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Many plant species show positive growth response to sodium. The effect of sodium,
however, has been ascribed to a poor supply of potassium, that is, sodium can in part compensate
for the shortage of potassium. (Flowers and Lauchli 1983). Brownell and Wood (1957) first
demonstrated the essentiality of sodium for the growth of Atriplex vesicaria, and their group has
done extensive studies on the function of this element. Brownell and Crossland (1972, 1974)
demonstrated that the sodium requirement is remarkable only in C4 and CAM plants, and
not in C3 plants. In two genera of Chenopodiaceae, Atriplex and Kochia, which contain both
C3 and C4 species, sodium was shown to be essential only for the C4 species (Brownell 1979).
In this paper, we report the sodium requirement of the NAD-malic enzyme type C4 plant
Amarantus tricolor L. cv. Tricolor, and discuss the possible functions of this element.
187
188 T. Matoh, D. Ohta and E. Takahashi
for calcium nitrate. The micronutrient composition was that of Arnon's solution cited in
Hewitt (1966) except that all the iron was supplied as ferrous-citrate. The culture solutions
were prepared using distilled and deionized water. Sodium was supplied as NaCl or Na2SO4
and in the reference treatment, KC1 or K2SO4 was supplemented to give the same anion con-
centration. Every pot had four or two seedlings and the solution was changed every four or
two days, as the plant growth proceeded.
The growth experiments were carried out for four times in the summer of 1983 and 1984.
In the 1983 experiments, the plants were allowed to grow until their seed formation stage.
The sodium-deprived plants had only about 20% seeds compared with the sodium-supplied
plants, but the germination efficiency was not different.
Analyses—The plants were dissected into leaves, stems and roots, and the parts were washed
with distilled water, blotted dry, weighed and dried in an oven at 70°C. The materials were
finely ground using a ball mill and appropriate amounts were digested with a nitric acid-sulfulic
Results
Growth—Dry matter production by A. tricolor under various growth conditions is shown in
Fig. 1. Growth of the plants receiving sodium salts was about 300% of those grown without
sodium salts. As the reference plants received potassium salts instead of sodium salts to give
the same anion concentrations, the growth stimulation induced by the application of sodium
salts is a sodium specific effect and is due neither to compensation of potassium shortage nor to
the accompanying anion.
Mineral contents—Mineral contents of A. triclor plants grown under various conditions are
shown in Tables 1 and 2. For elements other than Na and K, only the contents in the leaves
are presented. The plants supplied with K2SO4 at 0.5 mM showed poorer growth (Fig. 1) and
lower chlorine and potassium contents (Table 1) than the sodium-deprived plants. Sodium
contents in the leaves increased 40- to 50-fold with addition of sodium salts at 0.5 or 1 mM, but
the distribution of this element among the organs was almost the same as in the sodium-deprived
Table 1 Contents of sodium, potassium and chlorine in A. tricolor plants under various conditions
Na(ppm) K (%) Cl (ppm)
Leaves Stems Roots Leaves Stems Roots Leaves
Standard 32.6 48.9 110 6.69 12.7 9.03 5,940
+0.5 mM KC1 41.8 54.3 93.9 7.30 13.9 11.3 6,740
+1 mM KC1 32.5 38.9 68.8 6.53 12.9 10.8 7,070
+0.5 mM K2SO4 59.6 56.2 54.3 4.38 7.46 5.58 3,350
+0.5 mM NaCl 1,700 6,500 6,930 4.80 9.26 5.19 9,150
+ 1 mM NaCl 2,430 10,500 6,300 4.08 7.25 5.29 8,570
+0.5 mM Na 2 SO 4 2,650 12,400 7,520 4.30 9.16 5.86 8,220
The values are the means of three replicates.
Sodium requirement of Amaranthus tricolor 189
Standard
+ 0.5 mM KCI
+ I mM KCI
+ 0.5 mM K2SO«
+ 0.5 mM NaCI
+ I mM NaCI
+ 0.5 mM No2SO4
Fig. 1 Dry matter production of Amaranthus tricolor plants grown with or without sodium salts. The seeds were
sown on May 4, 1984. Sodium salts were given for 27 days and the plants were harvested on July 9, 1984. The
results are shown as an average of three plants and the bars show standard deviations. The growth experiment was
repeated three times with similar results.
Fig. 2 Acidification of the Amaranthus tricolor rooting medium by sodium salt application. The medium pH
changes were traced in the 1984 growing period using the same plants as in Fig. 1. Determinations were done from
the 17th day after sodium salt application. Standard culture solution (-O-); standard culture solution containing
0.5 meq./liter chloride salts ( ) of sodium (•) and potassium (O), the same at 1 meq./liter cation concentration
( ); the same at 1 meq./liter cation concentration, but given as sulfate salts ( ).
plants. A. tricolor plants responded to sodium but the sodium content and distribution in the
sodium-supplied plants are not different from those of other crop plants grown under ordinary
conditions, for which the sodium requirement has not been reported (Flowers and Lauchli 1983).
The chlorine content (Table 1) was somewhat higher in the sodium-recieving plants, but even
that of the reference plants far exceeded the critical content for chlorine deficiency (Broyer et al.
1954).
Comparing the mineral contents between + NaCI and +KC1 plants, significant differences
were found also for N, K and Ca contents. Nitrogen and potassium contents were higher in
the sodium-deprived plants. These results suggest that sodium improves the efficiency of
Table 2 Contents of nitrogen, phosphorus, calcium, magnesium and iron in A. tricolor plant leaves under various
conditions
Culture conditions N (%) P (%) Ca (%) Mg (%) Fe (ppm)
Standard 5.91 ±0.249 0.743 ±0.010 2.84±0.100 1.12 ±0.058 128±5.50
+0.5 mM KCI 6.20±0.370 0.855 ±0.058 2.85±0.064 1.21 ±0.067 N.D.
+1 mM KCI 5.99±0.050 0.807 ±0.071 2.54±0.50 1.08 ±0.130 249±12.4
+0.5 mM K2SO4 5.50±0.189 0.509±0.044 3.33±0.162 1.01 ±0.025 213±10.0
+0.5 mM NaCI 5.20±0.161 0.722 ±0.058 3.01 ±0.020 1.08 ±0.052 N.D.
+1 mM NaCI 5.28±0.I17 0.628 ±0.023 3.20±0.016 1.12 ±0.027 331 ±12.5
+0.5 mM Na2SO4 5.I0±0.U6 0.641 ±0.006 3.17 ±0.067 1.16±0.063 205 ±4.49
The values are the means of three replicates±standard errors. N.D., Not determined.
190 T. Matoh, D. Ohta and E. Takahashi
" 0.5 DIM KC1 or 0.5 HIM NaCl was supplemented to the standard culture solution.
* Leaves just reaching their maximum sizes were used.
Determinations were carried out six times using different plants.
Discussion
Under our growth conditions, the contaminating sodium in the culture solutions was less
than 20 ppb, and the sodium content in the leaves of plants grown without added sodium were
30 to 60 ppm (Table 1) although in the steins and roots they were generally higher. According
to Brownell (1979), his group reduced the concentration of contaminating sodium in the culture
solutions to less than 2 ppb, but the sodium content in their Atriplex vesicaria leaves, showing
sodium-deficiency symptom, was 10 mmol/kg dry matter, equaling 230 ppm. Kushizaki and
Yasuda (1964) reported that when sugar beet plants showed sodium-deficiency symptoms, the
internal sodium content was 6 mmol/kg dry leaves. Comparing these findings indicates that
the critical concentration of sodium at which plants suffer sodium deficiency may differ from
species to species.
Brownell (1979) also pointed out that the sodium requirement of the Ci-Atriplex plants was
met by application of about 0.1 meq./liter sodium. Our results (Fig. 1) indicate that application
of 0.5 meq./liter of sodium satisfies the sodium requirement of A. tricolor but the precise threshold
concentration of sodium for the plants was not determined. Assuming that all the sodium in
the leaves was free, the sodium concentrations were calculated to be about 0.2 nut in the sodium-
deprived leaves and about 10 DIM in the sodium-supplied leaves on the tissue water basis.
Sodium requirement of Amaranthus tricolor 191
Accordingly, if the system(s) which requires sodium occurs in the leaves, the internal threshold
concentration of sodium is in a mM range.
The physiological ground for the sodium requirement has never been made clear. Nable
and Brownell (1984) reported that in the sodium-deficient Amaranthus leaves, alanine accumula-
tion is significant compared with the contents in the sodium-sufficient plant leaves. They suggest
that sodium is required for in vivo activation of pyruvate orthophosphate dikinase. However,
we are not aware of any report on the sodium requirement of maize and sugar cane, representative
C4-plants having photosynthesis of the NADP-malic enzyme type. Hewitt (1983) summarized
the sodium effect on the C4 plant species and suggested that aspartate-forming C4-plants respond
to sodium. Comparison of the photosynthetic capacities between sodium-supplied and -deprived
plants is now in progress in our laboratory.
Nunes et al. (1983) demonstrated a NaCl-stimulated proton efflux from sugar beet leaf discs.
When the leaf discs absorbed sodium, stoichiometric release of potassium and proton occurred
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