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Improved Mass Screening of

Tolerance to Iron Toxicity in
Rice by Lowering Temperature
of Culture Solution
a b
Akifumi Shimizu , Corinta Q. Guerta , Glenn B.
b a
Gregorio & Hiroshi Ikehashi
a
Laboratory of Plant Genetics and Breeding, College
of Bioresource Sciences, Nihon University, Fujisawa,
Japan
b
International Rice Research Institute (IRRI), Makati
City, Philippines
Published online: 14 Feb 2007.

To cite this article: Akifumi Shimizu , Corinta Q. Guerta , Glenn B. Gregorio & Hiroshi
Ikehashi (2005) Improved Mass Screening of Tolerance to Iron Toxicity in Rice by
Lowering Temperature of Culture Solution, Journal of Plant Nutrition, 28:9, 1481-1493

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 Journal of Plant Nutrition, 28: 14811493, 2005
Copyright Taylor & Francis Inc.
ISSN: 0190-4167 print / 1532-4087 online
DOI: 10.1080/01904160500201352

Improved Mass Screening of Tolerance to Iron

Toxicity in Rice by Lowering Temperature
of Culture Solution
Downloaded by ["Queen's University Libraries, Kingston"] at 18:03 28 July 2013

Akifumi Shimizu,1 Corinta Q. Guerta,2 Glenn B. Gregorio,2

and Hiroshi Ikehashi1

1
Laboratory of Plant Genetics and Breeding, College of Bioresource Sciences,
Nihon University, Fujisawa, Japan
2
International Rice Research Institute (IRRI), Makati City, Philippines

ABSTRACT

Breeding for tolerance to iron (Fe) toxicity in rice (Oryza sativa L.) is hindered by
lack of a suitable screening technique. In the culture solution methods used to date, a
major difficulty has been maintaining an excess level of iron concentration in order to
reveal toxicity symptoms. Experimental results showed that this problem was solved
by lowering the solution temperature to around 20 C, at which leaf discoloration in
susceptible cultivars became more pronounced. At 20 C Fe uptake was increased from
a threshhold content of 300 mg/kg of dry weight of shoot for toxic symptom to more
than 1000 mg/kg in susceptible and in tolerant cultivars. Concentrations of other related
minerals in the plant tissue, i.e., potassium (K) and phosphorus (P), were not affected
by the low solution temperature itself but by Fe content, which can be inferred on the
basis of their response curves to the excess Fe treatment. Using the proposed screening
method makes it possible to obtain reproducible results in screening a large number of
plants or breeding lines.

Keywords: bronzing, iron toxicity, Oryza sativa, screening method, temperature of

culture solution

INTRODUCTION

Iron (Fe) toxicity in soil is a widespread problem for rice (Oryza sativa L.)
cultivation in India, South and Southeast Asia, South America, and West Africa

Received 3 November 2003; accepted 14 January 2005.

Address correspondence to Hiroshi Ikehashi, Laboratory of Plant Genetics and
Breeding, College of Bioresource Sciences, Nihon University, Kameino, Fujisawa
252-8510, Japan. E-mail: ikehashi@brs.nihon-u.ac.jp

1481
 1482 A. Shimizu et al.

(Mahadevappa et al., 1979; Ottow et al., 1991; Tanaka and Yoshida, 1970).
Leaves affected by a high level of iron (Fe) reveal brown spots on their tips
and finally turn entirely brown. This symptom is called bronzing and markedly
reduces grain yield of rice. Many attempts have been made to ameliorate such
soil conditions, but no sustainable method has been available in affected areas
where farmers cannot afford the necessary investment.
There are great intervarietal differences in iron toxicity tolerance in rice
(Gunawardena et al., 1982; Mohanty and Panda, 1991; Virmani, 1976), which
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many reports have confirmed to be a heritable trait by (Abifarin, 1985; Wu

et al., 1997). Thus, breeding cultivars tolerant to iron toxicity is considered
the most reliable means to avert it. To generate iron-toxicity-tolerant cultivars
most effectively, an efficient and reproducible method for mass screening is
necessary.
Few reports exist on the genetic analysis of iron toxicity in rice due to a lack
of such reproducible, rapid, and accurate screening methods. Mass screening
under culture solution is generally a useful method that allows stress conditions
to be easily controlled. For iron toxicity, however, maintaining an appropriate
level of stress is difficult due to an inevitable rapid decrease in dissolved Fe2+
concentration. To improve mass screening for iron toxicity, two factors that
lower the dissolved Fe2+ concentration should be taken into account. The first
is auto-oxidation of Fe2+ by water-dissolved oxygen. However, this problem
has been overcome by using EDTA-chelated iron (Fe-EDTA) as a source of
Fe (Fageria and Rabelo, 1987; IRRI, 1995). The second factor is the Fe2+ -
oxidizing power of rice root. Tadano (1974) reported that the Fe-excluding
power of rice roots at the seedling stage oxidized ferrous Fe not only at the root
surface but also in outer culture solution. According to Tadano (1975), such
an Fe-excluding power is stimulated by respiration. Therefore, lowering the
solution temperature is considered one of the most effective ways to depress
the oxidation by root and to promote Fe uptake into the rice plant.
In the present study, we investigated whether or not lowering the tempera-
ture of Fe-added solution with Fe-EDTA can make a further distinction between
tolerant and susceptible genotypes based on bronzing score. Iron, potassium (K),
and phosphorus (P) contents in the shoot were also measured to investigate the
effects of lowering temperature upon uptakes of Fe and other related minerals.
We tested this technique for an actual mass screening of F3 population.

MATERIALS AND METHODS

Basic Procedures

Germinated seeds were sown in holes of a polystyrene plate covered at the

bottom with a nylon net. The polystyrene plate was floated on a plastic tray
filled with deionized water for 4 d, after which the water was replaced with
 Mass Screening for Iron Toxicity 1483

Table 1
Bronzing score classified into seven ranks according to inspection
of leaf blades

Leaf order

Score 2nd 3rd 4th 5th 6th

1 N or T N N N N
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2 W N N N N
3 W P N N N
4 R R N N N
5 R R P N N
6 R R R P P
7 R R R R R

N: normal, T: discoloration of leaf tip, P: partly discolored, W:

whole leaf discoloration, R: rolled or dead leaf.

normal-strength Yoshidas solution (1976) including 1.78 mM silicon (Si) at

pH 5.0. After 5 d, the seedlings were exposed to iron toxicity by the addition of
ferrous Fe with 0.09 mM Fe-EDTA. The culture solution was renewed weekly,
and adjusted to pH 5.0 twice a day with 1 N NaOH/HCl. The seedlings were
grown until 30 d after sowing (six-leaf stage), recorded for bronzing score
according to Table 1, and harvested for measuring dry weight after being oven-
dried at 80 C for 48 h.

Test of Low-Solution Temperature for Mass Screening at IRRI

The experiment was arranged in a split-split plot design with three replications,
in which excess Fe factor, solution temperature, and cultivars were assigned
to the main plot, the subplot, and the sub-subplot, respectively. For each plot,
a plate with 10 10 holes was used in a 12 L solution. The experiment was
conducted in a greenhouse at the International Rice Research Institute (IRRI),
where the average air temperature was 30.2 C. Three levels of additional Fe so-
lution (0.13, 2.81, and 5.50 mM), including 0.09 mM Fe-EDTA and 0.04 mM
of Fe from Yoshidas solution and two levels of solution temperature (18 C
and 28 C), were compared. The 10 cultivars tested were six indica cultivars
(IR24, IR36, IR64, Suakoko8, BG90-2, and Kasalath) and four japon-
ica cultivars (Asominori, Gimbozu, Koshihikari, and Nekken2). Of these
cultivars, IR24 (Gunawardena et al., 1982), IR36 (Mahadevappa et al., 1979;
Gunawardena et al., 1982), Suakoko8 (Abifarin, 1985; IRRI, 1997; Virmani,
1976), IR74 (IRRI, 1997) had been reported as tolerant and IR36 (Virmani,
1976) and IR64 (IRRI, 1997) had been reported as susceptible. Reports on
IR36 are contradictory, which may be because of the differing methods of
 1484 A. Shimizu et al.

evaluation used, such as at a field in the Philippines (Mahadevappa et al., 1979),

by culture solution (Gunawardena et al., 1982), and at inland valley swamps
in Liberia (Virmani, 1978). Plastic trays were immersed in a temperature-
controlled water bath where water was circulated. Ground oven-dried samples
were digested with mixed acids and filtered with Whatman No. 42. Phosphorus
content was determined by the colorimetric method, and Fe and K contents
were determined by atomic absorption spectroscopy (Yoshida et al., 1976).
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Mass Screening of an F3 Population at Kyoto University

Eighty-five F3 lines from a cross between IR36 and Nekken2 were applied
to the mass screening of iron toxicity under 20 C culture solution. These par-
ents were previously tested with a pot experiment (Shimizu et al., 1996) and by
culture solution (unpublished), and showed different tolerance depending on
screening methods. IR36 was more tolerant than Nekken2 in a pot experi-
ment, but less tolerant than Nekken2 in culture solution. Five plants per line
were grown with two replications. For this screening, a polystyrene plate with
19 15 holes was used in a 35 L culture solution. This ratio of plant number
to solution volume is nearly equal to the ratio of 10 10 plants per a 12 L so-
lution in the previous experiment. After 30 d cultivation, the shoot dry weight
and bronzing score were recorded according to Table 1. The experiment was
conducted in a greenhouse at Kyoto University, and the average air temperature
was 28 C.

Statistical Procedures

Polynomial regression ANOVA was used to select a better-adapted regression

model for a response curve of a measured trait to three levels of Fe supply.
Among zero to two-degree polynomials, the best-fitted one with least squares
was selected based on the coefficients of determination (R 2 ) after being adjusted
for the degrees of freedom. Box-Cox conversion was used to find the transfor-
mation, which satisfies the condition of homogeneity of variance (Box and
Cox, 1964). As there was no difference within cultivars and/or within tempera-
ture treatment, data pooling was conducted and response curves were grouped
together.

RESULTS

Iron Toxicity Screening with Low-Solution Temperature

The response curve of bronzing score to Fe levels showed quantitative gradation,

although the score was given on a non-parametric scale (Figure 1). At 18 C of
 Mass Screening for Iron Toxicity 1485
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Figure 1. Response curves of bronzing scores at two solution temperatures. Formulas

for the curves were (a) at 28 C: Kasalath, y = 0.29 x + 2.8(r 2 = 0.79); others, y =
0.096x 2 0.47x + 2.9(r 2 = 0.51) and (b) at 18 C: Kasalath, y = 0.033x 2 + 0.81x
+ 2.8(r 2 = 0.96); IR64, y = 0.037x 2 + 0.86x + 1.0(r 2 = 0.99); BG90-2, y =
0.081x 2 + 0.77x + 1.3(r 2 = 0.90); others, y = 0.049x 2 + 0.73x + 1.3(r 2 = 0.93).

culture solution in all cultivars, the bronzing score increased with an increase of
Fe supply, while at 28 C of culture solution only the variety Kasalath showed
a higher score. Shoot dry weights and root dry weights of the tested varieties
also were adversely affected by iron toxicity (Figures 2 and 3), except for root
dry weight under 28 C of culture solution.

Change of Fe Content at Low-Solution Temperature and Under Excess Fe

There was no significant intervarietal difference in the Fe content of shoots

at 28 C of culture solution, while at 18 C of culture solution, the susceptible
 1486 A. Shimizu et al.
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Figure 2. Response curves of shoot dry weight (log-transformed) at two solution tem-
peratures. Scales of vertical axis in figure are re-transformed to original scales. Formulas
for the curves were (a) at 28 C: Kasalath, log(y) = 0.17x + 1.6(r 2 = 0.97); Gim-
bozu, log(y) = 0.052x + 0.91(r 2 = 0.68); IR74, log(y) = 0.013x 2 0.20x +
9.5(r 2 = 0.98); others, log(y) = 0.13x + 1.3(r 2 = 0.86) and (b) at 18 C: Kasalath,
log(y) = 0.029x 2 + 0.40x + 1.7(r 2 = 0.99); IR74, log(y) = 0.018x 2 0.21x +
1.1(r 2 = 0.91): others, log(y) = 0.021x 2 + 0.23x + 1.3(r 2 = 0.92).

cultivar Kasalath was differentiated from other cultivars (Figure 4). Iron con-
tents of Kasalath and other cultivars increased quadratically with the increased
Fe supply. The mean Fe content of Kasalath and other cultivars were increased
179% and 224%, respectively, in response to increased Fe supply of from 0.13
mM to 2.81 mM, and were increased 1093% and 281%, respectively, in response
to Fe supply of from 2.81 mM to 5.50 mM. Overall, the mean Fe content was
significantly increased at the lower temperature even under a low level of Fe
supply. For an instance, at the Fe supply level of 5.50 mM, the mean Fe content
of 489 133 mg/Kg at 28 C was one-third of mean Fe content of 1579 851
mg/Kg at 18 C.
 Mass Screening for Iron Toxicity 1487
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Figure 3. Response curves of root dry weight at two solution temperatures. Formulas
for the curves were (a) at 28 C: Kasalath, y = 0.017x 2 0.045x + 2.4(r 2 = 0.89);
BG90-2, IR24, and IR64, y = 0.10x 2 + 0.085x + 0.91(r 2 = 0.27); IR36, IR74,
and Nekken2 y = 1.4(0.35s.d.); others, y = 0.044x + 1.7 (r 2 = 0.16) and (b) at
18 C: Kasalath, y = 0.27x + 1.7(r 2 = 0.96); BG90-2 and IR24, y = 0.099x +
1.2(r 2 = 0.85); IR36, IR64, and Nekken2, y = 0.12x + 1.4(r 2 = 0.76); others,
y = 0.064x + 1.0(r 2 = 0.74).

Change in Other Nutritional Contents at Low-Solution Temperature

and Under Excess Fe

There was no significant intervarietal difference in P content at the two solution

temperatures (Figure 5). The P content decreased with the increase of Fe supply.
The P content was reduced by 69% in response to Fe supply of from 0.13
mM to 2.81 mM, and by 9% in response to iron supply of from 2.81 mM to
5.50 mM.
Potassium content at 18 C of culture solution decreased with the increase of
Fe supply (Figure 6). The mean K content was reduced in response to increased
 1488 A. Shimizu et al.
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Figure 4. Response curves of Fe contents (log-transformed) at two solution tempera-

tures. Scales of vertical axis in figure are re-transformed to original scales. Formulas for
the curves were (a) at 28 C: log(y) = 0.11x + 5.6(r 2 = 0.48) and (b) at 18 C: Kasalath,
log(y) = 0.13x 2 0.17x + 5.2(r 2 = 0.96); others, log(y) = 0.34x + 5.3(r 2 = 0.90).

Fe supply of from 0.13 mM to 2.81 mM by 14% in Gimbozu, by 15% in

Asominori and Koshihikari, and by 30% in the other cultivars. The mean K
content was also reduced in response to increased Fe supply of from 2.81 mM
to 5.50 mM for Gimbozu by 13%, Asominori and Koshihikari by 8%, and
for the others by 5%. At 28 C of culture solution, the K content did not decrease
with the increase of Fe supply.

Mass Screening

More than 3000 plants were screened at once by the culture solution, which was
controlled at 20 C. The scores in Table 1 can be roughly classified from tolerant
 Mass Screening for Iron Toxicity 1489
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Figure 5. Response curves of P contents (log-transformed) at two solution temperatures.

Scales of vertical axis in figure are re-transformed to original scales. There was no
significant difference among cultivars between results at 18 C and at 28 C. The formula
for response curve is log(y) = 0.011x 2 0.096x + 0.27(r 2 = 0.77).

(score 13, no rolled leaf) to moderately tolerant (score 4 to 5), to susceptible

(score 6 to 7, dying or almost dying). The distribution of bronzing score was
skewed to the tolerant side (Figure 7). The correlation coefficient (r ) between
replications of bronzing score was significant at the 1% level (r = 0.64). This
result was slightly lower than the r for shoot dry weight, 0.79.

DISCUSSION

Rice is one of the plants well-adapted to excess Fe environments (Snowden

and Wheeler, 1995). The oxidative activity of their roots is one of the factors
responsible for their tolerance to an excess of reduced Fe in waterlogged soils.
Although it still remains unknown which factor engenders the intervarietal dif-
ference in iron toxicity tolerance, the oxidative power or Fe2+ exclusion of
rice root (Ando et al., 1983; Chen et al., 1980; Green and Etherington, 1977;
Jayawardena et al., 1977; Luo et al., 1997), cannot be disregarded when eval-
uating the excess Fe tolerance in rice. Many measures to depress the oxidative
power of root have been tested, e.g., addition of toxic chemicals such as NaN3
and KCN (Tanaka and Tadano, 1969), or physical excision such as removing
roots and detaching leaves (Tanaka et al., 1966; Yamauchi and Peng, 1995).
However, all these measures are too deleterious on plant growth to be used in
precise evaluation of tolerance to excess Fe. Ando et al. (1983) pointed out
that lowering solution temperature was an effective way to reduce the oxida-
tion power of rice root. Also, in the present study, leaf bronzing and Fe uptake
were promoted under the high-level iron supply of 5.50 mM under the lowered
 1490 A. Shimizu et al.
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Figure 6. Response curves of K contents at two solution temperatures. Formulas for

the curves were (a) at 28 C: Kasalath, y = 0.16x 2 + 1.2x + 3.4(r 2 = 0.85);
Asominori, y = 2.9; others, y = 0.086x 2 0.48x + 3.2(r 2 = 0.32) and (b) at 18 C:
Gimbozu, y = 0.17x + 3.7(r 2 = 0.86); Asominori and Koshihikari, y = 0.016x 2
0.21x + 3.0(r 2 = 0.75); others, y = 0.068x 2 0.62x + 3.8(r 2 = 0.74).

solution temperature at around 20 C (Figures 1 and 4). In response to the in-

creased Fe supply, the shoot dry weight decreased at both of 18 C and 28 C
of solution temperatures (Figure 2), while root weight decreased only at 18 C
(Figure 3). When the solution temperature was lowered, the linear reduction
of root dry weight appeared in response to increased iron supply. This result
demonstrates the effectiveness of lowering solution temperature to suppress
the Fe-excluding power of rice root and to reduce linearly root dry weight in
parallel to the increase of Fe supply (Figure 3b compared to 3a).
The present study also shows that the lowering of solution temperature
makes it possible for rice plants to absorb excess Fe even under a high plant
density. At an Fe supply level of 5.50 mM, the mean iron contents of Kasalath
 Mass Screening for Iron Toxicity 1491
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Figure 7. Distribution of bronzing score in F3 lines from a cross between IR36 and
Nekken 2. Five plants each per line (total 85 lines) were exposed to 10.9 mM FeSO4 .
After 30 d bronzing scores were recorded according to Table 1. Horizontal bars show
ranges of bronzing scores of parental cultivars.

and other cultivars at 18 C of culture solution was 713% and 284% larger,
respectively, than the contents at 28 C of culture solution. A high level of Fe
accumulation was found in Kasalath at the 5.50 mM iron supply at two solution
temperatures. This result seems to be attributable not to the solution temperature,
but to the extreme susceptibility of this cultivar to excess Fe. Kasalath showed a
severe bronzing symptom even at 2.81 mM of Fe supply. The mean Fe content of
Kasalath at 2.81 mM Fe supply was higher than that of the second susceptible
variety, IR64, at 5.50 mM iron supply. This result implies that Kasalath has
a genetic factor that controls Fe toxicity susceptibility differently from other
cultivars.
There was no significant difference in P contents of shoots between 28 C
and 18 C of culture solution. According to Takahashi et al. (1955), P2 O5 ab-
sorption was highly inhibited by a low water temperature of 16 C. Since the
addition of excess Fe promotes Fe-P precipitation in the solution, the resulting
decrease of P2 O5 concentration in the culture solution may have inhibited the
P uptake. Accumulated Fe in the plant may also have inhibited P uptake. The
power of P-uptake inhibition by the accumulated Fe in the plant may have been
so strong that there was no significant intervarietal difference in P content.
Potassium content was not affected by the Fe content at 28 C. At 18 C
of culture solution, K content was not reduced in 0.13 mM iron supply, but
monotonically decreased with increased Fe supply. The resulting K content
was still much higher than the marginal K content of 1% (Yoshida et al., 1976).
The reduction in K uptake was caused not by the low solution temperature but
 1492 A. Shimizu et al.

by the increased uptake of excess Fe. The intervarietal difference in K content

was different from that of Fe content.
In summary, it was possible to observe clearly intervarietal differences in
iron toxicity tolerance under the low temperature, at which P and K content
was not directly affected. The low solution temperature seemed uniformly to
restrain the Fe-excluding power of rice roots under excess Fe supply (5.50
mM) and to differentiate clearly intervarietal tolerance to Fe toxicity in a test of
a large number of cultivars or genotypes. Therefore, by using of this method, it
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is possible to obtain reproducible results in screening a large number of plants

or genotypes in genetic analysis.

ACKNOWLEDGMENTS

The authors dedicate this work to the memory of Dr. Senadhira. This research
was funded by the Japan-IRRI Shuttle Research Project.

REFERENCES

Abifarin, A. O. 1985. Inheritance of tolerance to iron toxicity in two rice cul-

tivars. In Rice genetics, 423427. Manila, International Rice Research In-
stitute (IRRI).
Ando, T., S. Yoshida, and I. Nishiyama. 1983. Nature of oxidizing power of
rice roots. Plant and Soil 72: 5771.
Box, G. E. P., and D. R. Cox. 1964. An analysis of transformations. Journal of
the Royal Statistical Society: Series B 26: 211252.
Chen, C. C., J. B. Dixon, and F. T. Turner. 1980. Iron coatings on rice roots: Min-
eralogy and quantity influencing factors. Soil Science Society of America
Journal 44: 635639.
Fageria, N. K., and N. A. Rabelo. 1987. Tolerance of rice cultivars to iron
toxicity. Journal of Plant Nutrition 10: 653661.
Green, M. S., and J. R. Etherington. 1977. Oxidation of ferrous iron by rice
(Oryza sativa L.) roots a mechanism for water logging tolerance. Journal
of Experimental Botany 28: 678690.
Gunawardena, I., S. S. Virmani, and F. J. Sumo. 1982. Breeding rice for tolerance
to iron toxicity. Oryza 19: 512.
IRRI. 1995. Program report for 1995. Manila: IRRI.
IRRI. 1997. Program report for 1997. Manila: IRRI.
Jayawardena, S. D. G., T. Watabe, and K. Tanaka. 1977. Relation between root
oxidizing power and resistance to iron toxicity in rice. Rep. Soc. Crop Sci.
Breed. Kinki, Jpn. 22: 3847.
Luo, A. C., G. Jing, P., P. Wu, J. J. Ni, S. H. Jing, and Y. S. Zhang. 1997.
Rice genotype differences in nutrient status under excessive ferric-iron
conditions. Journal of Plant Nutrition 20: 13611373.
 Mass Screening for Iron Toxicity 1493

Mahadevappa, M., H. Ikehashi, and F. N. Ponnamperuma. 1979. The contribu-

tion of varietal tolerance for problem soils to yield stability in rice, IRRI
Research Paper Series No. 43. Manila: IRRI.
Mohanty, S. K., and K. Panda. 1991. Varietal behaviour of rice towards Fe
toxicity. Oryza 28: 513515.
Ottow, J. C. G., K. Prade, W. Bertenbreiter, and V. A. Jacq. 1991. Strategies to
alleviate iron toxicity of wetland rice on acid sulphate soils. In Rice produc-
tion on acid soils of the tropics, eds. P. Deturck and F. N. Ponnamperuma,
Downloaded by ["Queen's University Libraries, Kingston"] at 18:03 28 July 2013

205211. Kandy, Sri Lanka: Inst. Fund. Studies.

Shimizu, A., J. Wan, and H. Ikehashi. 1996. Genetic markers for tolerance to
iron toxicity in rice. Rice Genetics Newsletter 13: 132134.
Snowden, R. E. D., and B. D. Wheeler. 1995. Chemical changes in selected
wetland plant species with increasing Fe supply, with specific reference to
root precipitates and Fe tolerance. New Phytologist 131: 503520.
Tadano, T. 1974. Studies on the iron nutrition of rice plant. 5. Change of the
susceptibility to iron toxicity during the growth. Journal of Sci. Soil Manure
Jpn 45: 521524 (in Japanese).
Tadano, T. 1975. Devices of rice roots to tolerate high iron concentration in
growth media. JARQ 9: 3439.
Takahashi, J., M. Yanagisawa, M. Kono, F. Yazawa, and T. Yoshida. 1955.
Studies on nutrient absorption by crops. Bull. Natl. Inst. Agric. Sci. B. 4:
183 (in Japanese).
Tanaka, A., R. Loe, and S. A. Navasero. 1966. Some mechanism involved in
the development of iron toxicity symptom in the rice plant. Soil Science
and Plant Nutrition 12: 3238.
Tanaka, A., and T. Tadano. 1969. Studies on the iron nutrition of rice plant. Part
2. iron excluding capacity of the rice roots. J. Sci. Soil Manure Jpn. 40:
469472.
Tanaka, A., and S. Yoshida. 1970. Nutritional disorder of the rice plant in Asia,
IRRI Technical Bulletin 10. Manila: IRRI.
Virmani, S. S. 1976. Breeding rice for tolerance to iron toxicity. In Second rice
varietal improvement seminar. Bouake, Cote dlvoire: WARDA: 120.
Virmani, S. S. 1978. Some results of varietal screening for tolerance to iron
toxicity. In International rice research conference, Manila, Philippines:
IRRI.
Wu, P., A. Luo, J. Zhu, J. Yang, N. Huang, and D. Senadhira. 1997. Molecular
markers linked to genes underlying seedling tolerance for ferrous iron
toxicity. Plant and Soil 196: 317320.
Yamauchi, M., and X. X. Peng. 1995. Iron toxicity and stress-induced ethylene
production in rice leaves. Plant and Soil 173: 2128.
Yoshida, S., D. A. Forno, J. H. Cock, and K. A. Gomez. 1976. Laboratory
manual for physiological studies of rice. Manila: IRRI: 19.