Beruflich Dokumente
Kultur Dokumente
2016, 000, 19
DOI: 10.1002/jpln.201500625
School of Agriculture, Food and Wine, Waite Research Institute, University of Adelaide, Urrbrae, SA 5064, Australia
Australian Centre for Plant Functional Genomics, Waite Research Institute, Urrbrae, SA 5064, Australia
3 King Abdullah University of Science and Technology, Center for Desert Agriculture, Thuwal, Kingdom of Saudi Arabia, 23955-6900
2
Abstract
Nitrate (NO
3 ) and ammonium (NH4 ) are the predominant forms of nitrogen (N) available to
plants in agricultural soils. Nitrate concentrations are generally ten times higher than those of
NH
4 and this ratio is consistent across a wide range of soil types. The possible contribution of
these small concentrations of NH
4 to the overall N budget of crop plants is often overlooked. In
this study the importance of this for the growth and nitrogen budget of maize (Zea mays L.) was
investigated, using agriculturally relevant concentrations of NH
4 . Maize inbred line B73 was
grown hydroponically for 30 d at low (0.5 mM) and sufficient (2.5 mM) levels of NO
3 . Ammonium
levels, addition of NH
was added at 0.05 mM and 0.25 mM to both levels of NO
.
At
low
NO
4
3
3
was found to improve the growth of maize plants. This increased plant growth was accompanied
by an increase in total N uptake, as well as total phosphorus, sulphur and other micronutrients in
the shoot. Ammonium influx was higher than NO
3 influx for all the plants and decreased as the
total N in the nutrient medium increased. This study shows that agriculturally relevant proportions
of NH
4 supplied in addition to NO3 can increase growth of maize.
1 Introduction
Nitrogen (N) is one of the major nutrients required by plants
with N limitation commonly leading to reduction in both growth
and yield. Plants absorb N mainly in the form of nitrate (NO
3)
and ammonium (NH
4 ) from the soil solution (Glass et al.,
2002). Nitrate is the dominant form present in agricultural
soils and, hence, is the focus of most research on N uptake
(Marschner, 2011). Although the NH
4 concentration in the
soil solution is small ( 10%) compared to NO
3 , this ratio is
consistent in most agricultural soils (Miller and Cramer, 2005)
and could make a significant contribution to plant N uptake.
Previous studies have demonstrated that a combination of
NO
3 and NH4 is beneficial for plant growth (Haynes and
Goh, 1978; Bloom et al., 1993). This has also been shown in
maize (Schrader et al., 1972; Smiciklas and Below, 1992).
However, these studies were conducted using proportions of
NH
4 greater than 25% (Schrader et al., 1972), which is much
higher than the concentrations typically present in agricultural
soils. Cox and Reisenauer (1973) used small amounts of
NH
4 relative to NO3 and found that, even at these low proportions, growth of wheat was stimulated. Therefore, it is
important to understand whether agriculturally relevant proportions of NH
4 increase growth of maize and, if there is any
increase, why is this so.
the NO
2 then enters plastids where it is converted to NH4 ,
from which amino acids are formed (Bloom et al., 1992). This
additional processing means that NO
3 assimilation is a
more energy-intensive process [requiring 12 adenosine triphosphate (ATP) molecules] compared to NH
4 assimilation
(requiring two ATPs) (Schrader et al., 1972; Clarkson, 1985).
On this basis, it may be that energy conservation is one factor
underlying the increased plant growth observed with small
amounts of NH
4.
Another reason for the increased plant growth may be related
to the higher uptake capacity of NH
4 relative to NO3 when
both N sources are present. Previous studies have shown that
when both N forms are available, plants take up NH
4 preferentially over NO
(Hatch
and
Macduff,
1991;
Gazzarrini
et al.,
3
1999; Glass et al., 2002), and that the total N uptake of these
plants is increased. The reason for preferential uptake of NH
4
may be due to the faster assimilation of NH
4 in the roots (Lee
et al., 1992) by the enzymes cytosolic glutamate synthetase
(GS1) and glutamate synthase (GOGAT) resulting in a more
rapid incorporation of inorganic N into organic N.
This positive growth effect of NH
4 could also be due to effects
on other aspects of plant nutrition, given that in some cases
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NH
4 can improve the phosphorus (P), sulphur (S) (Kirkby,
1968; Flores et al., 2001), and micronutrient (Riley and Barber,
1971; Thomson et al., 1993; Jeong and Lee, 1996) nutrition of
plants. The uptake of NO
3 by plants can enhance the absorption of cations such as K+, Mg2+, and Ca2+, but can lead to
slower uptake of P, S, and some trace elements (Jackson and
Williams, 1968). Iron (Fe) deficiency has been observed in
plants grown solely with NO
3 compared to plants grown with
(Zou et al., 2001). On the other hand,
both NH
and
NO
4
3
when NH
4 is supplied to plants it can increase the uptake of
Fe from the nutrient solution, and the re-mobilisation of Fe inside the plant (Marschner et al., 1987; Zou et al., 2001).
The aim of this study was to compare and quantify the effect
of low and high levels of NH
4 on maize growth at both low
and sufficient levels of N. This study looked at the effect of
small levels of NH
4 on total N uptake, C : N ratio, NH4 and
NO3 influxes and on macro and micronutrient uptake. Maize
inbred line B73 was used as it is the source of the maize
reference genome and has been used in a large number of
physiological studies (Agrama et al., 1999; Martin et al.,
2006). In this manuscript, we report the effect of agriculturally
relevant proportions of NH
4 on the maize inbred line when
provided along with NO
.
3
(SN+LA), and 2.5 mM NO3 + 0.25 mM NH4 (SN+HA). All
these treatments contained different N concentrations. A second experiment was conducted in a similar manner, but the N
concentration remained the same in both treatments of this
experiment which consisted of 0.75 mM NO
3 alone (LN+0A)
Experiment
Treatments
NO
3 (mM)
NH
4 (mM)
Experiment 1
LN+LA
LN+HA
SN+LA
SN+HA
0.50
0.50
2.50
2.50
0.05
0.25
0.05
0.25
Experiment 2
LN+0A
LN+HA
0.75
0.50
0.25
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3 Results
3.1 Plant biomass
The shoot biomass of plants grown in low N was greater (at
30 DAE) with added NH
4 in the LN+HA treatment; however,
when grown with sufficient N, the plants showed no response
to NH
4 (Fig. 1A). Plants grown in higher concentration of
NH
4 showed no change in root biomass regardless of N
treatment (Fig. 1B), except on 23 DAE, which had a higher
root biomass in SN+LA compared to SN+HA. Plants grown in
LN+LA and LN+HA had a higher root : shoot ratio compared
to plants in both sufficient N treatments at 10 DAE (Fig. 1C).
Thereafter, this ratio decreased for plants in all treatments
and at final harvest, a decrease was observed in the root:
shoot of plants in LN+HA compared to LN+LA and both sufficient N treatments.
by the added NH
4 or the presence of high NH4 in LN+HA.
mixture of NO
3 and NH4 (LN+HA) had higher shoot dry matter (Fig. 2A) than plants grown in NO
3 alone (LN+0A). No difference in root dry matter was observed between treatments
(Fig. 2B). The root : shoot ratio of plants was higher with NH
4
at 10 DAE (Fig. 2C). Thereafter, the root : shoot ratio de
creased in plants treated with NH4 and was lower at 20 and
23 DAE than for plants grown with NO
3 alone.
Root DM / g
Root DM / g
Shoot DM / g
Shoot DM / g
Experiment 2 was conducted to determine whether the observed higher biomass of maize plants in LN+HA compared
Figure 1: Shoot dry matter, root dry matter, and root : shoot ratio of
maize inbred line B73 (A, B, C) grown in 0.5 mM NO
3 + 0.05 mM
NH
4 (LN+LA), 0.5 mM NO3 + 0.25 mM NH4 , (LN+HA), 2.5 mM NO3
+ 0.25 mM NH
+ 0.05 mM NH
(SN+LA),
and
2.5
mM
NO
4
4
3
(SN+HA). Values are means SE, n = 6. Significant differences for
individual harvest days are represented by different letters (P 5%).
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NO
3 and NH4 (LN+HA) had a higher shoot N concentration
(Fig. 4A) than plants grown in NO
3 alone (LN+0A). Total N
and net uptake relative to root dry matter content also showed
an increase in LN+HA compared to LN+0A (Fig. 4B, C). No
difference in the C : N ratio of plants was observed between
the two treatments.
The NH4 influx of all the plants was several fold higher in all
the treatment compared to their corresponding NO
3 influx
(Fig. 5). The NO
influx
was
lower
only
for
maize
plants
3
grown in SN+HA (Fig. 5). In comparison, NH
influx
de4
creased as N level in the medium increased.
4 Discussion
This study investigated the effect of agriculturally relevant
proportions of NH
4 and NO3 on the growth of maize. Maize
plants were grown in two levels of NO
3 , either low (0.5 mM)
or sufficient (2.5 mM) and with two levels of NH
4 (0.05 mM or
0.25 mM) for each NO
treatment
(Table
1).
These
results in3
dicate that shoot biomass of maize plants was increased
when more NH
4 was added to the low NO3 treatment
(Fig. 1A). In this experiment, the extra total N provided in the
added NH
4 may have contributed to the increased biomass
of the plants. However, the increased biomass in the low
NO
3 with high-NH4 treatment compared to NO3 -only treatment in Experiment 2 suggests that the increase in plant
C : N ratio
Net uptake N / g N g
root DW
Total N / g
Shoot N%
In Experiment 1, shoot sulphur (S) and phosphorus (P) concentrations in maize plants grown in LN+HA were higher than
for those grown in LN+LA (Fig. 6A). Increasing NH
4 concentration at low NO
levels
increased
the
tissue
concentration
3
Figure 3: Shoot N concentration, total N content, net uptake, and C : N ratio of the maize inbred line B73 (A, B,
C, D) grown in 0.5 mM NO
3 + 0.05 mM NH4 (LN+LA), 0.5 mM NO3 + 0.25 mM NH4 , (LN+HA), 2.5 mM NO3
+ 0.25 mM NH (SN+HA). Values are means SE, n = 6. Signifi+ 0.05 mM NH
(SN+LA),
and
2.5
mM
NO
4
4
3
cant differences are represented by different letters for each group of bars at individual harvest days (P 5%).
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Total N / g
C : N ratio
Shoot N%
Figure 4: Shoot N concentration (A) and net uptake (B) of the maize inbred line B73 grown in 0.75 mM NO
3 (LN+0A) and
0.5 mM NO
3 + 0.25 mM NH4 (LN+HA) on 23 DAE. Values are means SE (n = 8). Significant differences between treatments with *P 5%).
2.5 mM NO
3 + 0.05 mM NH4 (SN+LA), and 2.5 mM NO3 + 0.25 mM
enhanced macro and micro nutrient nutrition. Nitrogen assimilation is very closely related to C metabolism because C skeletons produced by photosynthesis are required for amino
acid synthesis (Kaiser and Forster, 1989; Pace et al., 1990).
The decreased C : N ratio and higher N concentration of
plants in low-NO
3 with high-NH4 is consistent with lower N
assimilation costs with added NH
4 compared to plants in low
NO
3 with low NH4 treatment. The lower root : shoot of NH4 treated plants throughout the growth period, except on day 10
in Experiment 2, suggests that less C is required for root proliferation in comparison to the increased shoot growth in
these plants. Increased biomass allocation to the shoot
compared to root increases carbon assimilation capacity and
plant relative growth rate (Van der Werf, 1996). This may explain the increased growth of plants in the low-N treatments
following the addition of NH
4.
Another hypothesis is that increased plant growth may result
from improved N status with the addition of NH
4 . This may be
due to a higher uptake capacity of NH
4 , which would result in
faster incorporation of N into an organic form and a corresponding increase in plant growth with high NH
4 concentration. For tomato plants provided with various ratios of radio
labeled NO
3 and NH4 , Glass et al. (2002) reported that 50%
of total N was from NH
4 even though it constituted only 10%
of the total N in the growth medium. Moreover, the extra NH
4
assimilated in the roots may have resulted in increased translocation of N assimilatesmainly amino acidsto the shoots,
resulting in increased shoot growth. This hypothesis has also
been proposed in earlier studies on maize (Alexander et al.,
1991; Gentry, 1992; Cramer and Lewis, 1993) and sorghum
(Lewis et al., 1982), in which the higher concentration of NH
4
in the growth solution was found to increase total N uptake.
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Macronutrients / mg kg
1
Micronutrients / mg kg
Figure 6: Macronutrient (A) and micronutrient (B) concentrations of the shoots of maize
inbred lineB73 grown in 0.5 mM NO
3 + 0.05 mM NH4 (LN+LA), 0.5 mM NO3 + 0.25 mM
NH4 , (LN+HA), 2.5 mM NO3 + 0.05 mM NH4 (SN+LA), and 2.5 mM NO
3 + 0.25 mM
NH
4 (SN+HA) on 30 DAE. Values are mean SE, n = 6. Significant differences are represented by different letters for each group of bars (P 5%).
and
NH
main NO
4 uptake systems in plants are the satura3
ble high affinity transport system (HATS) and non-saturable
low affinity transport system (LATS; Forde, 2000). In the cur
rent study, the influx via HATS of both NH
4 and NO3 played a
major role in the N uptake response of maize to N demand
(Malagoli et al., 2004; Garnett et al., 2013). The finding here
that NH
4 influx was higher than NO3 influx is consistent with
previous work (Lee and Rudge, 1986; Teyker et al., 1988;
Hole et al., 1990; Glass et al., 2002). Furthermore, it was
found that, even when the concentration of NO
3 is ten times
higher than that of NH
4 , plants have the tendency to absorb
NH
4 more rapidly than NO3 (Gessler et al., 1998).
The NH
4 influx of plants decreased as N levels in the growth
solution increased, indicating that these plants alter their influx based on total N supply. This result is expected as a number of studies show that nutrient deprivation augments the
transport capacity of the deficient ion (Clarkson et al., 1983;
NO
3 and NH4 was 12 : 2 and 10 : 4 (Flores et
al., 2001). Earlier studies suggested that enhanced P uptake in plants supplied with NH
4
may be due to acidification at the root surface
caused by absorption and assimilation of
NH
4 (Miller et al., 1970). Recent research indicated the involvement of plasma membrane H+ ATPase in the enhanced uptake of
P by rice roots supplemented with NH
4
(Zeng et al., 2012). The authors suggested
that plasma membrane H+ ATPase activity is
increased by low pH resulting from NH
4 nutrition which facilitates uptake.
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Macronutrients / mg kg1
Acknowledgments
Micronutrients / mg kg1
The authors would like to acknowledge the technical assistance provided by research and technical staff at Australian
Centre for Plant Functional Genomics (ACPFG) and Plant
Research Centre in the University of Adelaide. This study
was funded by the Australian Centre for Plant Functional Genomics (ACPFG), the Grains Research and Development
Corporation (GRDC), and Australian Research Council Linkage Grant (LP130101055).
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