placements, temperature and soil types, at
growth stage 21 (open bars) and at growth stage 49 (lled bars). Vertical bars represent standard errors of the means.
156
Figure 2. Relationships between root hair length (mm) and P concentration (g P 100 g DM1) of plants grown on three soils, at two
P-AL levels, two temperatures and with two dierent P placements.
157
also favourable for aggregate stability (Riley
1983). Such factors may in part explain the lower
shoot DM that were obtained on the sandy silt
soil, but could not explain the dierences in yield
between the loam soil and the clay loam soil.
Conclusion
P fertilizer placement gave only few and small effects on shoot DM and P content. No interactions
between P placement and temperature, soil P-AL
level and soil types were found. Low temperature
within the rst weeks of the growing season decreased P uptake, both when P fertilizer was seedplaced and when it was applied 5 cm below the
seed. The plants grown at the lowest initial temperature never caught up with the plants grown at
higher initial temperature. At GS 49, both shoot
DM and P content increased by 20% from low to
high temperature, on average for all soils. The soil
P level inuenced the shoot P content to a high
degree. The shoot P content at GS 21 increased by
190% on average for all soils at high P-AL compared to low P-AL, and by 170% at GS 49. The
higher content of P at higher soil P-AL levels was a
result of both higher P concentration in the plants
and higher shoot DM yield. There was high variation in root hair length within all treatments, and
no signicant dierences were found between the
treatments. So the ability to adapt morphologically to suboptimal conditions was not great enough to avoid reduced growth because of P
deciency. Placing some P together with the seeds
could either inhibit limited P uptake at suboptimal
growth conditions in this trial. Seed placement of
P is recommended in Norway, especially on silty
soils. However, the results from this study support
what are observed in eld, that the positive eects
of seed placed fertilizer are variable. The subject
indeed needs further investigation before the eect
of seed placed P are fully understood.
Acknowledgements
This work was funded by the Norwegian Research
Council. Thanks to Prof Tore Krogstad, UMB
and Dr Anne Falk gaard, UMB for advice
during the experimental work.
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