Springer 2005

Nutrient Cycling in Agroecosystems (2005) 73:147159

DOI 10.1007/s10705-005-0076-2

Eects of P fertilizer placement and temperature on root hair formation, shoot

growth and P content of barley grown on soils with varying P status
Annbjrg verli Kristoersen1,*, Hugh Riley2 and Trine A. Sogn3
1

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 kg
1 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

most nutrient uptake by plants takes place within

the rst 710 weeks of the growing season
(Gahoonia et al. 1999), the temperature during this
period is very important for nutrient availability

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).

Plants have various strategies for increasing P

uptake when this nutrient is scarce. They may
increase root length by producing longer and
thinner roots with less branching, they may increase
root hair density, develop mycorrhizal associations
or modify the rhizosphere by excreting chelating
compounds, organic acids or phosphatase enzymes
(Anuradha and Narayanan 1991; Smith et al. 1994;
Gahoonia et al. 1997; Hinsinger 1998).
Root hairs increase the uptake radius of plant
roots, thus reducing the distance that P must diffuse (Baon et al. 1994a). There is high variability,
both between and within plant species, in their
distribution, density and length (Hofer 1996;
Krasilniko et al. 2003). The importance of root
hair length for P uptake has been studied by several workers Fohse and Jungk 1983; Itoh and
Barber 1983; Gahoonia et al. 1999; Bates and
Lynch 2001; Lcs 2003). It has been suggested that
root hair length increases at low P concentrations
in the growing medium (Bates and Lynch 1996).
On the other hand, Fohse and Jungk (1983)
maintained that root hair length is governed by
plant P concentration, and only indirectly by P
concentrations in the soil. In fact, both factors
probably play a role in the regulation of root hair
growth (Forde and Lorenzo 2001). However, there
is still considerable uncertainty about how soil
factors such as soil temperature, soil density and
nutrient availability aect root hair development.
The aim of this work was to study (1) whether
starter fertilizer has a greater eect at low than at
higher temperatures, and (2) whether it performs
better on silty soils, which are known to be relatively cold in spring, than on other common soils,
at varying levels of P availability in the soil. Furthermore, we wished to see (3) whether possible
eects of low early season temperatures remain of
importance later in the growing season. Finally we
wished to see (4) whether dierent soil types with
varying soil P availability resulted in dierences in
root hair length.

Material and methods

Soil characteristics
Bulk soil samples from a loam, a clay loam and a
sandy silt were collected from the upper layer
(020 cm) of three cultivated soils in southeastern

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 (60
47 N,
11
10 E) and the second from As in the county of
Akershus (59
39 N, 10
45 E). The sandy silt soil
was collected from a farm at Romerike, also in the
county of Akershus (60
14 N, 11
31 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 ha
1 year
1 (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

25 mg kg
1 are considered low, whilst those above

70 mg kg
1 are considered high, according to the
guidelines used in Norway (Krogstad 1992). To
give a broader picture of the soils P content,
various other P extraction methods were used
(Table 1). Soil was extracted with 0.5 M NaHCO3
buered at pH 8.5 to measure Olsen-P (Olsen et al.
1954), another common index of plant available P.
Water extractable P was measured by shaking
1.5 g soil and 30 ml distilled water for 22 h. Total
and inorganic P were analysed by the method of
Mberg and Petersen (1982), and the dierence
between these two was assumed to be organic P.
The P concentrations in the various extracts were
measured using the molybdenum-blue method
(Murphy and Riley 1962).
Particle-size distribution was determined by the
pipette method (Elonen 1971). Soil pH was measured in a soilwater suspension (1:2.5 v/v). To
evaluate the general nutrient status of the soils,
plant available K (K-AL), Mg (Mg-AL) and Ca
(Ca-AL) was measured as well. Acid soluble K
(K-HNO3) was extracted with 1 M nitric acid as
described by Pratt (1965). The K-HNO3 minus the
K-AL value is used to estimate the K reserves that
may become available to plants over time.

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-

Table 1. Phosphorus status of the soils used in the pot experiment.

Soil P status (mg P kg
1)

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 kg
1)

K-HNOc3
(mg kg
1)

Mg-ALb
(mg kg
1)

Ca-ALb
(mg kg
1)

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

Soil organic matter, corrected for clay content (Riley 1996).

K-AL, Mg-Al and Ca-AL: K, Mg and Ca extracted with ammonium lactate and acetic acid as described by Egner et al. (1960).
c
K-HNO3, acid soluble K extracted with nitric acid as described by Pratt (1965).
d
Weight% <2 mm, after USDA Soil Survey classication.
e
Weight% < 2 mm, ignition loss at 550
C.
b

tilizer. Three kg of air-dried soil were placed in each

plastic pot, and each pot received 10 mg P kg
1
soil (20 kg ha
1). In half of the pots, all P was
given as triple superphosphate, applied 5 cm below
the seeds. In the other pots, 5 mg P kg
1 soil was
applied in the same way, and the remaining P was
given as starter fertilizer. This meant that each seed
had a fertilizer grain in its immediate vicinity at
sowing time. As starter fertilizer, Opti StartTM NP
12-23 was used. Basal doses (120 mg kg
1 soil) of
N (ammonium nitrate) and K (potassium sulphate)
were applied in granular form 5 cm below the
seeds. In the pots with starter fertilizer, basal doses
were reduced corresponding to the N content of
starter fertilizer. Twelve seeds of barley (Hordeum
vulgare L. cv. Ven) were sown per pot and thinned
to nine once germinated. Soils were compacted by
hand to a relative degree of compactness of 75%
(Kristoersen and Riley 2005).
At the start of the trial, the mass wetness (w) in
the pots was adjusted to 30% and maintained at
that level by regular watering with deionized water. The loss of water was determined by weighing
pots before each watering. As the soils were of
varying texture and pore size distributions, soil
moisture tensions varied among soils, but the total
amounts of water in each pot were the same.
The experiment was carried out under two different temperature regimes, either 9
C by day/7
C
by night or 15
C by day/13
C by night until
plants reached growth stage (GS) 21, one tiller
formed (Zadoks et al. 1974). Then, two replicates
were harvested. After harvesting, the temperature
was adjusted to 17
C by day/15
C by night for the
three remaining replicates. When plants reached

GS 49, start of heading, these three replicates were

harvested. The temperature regimes were chosen
to represent the mean temperature of some representative areas for crop production in Norway
(Table 3). The light period was 14 h during the
whole experimental period. At harvest, the plants
were cut 1 cm above ground and their biomass was
recorded. The P concentration was determined
using inductively coupled plasma spectrometry
(ICP) after ignition and subsequent solubilization
of ashes in aqua regia. The P content in shoot DM
was calculated as dried shoot DM (g DM pot
1)
multiplied by dried shoot P concentration
(mg g
1 DM).

Sampling and length measurement of roots

At the rst harvest, soil from each pot of one
replicate was cut into four equal slices, and roots
from one quarter were gently washed out of the
soil. Roots were scanned using an optical scanner

Table 3. Mean temperature (
C) for the months AprilAugust

in three representative municipalities for grain production in
Norway.

April
May
June
July
August

Follo
(59
40 N,
10
51 E)

stre Toten
(60
42 N,
10
53 E)

Stjrdal
(63
28 N,
10
54 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

where RW3/4 is ash-free root dry weight of 3/4 of

the whole pot, and RW1/4 is ash-free root dry
weight of the rest of the roots. RL1/4 is total root
length of the 1/4 sample.

measured on each image, giving a total of 120 root

hairs per treatment. The analysis of root hair
lengths was performed using a digital camera
DC100 (Leica) tted to a light microscope (Leica
MZ6) at 10 magnication, and image processing
software (UTHSCSA ImageTool programme).

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.

Results and discussion

Root hair analysis
From one replicate at each harvest time, six representative images were taken of the roots from
each treatment. Twenty random root hairs were

The splitsplitplot analyses of variance (Table 4)

showed that no three-factor interactions between
fertilizer P placement, temperature, soil types and
soil P-AL levels were signicant for shoot P con-

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 pot
1 Shoot P content mg pot
1 Shoot dry matter g pot
1 Shoot P content mg pot
1
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.

Eects of dierent placement of P fertilizer

The eect of placing some P with the seeds compared with giving all the P fertilizer in a layer 5 cm
below the seeds was very small. There were only
two signicant eects of starter fertilization,
namely a slight increase in shoot P content at rst
harvest, from 3.16 to 3.44 mg P pot
1, and an
increase in shoot DM from 14.06 to 14.47 g DM
pot
1 at second harvest. There were no interactions between the level of plant available P in soils
and the seed-placement, either at GS 21 or GS 49.
And no interaction was observed between temperature and the placement of P on either shoot
DM or shoot P concentration.
Many eld trials have shown positive eects on
yield and nutrient uptake when some nutrients are
placed in the seed row (Bates 1971; Lauzon and
Miller 1997; Goos and Johnson 2001). In Finland,
Kleemola et al. (1998) found that starter application increased cereal crop yields, and that the response was higher for barley than for oats. The
best eect of starter fertilizer was obtained when
the soil P level was low. Alessi and Power (1980)
found that banding P with spring wheat seed was
benecial, even when soil P levels were high. Miller
et al. (1971) observed in a greenhouse study, that

maize plants receiving 6.5 kg P ha
1 with the seed,

grew and absorbed P more rapidly during the rst
four weeks compared to plants receiving
24.6 kg P ha
1 in a band to the side and below the
seed. After 7 weeks, however, they no longer
found any dierence in growth due to fertilizer
placement.
Several workers have suggested that particularly
when soil temperatures are low at sowing, it may
be an advantage to place some P close to the seed
(Kleemola et al. 1998; Chassot et al. 2001; Goos
and Johnson 2001). In a greenhouse study by
Costigan (1986), the presence of starter fertilizer
increased the nutrient concentration in lettuce
plants at both 10 and 20
C, but the growth was
only aected of the starter fertilizer at 10
C. At
the high temperature, there was no growth
response of starter fertilizer.
Field trials in Norway have shown that barley
plants grown on silty soils respond more to starter
fertilizer than those grown on loamy soils
(Kristoersen et al. 2004). However, no dierences in the response to starter fertilizer on different soils were found in our pot experiment.
It is dicult to fully explain why there were
such small dierences in shoot DM and P content
between the treatments with and without starter
fertilizer P in our experiment. One reason may be
the experimental set up. The soil volume that
roots could occupy was much smaller in the pots
than under eld conditions. This fact may inuence smaller dierence between the band placed P
and P given as starter fertilizer in the pots,
compared to eld conditions. One other explanation could be that the variety of barley used in
our experiment, Ven, has a lower response to
starter fertilizer than other barley varieties.
Rhoads and Wright (1998) showed that there
were large variations between maize hybrids, with
some responding well to starter P fertilizer, while
others not responding at all. In their experiment,
the starter fertilizer was mixed with the upper
25% of the soil volume in the pot, and compared
with mixing the P fertilizer within the total soil
volume. In Norway, greater response to starter
fertilizer have been found in barley than in oat
(Kristoersen et al. 2004) as was also found in
Finland (Kleemola et al. 1998). However, no
specic comparison of the responses of Norwegian barley varieties to starter fertilizer has been
made.

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.

Eects of dierent temperature

The plants took 9 days longer to reach GS 21 at low
than higher temperatures. The high temperature
gave the highest shoot DM on all three soils, even
though this was not statistically signicant
(Table 5). There was also a tendency for interaction
between soil type and temperature (P = 0.08) on P
content (mg P pot
1) at rst harvest. Higher
temperature increased the P content on all soils, but
the eect was greatest on the loam soil, intermediate on the clay and least on the sandy silt (Table 5).
The eect of temperature regime in the early
growth period was detectable on shoot DM at the
second harvest, with plants grown at high temperature having the highest shoot DM yield. These
plants reached GS 49 7 days before those grown at
low temperature in the earlier growth period.
Shoot DM increase from low to high temperature
was 21% on the loam and the sandy silt soil, and
14% on the clay loam soil (Table 5).
There was also a tendency to greater P content
at GS 49 for plants grown at high initial temperature compared to plants grown at the lower
temperature (P = 0.07) (Table 5). Singh and
Subramanian (1997) reported that the P content in
oats was 2.5 times higher at 18
C than at 9
C, and
Mackay and Barber (1984) found lower P content
in maize (Zea mays L.) at 18
C than at 25
C.
Thus, positive eects of increased temperature on
P uptake seem to occur over a relatively wide
range of temperatures.

Table 5. Shoot dry matter (g DM pot
1) and shoot P content (mg P pot
1) of barley grown at two temperature regimes and three soil
types.

Shoot dry matter (g pot
1)

1.12
Clay loam
Sandy silt
LSD(P = 0.05)

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

Shoot P content (mg pot )

Loam
Clay loam
Sandy silt
LSD(P = 0.05)
n.s. = not signicant.

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 pot
1), root lengths
(m pot
1) 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 dry matter

(g pot
1)

Root length
(m pot
1)

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

Eects of dierent soil P-AL levels

All pots were given 30 mg P as triple superphosphate. This was obviously not enough to give
optimal P availability for the plants grown at low
P-AL levels, since the shoot DM increased from
low to high P-AL level on all soils. As Table 7
shows, the highest shoot DM was at both harvests
measured on the loam soil at high P-AL, whilst the
lowest shoot DM was measured on the sandy silt
soil at low P-AL, both with and without starter
fertilizer.
The soil P-AL level also inuenced shoot P
content to a high degree (Table 7). On all soils,
both at GS 21 and GS 49, shoot P content was
about three times as high at high P-AL as that at
low P-AL. This was a result of both higher plant
P concentration and higher shoot DM yield.

Phosphorus uptake from soil depends on the

ability of the soil to maintain sucient P supply
to the roots. This property depends on the level of
P in soil solution and the ability of the soil to
supply P to the solution (Schenk and Barber
1979). A soil with a high level of AL-extractable P
is able to supply plant roots with more P compared to soils with low P-AL levels.
There was a signicant interaction between
temperature and P-AL level at the second harvest
for shoot P content. The shoot P content
increased from low to high P-AL level in the soil,
and the increase was somewhat higher for plants
grown at high temperature early in the growth
period compared to those grown at lower temperature. More P was probably released through
mineralization of organically bound P at the
higher temperature. Nielsen et al. (1960) also observed increased yield and P uptake in oats when
soil temperature was increased from 5 to 20
C.
They found that the eect of soil temperature on
uptake was more consistent of P than for other
elements.

Eects of soil P-AL level and temperature on root

hair growth
Root hairs extend the P depletion zone from the
root epidermis, thereby increasing the rate of P
uptake and the amount of P accessible to the roots
(Jungk 2001). In our experiment, there was high
variation in root hair length within all treatments
at both harvests, and no signicant dierences

Table 7. Shoot dry matter (g DM pot
1) and shoot P content (mg P pot
1) 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

Shoot dry matter (g pot
1)

Loam
Clay loam
Sandy silt
LSD(P = 0.05)

0.97
0.70
0.53

1.54
1.15
0.85

15.4
10.6
7.4

19.5
16.7
15.9

Shoot P content (mg pot
1)

Loam
Clay loam
Sandy silt
LSD(P = 0.05)

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.

were found between the treatments, or their

interactions. Therefore, only the main eects of PAL levels, P placement, temperature and soil types
are given in Figure 1. The gure shows that the
root hairs were equally well-developed in the soil
at both high and low P-AL levels, and have
probably been eective in the P uptake at both
levels. In other studies, a clear connection between
P concentration in the growing media and root
hair length has been found (Fohse and Jungk
1983; Bates and Lynch 1996; Gahoonia et al.
1999). Bates and Lynch (1996), when growing
Arabidopsis thaliana in a nutrient salt plant culture
media, found that local changes in P availability
inuenced root hair growth, regardless of the P
status of the plants. Fohse and Jungk (1983) found
the opposite result, and have concluded that the
formation of root hairs does not depend directly
on the P concentration at the root surface, but
rather on the P content of the plants. Forde and
Lorenzo (2001) have concluded that both the
internal P status of the plants and the P concentration outside the root hairs most probably contribute to the regulation of root hair growth.
However, we did not found any correlation
between the P content of the plants and root hair
length (Figure 2). The variation in root hair length
for plants grown under similar conditions was
greater than the variation in root hair length

between plants grown in soil with dierent soil PAL levels.

In our trial, the soil P-AL levels were about 25
and 130 mg P kg
1 soil. However, each pot received 30 mg P fertilizer in addition. This may
have been enough P to suppress possible dierences in root hair length that the original soil P
levels might have caused. Gahoonia et al. (1999)
studied variations in root hair growth between
cereal cultivars in soil, and found the largest
variations between cultivars grown in soil not
supplied with P fertilizers for about 30 years.
When 10 kg P ha
1 was applied, the root hair
length did not dier between the cultivars.
There have been relatively few studies of root
hair development in plants growing in soils. Most
studies have been performed in nutrient solution
(Fohse and Jungk 1983; Bates and Lynch 1996) or
in various modied growth media (Bates and
Lynch 2001). It is easiest to study root hairs in
nutrient solution. In soils, there may be a problem
that root hairs become damaged when the roots
are removed form the soil for observation, even
when one is very careful. When using natural soil,
there may also be confounding eects of phosphatases, protons, organic acids and other rhizosphere modifying exudates on P availability from
diverse soil pools, which lead to variations in the P
concentration along the roots. However, results

156

Figure 2. Relationships between root hair length (mm) and P concentration (g P 100 g DM
1) of plants grown on three soils, at two
P-AL levels, two temperatures and with two dierent P placements.

from a pot experiment with plants growing in soil,

will be more relevant to what may be expected in
the eld. Our results show that root hair length
seems to play a minor role in causing the variations in shoot P content of Ven barley grown in
soils with dierent P-AL levels.
No measurements of mycorrhiza were made in
the present study. As the growth period of spring
cereals is short, it has been suggested that mycorrhiza probably play only a minor role in their P
supply (Johnson et al. 1997). Mycorrhiza development takes place after 36 weeks (Baon et al.
1994b), whereas the plants P uptake start some
days after germination and continue in 710 weeks
after germination. The cost of such a symbiosis in
terms of carbohydrate loss early in the growing
season, may thus be greater than any benet to the
plant of such a symbiosis (Stribley et al. 1980;
Marschner and Dell 1994).

Temperature regime and soil types

It is worth mentioning that, in the present study,
the shoot DM production on loam at low temperature was actually higher than the shoot DM
production on both clay loam and sandy silt at
high temperature. This suggests that soil type is at
least as important for nal yield as the temperature

at which plants grew during the early growth

phase. Silty soils are renowned for often having
poor soil structure, with limited air capacity and
low permeability (Riley 1983). Therefore all pots
were compacted to the same level, 75% of the
standard degree of compactness. At this level, the
amount of large pores, suitable for unimpeded
root growth and gaseous exchange was high in all
soils (Kristoersen and Riley 2004), which means
that soil structure was probably not a limiting
factor here. The nutrient content in silty soils are
also often more limited compared to more loamy
soils, but it is dicult to see any explanations in
terms of the K, Ca and Mg availability in the soils
(Table 2). Although, there were dierences between the soils in K availability, with the highest
value in the loam soil, the K level was high enough
in all soils to rule out the likelihood of deciency,
and in any case all pots received a large dose of K
fertilizer.
There were dierences in the organic matter
contents between the soils. The sandy silt soil had
the lowest content, with approximately 3%, while
the loam soil and the clay loam soil had 47%
(Table 2). Since the ability of organic matter to
adsorb P is relative low, the P availability is often
higher in soils with high organic matter content,
compared to soils with low organic matter content
(Barber 1995). A high content of organic matter is

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.

References
Alessi J. and Power J.F. 1980. Eects of banded and residual
fertilizer phosphorus on dryland spring wheat yield in the
Northern Plains. Soil Sci. Soc. Am. J. 44: 792796.
Anderson W.B. and Kemper W.D. 1964. Corn growth as affected by aggregate stability, soil temperature, and soil
moisture. Agron. J. 56(5): 453456.
Anuradha M. and Narayanan A. 1991. Promotion of root
elongation by phosphorus deciency. Plant Soil 136: 273
275.
Baon J.B., Smith S.E. and Alston A.M. 1994a. Growth response and phopshorus uptake of rye with long and short
root hairs: interactions with mycorrhiza infections. Plant Soil
167: 247254.
Baon J.B., Smith S.E. and Alston A.M. 1994b. Phosphorus
uptake and growth of barley as aected by soil temperature
and mycorrhizal infection. J. Plant Nutr. 17(23): 479492.
Barber S.A. 1995. Soil Nutrient Bioavailability: A Mechanistic
Approach. 2nd edn. John Wiley & Sons, New York, p. 414.
Bates T.E. 1971. Response of corn to small amounts of fertilizer
placed with the seed: II. Summary of 22 eld trials. Agron. J.
63: 369371.
Bates T.R. and Lynch J.P. 1996. Stimulation of root hair
elongation in Arabidopsis thaliana by low phosphorus availability. Plant cell Environ. 19: 529538.
Bates T.R. and Lynch J.P. 2001. Root hairs confer a competitive advantage under low phosphorus availability. Plant Soil
236: 243250.
Bravo F-P. and Uribe E.G. 1981. Temperature dependence of
the concentration kinetics of absorption of phosphate and
potassium in corn roots. Plant Physiol. 67: 815819.
Chassot A., Stamp P. and Richner W. 2001. Root distribution
and morphology of maize seedlings as aected by tillage and
fertilizer placement. Plant Soil 231: 123135.
Costigan P.A. 1986. The eects of soil temperature on the response of lettuce seedlings to starter fertilizer. Plant Soil 93:
183193.
Egner H., Riehm H. and Domingo W.R. 1960. Untersuchung
uber die chemische Boden-Analyse als Grundlage fur die
Beurteilung des Nahrstozustandes der Boden. Kungl.
Landtbrukshogskolans Annaler. 26: 199215.
Ekeberg E. 1982. Eects of irrigation and fertilizer placement
on cereal growth. I. Yields and quality. Forsk. Fors. Landbr.
33: 99110.
Ekeberg E. and Riley H. 1995. The long-term fertilizer trials at
Mystad, S.E. Norway. SP report. 100th Anniversary
Workshop. Ministry of Agriculture and Fisheries, Danish
Institute of Plant and Soil Science. 29: 8397.
Elonen P. 1971. Particle-size analysis of soil. Acta Agriclia
Fennica 122: 1122.
Engels C. 1993. Dierences between maize and wheat in
growth-related nutrient demand and uptake. Plant Soil
150(1): 129138.
Engels C. and Marschner H. 1992. Root to shoot translocation
of macronutrients in relation to shoot demand in maize (Zea
mays L.) grown at dierent root zone temperatures. Z.
Panzenernahr. Bodenk. 155: 121128.
Equiza M.A., Mirave J.P. and Tognetti J.A. 2001. Morphological, anatomical and physiological response related to

158
dierential shoot vs. root growth inhibition at low temperature in spring and winter wheat. Ann. Bot. 87: 6776.
Forde B. and Lorenzo H. 2001. The nutritional control of root
development. Plant Soil 232: 5168.
Fohse D. and Jungk A. 1983. Inuence of phosphate and nitrate supply on root hair formation of rape, spinach and
tomato plants. Plant Soil 74: 359368.
Gahoonia T.S., Care D. and Nielsen N.E. 1997. Root hairs and
phosphorus acquisition of wheat and barley cultivars. Plant
Soil 191: 181188.
Gahoonia T.S., Nielsen N.E. and Lyshede O.B. 1999. Phosphorus (P) acquisition of cereal cultivars in the eld at three
levels of P fertilization. Plant Soil 211: 269281.
Goos R.J. and Johnson B.E. 2001. Response of spring wheat to
phosphorus and sulphur starter fertilizers to diering acidication potential. J. Agric. Sci. 136: 283289.
Hinsinger P. 1998. How do plant roots acquire mineral nutrients? Chemical processes involved in the rhizosphere. Adv.
Agron. 64: 225265.
Hofer R.M. 1996. Root hairs. In: Waisel Y., Eshel A. and
Kafka U. (eds), Plant Roots The Hidden Half. Marcel
Dekker, Inc., New York, pp. 111126.
Itoh S. and Barber S.A. 1983. A numerical solution of whole
plant nutrient uptake for soilroot systems with root hairs.
Plant Soil 70: 403413.
Johnson N.C., Graham J.H. and Smith F.A. 1997. Functioning
of mycorrhizal associations along the mutualismparasitism
continuum. New Phytol. 135: 575585.
Jungk A. 2001. Root hairs and the acquisition of plant nutrients
from soil. J. Plant Nutr. Soil Sci. 164: 121129.
Ketcheson J.W. 1968. Eect of controlled air and soil temperature and starter fertilizer on growth and nutrient composition of corn (Zea mays L.). Soil Sci. Soc. Am. Pro. 32:
531534.
Kleemola J., Jarvi A. and Kauppila R. 1998. Placing nutrients
with seed. Kungl. Skogs- och Lantbruksakademiens Tidskr.
137(7): 9398.
Krasilniko G., Gahoonia T. and Nielsen N.E. 2003. Variation
in phosphorus uptake eciency by genotypes of cowpea
(Vigna unguiculata) due to dierences in root hair length and
induced rhizosphere processes. Plant Soil 251: 8391.
Kristoersen A.. and Riley H. 2005. Eects of soil compaction
and moisture regime on the root and shoot growth and phosphorus uptake of barley plants growing on soils with varying
phosphorus status. Nutr. Cycl. Agroecosyst. 72(2): 135146.
Kristoersen A.., Bakkegard M., Hoel B. and Tandsther H.
2004. Startgjdsling til korn- oppsummering av 6 ar med
forsk. In: Bakkegard M. (ed.), Jord- og plantekultur 2004.
Apelsvoll, pp.138149.
Krogstad T. 1992. Methods of Soil Analysis. Report 6/92.
Department of Soil and Water Sciences, Agricultural University of Norway, p. 32
Lauzon J.D. and Miller M.H. 1997. Comparative response of
corn and soybean to seed-placed phosphorus over a range of
soil test phosphorus. Commun. Soil Sci. Plant Anal. 28((35):
205215.
Lcs A.K. 2003. Studies of the Availability of soil Phosphorus
(P) and Potassium (K) in Organic Farming Systems, and of
Plant adaptions to low P- and K- availability. Ph.D. Thesis
29, NLH, Norway.

Lyngstad I. 1977. Radgjdsling til korn. Forsk i perioden

196675. Norges landgrukshgskole. Forsk. Fors. Landbr.
28: 159177.
Mackay A.D. and Barber S.A. 1984. Soil temperature eects on
root growth and phosphorus uptake by corn. Soil Sci. Soc.
Am. J. 48: 818823.
Malhi S.S., Nyborg M., Penney D.C., Kryzanowski L. and
Robertson J.A. 1993. Yield response of barley and rapeseed
to P fertilizer: Inuence of soil test P level and method of
placement. Commun. Soil Sci. Plant Anal. 24(12): 110.
Marschner H. 1995. Mineral Nutrition of Higher Plants.
Academic Press, London, pp. 889
Marschner H. and Dell B. 1994. Nutrient uptake in mycorrhizal
symbiosis. Plant Soil 159: 89102.
Miller M.H., Bates T.E., Singh D. and Baweja A.S. 1971.
Response of corn to a small amounts of fertilizer placed with
the seed: I. Greenhouse studies. Agron. J. 63: 365368.
Moorby H. and Nye P.H. 1984. The eect of temperature
variation over the root system on root extension and phosphate uptake by rape. Plant Soil 78: 283293.
Murphy J. and Riley J.P. 1962. A modied single solution
method for the determination of phosphate in natural waters.
Anal. Chim. Acta 26: 3136.
Mberg J.P. and Petersen L. 1982. velsesvejledning til geologi og jordbundslre., Den kgl. Veterinr og Landbohgskole Part 2. p.136.
Nielsen K.F., Halstead R.L., MacLean A.J., Holmes R.M. and
Bourget S.J. 1960. The inuence of soil temperature on the
growth and mineral composition of oats. Can. J. Soil Sci. 40:
255263.
Olsen S.R., Cole C.V., Watanabe F.S. and Dean L.A. 1954.
Estimation of Available Phosphorus in Soil by Extraction
with NaHCO3. US Department Agric. Circ. 939 pp.
Pettersson S. 1995. Low root zone temperature eects on net
mineral nutrient uptake and distribution in barley (Hordeum
vulgare). J. Plant Physiol. 145: 459464.
Pratt P.F. 1965. Potassium. In: Black C.R. (eds), Methods of
Soil Analysis Part 2: Chemical and Microbiological Properties. American Society of Agronomy, Madison, Wisconsin,
USA, pp. 10231031.
Randall G.W. and Hoeft R.G. 1988. Placement methods for
improved eciency of P and K fertilizer: a review. J. Prod.
Agric. 1(1): 7079.
Rhoads F.M. and Wright D.L. 1998. Root mass as a determinant of corn hybrid response to starter fertilizer. J. Plant
Nutr. 21(8): 17431751.
Riley H. 1983. Soil physical properties of clay and silt soils.
Eects of organic matter content and soil conditioners.
Forsk. Fors. Landbr. 34: 155165.
Riley H. 1996. Estimation of physical properties of cultivated
soils in southeast Norway from readily available soil information. Nor. J. Agri. Sci. 25: 151.
Schenk M.K. and Barber S.A. 1979. Phosphate uptake by corn
as aected by soil characteristics and root morphology. Soil
Sci. Soc. Am. J. 43: 880883.
Singh B.R. and Subramaniam V. 1997. Phosphorus supplying capacity of heavily fertilized soils. II. Dry matter
yield of successive crops and phosphorus uptake at different temperatures. Nutr. Cycl. Agroecosyst. 47: 123
134.

159
Smith S.E., Gianinazzi-Pearson V., Koide R. and Cairney
J.W.G. 1994. Nutrient transport in mycorrhizas: structure,
physiology and consequences for eciency of the symbiosis.
Plant Soil 159: 103113.
Stribley D.P., Tinker P.B. and Rayner J.H. 1980. Relation of
internal phosphorus concentration and plant weight in plants
infected by vesicular-arbuscular mycorrhizas. New Phytol.
86: 261266.

Uhlen G. and Rd D. 1983. Frame experiments in comparison

with pot- and eld experiments for measuring phosphorus
and potassium fertilizer responses (in Norwegian). Meld. fra
Norges l.hgskole 62(23): 114.
Zadoks J.C., Chang T.T. and Konzak C.F. 1974. A decimal
code for the growth stages of cereals. Weed Res. (Oxford) 14:
415421.