Plant and Soil 69, 31-44 (1982). 0032-079X/82/0691-003152.10. Ms.

5126
9 1982 Martinus No'hoff/Dr W. Junk Publishers, The Hague. Printed in The Netherlands.

Influence of aluminum in nutrient solutions on chemical composition

in u p l a n d rice cultivars

N. K. FAGERIA and J. R. P. CARVALHO

National Rice and Bean Research Center, Caixa Postal 179-74000 Goifinia, Goids, Brazil

Key words Aluminum toxicity Chemical composition Oryza sativa L. Upland rice

Summary Aluminum toxicity is an important growth limiting factor for upland rice production on
oxisols of'cerrado' region in Brazil. Data related to the effect orAl on uptake of nutrients for rice crop
are limited. The effect of five AI concentrations (0, 10, 20, 40 and 60 ppm) in culture solution on the
chemical composition of 30 upland rice cultivars was studied.
Aluminum concentration and content in plant tissues were increased with higher levels of A1 in all
cultivar. In the roots AI content was higher as compared with the tops. Critical toxic level of A1 in the
tops of 21 days old plants varied from 100 to 417 ppm depending on the cultivars. Rice cultivars
responded differently to AI treatments with respect to nutrients uptake. Increased AI concentrations
in the solution exerted an inhibiting effect on the concentrations and contents of N, P, K, Ca, Mg, S,
Na, Zn, Fe, Mn, B and Cu. Thus the inhibition was more effective for macronutrients in the plant tops
in following order: Mg > Ca > P > K > N > S > Na. Whereas for micronutrients it was in the
order of Mn > Zn > Fe > Cu > B. Morphological, physiological and biochemical effects of AI,
toxicity responsible for the reduction in plant nutrient uptake, are discussed.

Introduction
Brazil is the world's largest producer of upland rice. About 22 per cent of the
total area of the Country is mainly covered by tropical savana, locally refered to
as the 'cerrado'.
Seventy percent of the rice production comes from cerrado predominating
central regions.
Upland rice in the cerrado region, may be exposed to many factors that are not
be found under lowland conditions, such as water stress, hot and dry climatic
conditions, and adverse soil conditions such as toxicity of aluminum and N, P,
Ca, Mg and Zn deficiency 5'21"23.
Several studies have shown modification in the mineral nutrition of plants
exerted by aluminum, which is absorbed in the free trivalent form in acidic soils.
The results were mostly restricted to P, K, Ca and Mg 1~ However,
information in this respect is limited in relation to other nutrients and especially
in relation to the rice crop.
Therefore, the objective of this investigation was to study the effect of A1 on the
concentration and content of nutrients in upland rice cultivars to understand
nutritional aspects of the rice crop under acidic soil conditions. The effect of Al on
growth of these cultivars was discussed in a separate paper 7.

31
32 FAGERIA AND CARVALHO

Materials and methods

An experiment was conducted in a glasshouse at the National Rice and Bean Research Center,
Goi~mia, Goi~s, to study influence of A1 on chemical composition in upland rice cultivars.
Seeds of 30 upland rice (Oryza sativa L.) cultivars were germinated in pure sand using plastic trays
of 30 x 45 x 8 cm. Eight to 10 days after sowing, 4 uniform seedlings were transplanted to acrilic
discs with holes in the center. The seedlings were held in place with cotton. These discs were then
transferred to plastic pots containing 7.5 liters of nutrient solution. Each pot had 3 discs with 12
plants.
With slight modifications the nutrient solutions used were those developed by the International
Rice Research Institute in the Philippines for rice 28. It had the following composition in I~M:
NH4NO3 2857; NaHzPO4.H20 129; KzSO 4 1023; CaCI 2 1000; MgSO4.7HzO 1645;
(NH4)6Mo7024.4H20 0.5; MnC12.4H2 9; H3BO 3 18.5; ZnSO4.5H20 0.15; CuSO 4 0.16 and
FeC13.6H20 36:AIC13in amounts required for A1concentration of 0; 371; 742; 1484; and 2226 ~tM (0,
10, 20, 40 and 60 ppm A1).The nutrient solutions were changed once a week. The pH of the solution
was adjusted to 4 + 0.2 initially and once every two days thereafter with 0.1 N NaOH or 0.1 N HCI.
The experiment was conducted in a randomized block design with two replications. Maximum and
minimum air temperature means during the experiment were 28 + 2~ and 18 + 2~ respectively.
After 21 days growth in AI treated solutions, plant tops and roots were harvested separately. Roots
were rinsed thoroughly in distilled water and blotted. Roots and tops were separated, dried to a
constant weight at 80~ and weighed. Chemical analysis of plant tissue was done at the Soil and Plant
Analysis Laboratory, University of Wisconsin, Madison, U.S.A.
Analysis for P, K, Ca, Mg, S, Na, Zn, Fe, Mn, B, Cu, and AI was done simultaneouslywith a plasma
emission spectrophotometer and N was determined on a separate sample via a semi-micro Kjeldahl
procedure 20.
Data are means of two replications. Regression analysis was used to express relationship between
AI concentrations and aluminum content in the plant tops.

Results and discussion

I n this p a p e r we h a v e used the t e r m s c o n c e n t r a t i o n a n d c o n t e n t c o n s i d e r i n g as

suggested b y F a r h o o m a n d a n d P e t e r s o n 9, these are the p r o p e r t e r m s to express
p l a n t s tissue a n a l y s i s results. T h e s e a u t h o r s p r o p o s e d the f o l l o w i n g d e f i n i t i o n s of
the t e r m s ' C o n c e n t r a t i o n ' a n d ' C o n t e n t ' .
C o n c e n t r a t i o n was defined as the a m o u n t of a p a r t i c u l a r e l e m e n t o r s u b s t a n c e
w h i c h is p r e s e n t in a n u n i t a m o u n t of a s o l u t i o n or c o m p o u n d (solid material);
w h e r e a s c o n t e n t was difined as the a m o u n t of a p a r t i c u l a r e l e m e n t o r s u b s t a n c e
p r e s e n t in a specific a m o u n t of a s o l u t i o n o r c o m p o n e n t . T h e f u n d a m e n t a l
d i s t i n c t i o n b e t w e e n these two t e r m s c a n best be seen b y a c o n s i d e r a t i o n of
i n t e n s i v e a n d extensive properties.
I n t e n s i v e p r o p e r t i e s are those w h i c h d o n o t d e p e n d u p o n the q u a n t i t y of the
h o m o g e n e o u s s u b s t a n c e u n d e r c o n s i d e r a t i o n ~3. These p r o p e r t i e s r e m a i n
c o n s t a n t t h r o u g h o u t a h o m o g e n e o u s s a m p l e regardless of its size. D e n s i t y ,
t e m p e r a t u r e , a n d color are e x a m p l e s of i n t e n s i v e properties. Extensive
properties, however, d e p e n d u p o n the q u a n t i t y of a h o m o g e n e o u s s u b s t a n c e ~a.
These p r o p e r t i e s o b v i o u s l y v a r y with the size of the sample. V o l u m e a n d
ALUMINUM IN UPLAND RICE CULTIVARS 33

weight are examples of extensive properties. It should be noted that extensive

properties are additive whereas intensive properties are not. Concentration is an
intensive property and will be the same regardless of the size of the sample,
whereas the content is an extensive property and will depend on the size of the
sample.

Concentration and content of aluminum as influenced by Al treatments

Regression equations and correlation coefficients between AI treatments and
content in the tops of rice cultivars are presented in Table 1. In most of the
cultivars a positive linear relationship was obtained with respect to
concentration of AI in solution and AI content in the plants tops as expected.

Table 1. Regression equations and correlation coefficients between aluminum levels and AI content in
the rice cultivars tops

Cultivar Regression equation Correlation coefficient

IAC 1246 "~ = 266,734483 + 24.524827x 0.98

IAC 47 ~= 229.001724 + 21.442241x 0.97
IAC 5544 ~= 355.410345 + 18.653448x 0.97
IAC 5100 ~, = 370.739655 + 17.611552x 0.95
IAC 1131 ~= 344.042672 + 29.267974x 0.93
IAC 25 "~ = 356.817241 + 24.727414x 0.98
MatZo Not significant
Pratao ~, = 244.959914 + 25.065388x 0.96
D o u r a d o Precoce Not significant
Prat~o Precoce ~, = 155.984483 + 18.959827x 0.99
M o n t a n h a liso ~, = 273.421552 + 26.088017x 0.93
Batatais "~ = 196.17112 + 20.174957x 0.97
Q u a t r o meses "~ = 161.732758 + 8.542586x 0.95
P~rola ~= 102.173707 + 12.088319x 0.99
Ponta Preta "~ = 190.723278 + 7.945258x 0.93
Ipeaco-163 "~ = 80.730603 + 14.002284x 0.98
Catal~o "~ = 72.026293 + 18.866681x 0.96
Ipeaco-462 "~ = 166.089655 + 12.086552x 0.95
IRAT 8 Not significant
I R A T 12 Not significant
IRAT 2 ~= 320.118497 - 11.368902x + 0.248274x 2 0.97
Ipeaco 162 "~ = 426.717057 - 17.648515x + 0.632434x 2 0.98
Ipeaco 562 Not significant
Paulistinha Not significant
Fernandes "~ = 191.272414 + 4,679138x 0.93
I R A T 11 ~' = 146.925862 - 8,99361x 0.96
I R A T 13 "~ = 143.119396 + 6.537715x 0.98
Esa6 Not significant -
Bico G a n g a Not significant
M R C 172-9 "~ = 240.507759 + 5.680086x 0.91
 34 FAGERIA AND CARVALHO

Table 2. Influence of aluminum on concentration and content of macronutrients in tops and roots of rice cultivars

Cultivars Aluminum concentration (ppm)

Tops

0 10 20 40 60

Conc. Content Conc. Content Conc. Content Conc. Content Conc. Content
~o mg/4 pl ~o mg/4 pl ~o mg/4 pl ~o mg/4 pl ~ mg/4 pl

Nitrogen
IAC 1131 4.22 144.88 4.26 147.61 4.46 144.00 4.66 72.15 4.19 58.6
Fernandes 4.92 138.78 5.21 170.29 4.86 135.30 5.00 142.33 4.88 94.5
Matao 3,90 t38.51 4.54 132.08 4.50 116.94 4,41 88,97 4.34 44.97
IPEACO 562 5.38 133.48 4.36 71.04 4.75 53.67 4,66 13.50 3.88 6.35
IRAT 2 4.95 95.87 4.29 49.76 4.89 51.24 4,81 23.92 4.45 10.91
IPEACO 162 5,01 92.95 4.32 49.75 5.09 32.80 3.46 8.13 3.16 14.36
Phosphorus
IAC 1131 0.50 17.24 0.45 15.82 0.41 13,44 0.42 6.65 0.37 5.20
Fernandes 0.84 23.81 0.76 25.08 0.61 17.03 0.58 16.76 0.62 12.12
Matao 0.51 18.33 0.54 16.42 0.54 13.99 0.56 11.48 0.59 6.31
IPEACO 562 0.60 14.96 0.44 7.38 0.41 4.62 0.22 0.66 0.11 0,17
IRAT 2 0.60 11.67 0.39 4.61 0.50 5.33 0.24 1.59 0.13 0,34
IPEACO 162 0.67 12.59 0.45 5.76 0.51 3.45 0.16 0.40 0.14 0.64

Potassium
IAC 1131 4.12 141.42 4.21 145.88 4.02 1.29.66 3.65 56.59 2,86 40.02
Fernandes 4.31 121.60 4,59 150.69 4.24 118.14 4.30 122.25 3,43 66.42
Matao 3.90 138.63 4,04 117.72 3.75 97,15 3,80 75.88 3.51 36.44
IPEACO 562 4.87 120.64 4,08 66.44 4,10 46.33 2,57 7.43 1,86 2.69
IRAT 2 4.99 96.79 4.15 48.09 4.30 45.1 3.07 15.22 2.11 5.17
IPEACO 162 4.43 90,42 4.01 47.34 3.56 26.45 2.29 6.04 2.15 1.68

Calcium
IAC 1131 0.27 9.57 0.21 7.52 0.18 6.00 0.14 2.27 0.12 1.77
Fernandes 0.21 6.18 0.18 6.15 0.16 4.52 0.15 4.39 0.14 2.84
Matao 0.17 6.33 0.17 4.97 0.16 4.16 0.14 2.97 0.13 1.45
IPEACO 562 0.11 3.97 0.18 2.93 0.17 1.93 0.14 0.35 0.14 0.20
IRAT 2 0.22 4.37 0.22 3.15 0.20 2.14 0.16 0.86 0.13 0.33
IPEACO 162 0.22 4.07 0.17 2.09 0,18 1.17 0.15 0.36 0.15 0.66

Magnesium
IAC 1131 0.39 13.40 0.28 9.98 0.27 8.92 0.28 4.33 0.20 2.84
Fernandes 0.33 9.24 0.31 10.25 0.22 6.07 0.21 5.85 0.19 3.65
Matao 0.34 11.90 0.28 8.30 0.28 7.37 0.21 4.27 0.25 2.58
IPEACO 562 0.36 8.94 0.24 3.98 0.22 2.48 0.28 0.80 0.21 0.3t
IRAT 2 0.50 9.61 0.32 3.71 0.32 3.38 0.30 1.50 0.34 0.84
IPEACO 162 0.36 6.60 0.22 2.55 0.28 1.79 0.28 0.65 0.28 1.27
Sodium
IAC 1131 0.27 9.20 0.26 8.92 0.23 7.32 0.32 4.90 0.22 3.08
Fernandes 0.13 3.86 0.13 4.14 0.15 4.06 0.18 5.22 0.20 3.96
Matao 0.24 8.63 0.13 3.86 0.22 5.69 0.23 4.54 0.27 2.79
IPEACO 562 0.11 2.69 0.14 2.27 0.16 1.83 0.42 1.19 0.07 0.10
IRAT 2 0.13 2.54 0.24 2.75 0.15 1.54 0,30 1.60 0.66 1.61
IPEACO 162 0,14 2.60 0.13 1.50 0.21 1.40 0.40 0.94 0.39 1.48

Sulfur
IAC 1131 0.50 17.03 0.48 16.48 0.58 18,71 0.63 9.83 0.48 6.73
Fernandes 0~57 15.99 0.51 16.75 0.50 13.94 0.59 16.78 0.58 11.28
Matao 0.50 17.60 0.51 14.64 0.64 16.62 0,53 10.24 0.52 5.43
1PEACO 562 0.52 12.79 0,60 9,70 0,55 6.17 0.63 1,84 0.66 0,95
IRAT 2 0.58 11.24 0.65 7.55 0.62 6.44 0,64 3.09 0,65 t,60
IPEACO 162 0,58 10,75 0,63 7.45 0.63 4,06 0.68 1.60 0.63 2,73
 A L U M I N U M IN U P L A N D RICE C U L T I V A R S 35

Roots

0 10 20 40 60

Conc. Content Conc, Content Conc. Content Conc. Content Conc. Content
~o mg/4 pl '~/ rag/4 pl ~o rag/4 pl ~,,i rag/4 pl ~ rag/4 pl

2.38 19.99 2.62 21.08 2.41 16.63 4.37 19.19 2.91 8.29
2.79 21.14 3.18 26.59 3.01 24.80 2.97 24.12 2.06 12.27
2,69 18.52 2.71 ~6.38 2.53 13.78 2.20 10.21 2.01 5.42
2.53 11.87 2.63 10.62 2.78 8.06 2.45 4.41 2.01 0.83
2.70 10.26 3.15 11.06 3.35 10.40 3.73 8.25 2,43 4.0
2.99 10.02 3.30 13.84 3.55 7.09 3.65 5.11 2.15 1,19

0.45 3,57 0.58 4.57 0.43 2.93 0.50 2.18 0.49 1.39
0,79 5,98 0.83 6.97 0.77 6.39 0.64 5.19 0.62 3.65
0,40 2.78 0.75 4.51 0.44 2.39 0.55 2.53 0.55 1.48
0.44 2.10 0.46 1.85 0.24 0.70 0.11 0.20 0.11 0.10
0.29 1.08 0.23 0.79 0.26 0.80 0.23 0.50 0.18 0.31
0.45 1.52 0.26 1.09 0.45 0.90 0.14 0.20 0.13 0.07

2.85 22.74 2.33 18.75 2.83 19.49 3,22 14.18 2.37 6.73
3.12 23.60 3.02 25.24 2,73 22.48 3.52 28.60 3.39 20.09
2.32 15.98 2.27 13.72 2.62 l 4.19 2.54 11.78 2,03 5,50
2.63 12.37 2.66 10.80 2.88 8.33 2.02 3.64 1,89 1.86
2.69 10.22 3.01 10.53 3.21 9.91 2.94 6.52 2.50 4.14
2.70 9.04 3.06 12.82 3.14 6,29 2,04 2.86 1.94 1.07

0.10 0.81 0.10 0.81 0.10 0.69 0.10 0.44 0.10 0.30
0.10 0.76 0.10 0.84 0.10 0.82 0.10 0.82 0.09 0.57
0.10 0.69 0.10 0.61 0.11 0.61 0.10 0.47 0.09 0.26
0.10 0.48 0.10 0.41 0.10 0.29 0.12 0.23 0.07 0.08
0.11 0.41 0.10 0.35 0.10 0.32 0.10 0.22 0,08 0.15
0.10 0.34 0.10 0.39 0.13 0.26 0.12 0.27 0.09 0.05

0.24 1.88 0.16 1.42 0.09 0.61 0.09 0.41 0.08 0.24
0.15 1,14 0.14 1.19 0.12 1.00 0.12 0.99 0.11 0.65
0.20 1.37 0.13 0.77 0,08 0.46 0.08 0.39 0,11 0.31
0.17 0.79 0,15 0.6I 0.I5 0,44 0.t2 0.23 0.08 0.09
0.2l 0.79 0.18 0.61 0.I9 0.58 0.15 0.35 0.09 0.15
0.18 0.60 0.17 0.72 0.21 0.43 0.12 0.17 0.08 0.05

0.61 4.85 0.58 6.16 0.48 3.30 0.42 1.85 0.29 0.83
0.21 1.63 0.53 4.43 0.45 3.73 050 4.10 0.48 2.86
0.42 2.92 0.36 2.19 0.48 2.65 0.55 2.57 0.43 1.16
0.22 1.04 0.15 0.62 0.13 0.38 0.16 0.28 0.15 0.14
0.23 0.87 0.13 0,45 0.22 0.69 0.22 0.46 0.26 0.42
0.16 0.53 0,14 0.58 0.21 0.41 0.16 0.22 0.11 0.07
 36 FAGERIA AND CARVALHO

Table 3. Influence of aluminum on concentration and content of aluminum and micro nutrients in tops and roots of
rice cultivars

Cultivars Aluminum concentration (ppm)

Tops

0 10 20 40 60

Conc. Content Conc. Content Conc. Content Conc. Content Conc. Content
ppm pg/4 pl ppm ktg/4 pl ppm ~tg/4 pl ppm I~g/4 pl ppm ~tg/4 pl

Aluminum
IACll31 82.75 283,50 150.50 522.20 297.00 959.75 1240.00 1922.00 1310.00 1837.60
Fernandes 129.00 364.40 126,30 413,70 117.00 3 2 5 . 7 5 236,00 671.00 860.50 1669.30
Matao 59.10 210.10 119.00 345.90 475.50 1210.50 705.00 1405.20 1383.00 1433.00
IPEACO 562 93.70 232.40 145.00 236.55 176.00 199.20 718.00 208.30 3071.50 445.35
IRAT 2 89.50 173.85 166.50 193.15 239.00 2 5 0 . 7 0 794.50 397.15 2247.50 550.65
IPEACO 162 85.10 157.80 144.50 168,75 281.50 181.85 876.00 205.90 3885.00 1732.50

Zinc
IAC 1131 31.70 109.00 26.05 90.50 23.50 76.00 20.90 32.50 16.85 23.50
Fernandes 35.50 9 8 . 9 0 24.90 81.65 25.35 70.75 23.25 66.30 15.30 29.60
Matao 17.85 6 3 . 5 0 18.25 54.00 15.20 39.50 12.55 25.00 11.70 12.00
IPEACO 562 29,70 73.75 26.50 43.25 22.85 25.75 17.20 5.15 14.80 2.10
IRAT 2 24.50 4 7 . 6 5 31.10 36.10 27.85 29.40 17.25 8.65 12.80 3.15
IPEACO 162 49.20 91.45 27.75 32.55 23.85 15.40 16.45 3.90 15.05 6.06

Iron
IAC 1131 300,00 1030.50 335.00 1162.10 277.50 896.70 189.50 293.20 251.50 352.15
Fernandes 3 0 4 , 5 0 857.85 314,50 1033,20 190.50 530.90 229,50 653.95 137.50 268.15
Matao 233.00 828.35 251.00 730.80 276.50 718.65 161.00 320.70 214.00 221,70
IPEACO 562 260.50 645.35 353.00 575.40 208.50 235.60 135.50 39.40 182.50 26,40
IRAT 2 307.00 595.85 243.00 281.80 238.00 249.85 154.50 77.60 153.00 37.50
IPEACO 162 390.50 724,75 288,50 336,75 232,00 149.10 166.00 39.85 1 2 2 . 5 0 62.30

Manganese
IAC ll31 499.50 1715.95 323.00 1120.60 269.00 8 6 9 . 0 0 205.00 317.50 79.75 111.60
Fernandes 7 7 0 . 5 0 2181.95 365.00 1023.80 366.50 1020.25 329.50 937.40 151.50 293.70
Matao 259.50 922.50 276.00 801.95 168.00 4 3 8 . 5 5 129.50 258.25 83.35 87.40
IPEACO 562 628.50 1559.10 591.00 962.30 703.00 7 9 4 . 2 5 327,00 95.00 2 1 2 . 5 0 30.80
IRAT 2 661.50 1284.15 669.50 776.35 561.00 5 6 6 . 0 0 299,50 149.65 1 3 3 . 5 0 32.73
IPEACO 162 1350.00 2505.80 805.50 936.45 628.50 4 0 5 , 2 5 369,00 86.70 2 1 3 . 0 0 94.10

Boron
IAC 1131 24.15 82.90 27.25 94.56 29.55 95.55 24.95 38.81 40,50 56,53
Fernandes 18.85 53.12 24.45 83.53 24,80 69,05 23.85 67,84 18.90 36,68
Matao 13.95 49.59 16.05 46,93 23.95 62.29 20.70 41.32 26.80 27.68
IPEACO 562 17.45 43.27 21.95 36.48 15.85 17.90 66.50 19.06 65.50 9.48
IRAT 2 26.50 51.45 30.90 34.36 28.70 30.16 40.85 20.71 53.55 13.14
IPEACO 162 24.95 46.28 23.80 27.70 14.90 9.62 52.10 12.23 68.80 30.68

Copper
IAC 1131 16.15 55.42 19.65 68.14 19.85 64.18 24.65 38.32 16.95 23.69
Fernandes 27.60 64.14 24.60 80.81 26.15 72.90 26.85 76.45 25.65 49.58
Matao 14.80 52.61 25.80 75.24 26.05 67.49 16.75 33.51 19.75 20.61
IPEACO 562 19.95 49.60 27.05 44.16 21,85 24.65 15.00 4.32 18.05 2.61
IRAT 2 16.90 32.77 21.75 25.25 24.60 25.86 15.55 7.91 10.10 2.48
IPEACO 162 31.80 59.12 27.95 32.86 13.90 9.06 10.04 2.36 21.55 9.51
 A L U M I N U M IN UPLAND RICE CULTIVARS 37

Roots

0 10 20 40 60

Conc. Content Conc. Content Conc. Content Conc. Content Conc. Content
ppm ~tg/4 pl ppm lag/4 pl ppm ~g/4 pl ppm lag/4 pl ppm p.g/4 pl

418.00 334.50 5585.00 4495.70 4052.50 2796.30 2905.00 1 2 8 0 . 9 0 3685.00 1050.50

439.50 331.80 1758.00 1468.00 3268.00 2679.90 3050.00 2472.50 3160.00 1880.40
126.00 86.90 1 4 0 5 . 0 0 849.80 1 5 0 0 . 0 0 817.50 2279.50 1059.90 2245.00 606.50
282.00 132.60 1 0 0 5 . 5 0 407.50 1 5 4 5 . 0 0 447.80 2965.00 534.10 3739.50 372.90
237.50 90.30 1149.00 402.10 1585.00 490.90 28.95 639.70 3450,00 570.00
249.00 83.40 850.00 356.00 1 6 8 0 . 0 0 335.40 2714.00 380.20 3685.00 203.00

29.40 23.54 37.40 30.09 28.80 19.76 40.55 17.88 31,50 8.98
30.00 22.80 24.85 20.75 23.60 19.51 28.30 22.98 30,50 18.14
12.75 8.79 13.80 8.34 12.65 6.95 14.85 6.91 13,65 3.73
26.50 12.46 24.50 9.86 27.85 8.06 26.55 4.78 23.50 2.34
27.15 10.30 25.85 9.06 27.90 8.62 26.00 5.88 21.00 3.45
19.90 6.67 21.00 8.80 35.65 7.15 31.65 4.44 29.00 1.59

3065.00 2450.40 2680.00 2168.30 1270.00 876.40 906.50 400.50 899.50 256.70
2585.00 1956.90 1422.001189.50 1165.00 965.20 874.00 703.30 632.50 375.90
1535.00 1 0 4 4 . 3 0 1790.00 1091.80 1172.00 684.20 1090.00 507.10 1085.00 295.40
1445.00 679.20 1 3 8 0 . 0 0 561.20 1295.00 374.70 637.50 115.80 514.00 52.40
1322.50 513.10 1 2 1 0 . 0 0 417.90 972.00 302.20 839.00 183.90 715.00 118.80
1980.00 647.90 1 8 4 0 . 0 0 774.60 1630.00 332.90 589.00 82.40 430.00 24.00

116.50 93.20 80.25 64.40 35.65 24.60 39.85 17.60 22.30 6.40
77.05 58.60 71.00 59.30 65.20 54.10 54.60 45.80 43.05 26.60
36.40 25.20 41.05 24.90 28.45 15.50 24,80 11.60 18.70 5.10
97.50 45.80 94.00 38.10 80.05 23.02 51.05 9.20 45.00 4.50
129.15 49.30 135.50 47.70 73.50 22.74 57.00 12.60 46.50 7.70
232.50 78.00 181.50 76.30 155.50 31.27 113.50 15.90 100.50 5.50

64.50 51.57 59.00 47.41 59.95 41.37 73.55 32.52 65.00 18.85
51.30 39,07 44.65 37.29 49.05 40.54 55.45 45.02 37.95 22.55
71.25 49,20 54.55 32.71 65.75 35.58 61.70 28.68 60.10 16.24
90.25 42.42 83.50 34.11 83.45 24.13 100.50 18.10 96.50 9.60
62.50 23.74 74.50 26.11 93.95 29.01 100.00 22.00 79.40 13.08
84.65 28.38 85.80 36.09 92.80 18.56 145.00 20.35 85.00 4.66

24.55 18.20 20.30 16.34 17.65 12.18 35.65 15.73 38.80 11.07
64.30 48.88 39.50 32.98 39.15 32.38 49.60 40.16 53.65 31.73
15.05 10.39 36.20 22.05 19.00 10.39 26.50 12.32 25.00 6.75
31.00 14.57 37.50 15.12 49.55 14.34 31.90 5.74 33.50 3.37
16.50 6.26 20.95 7.34 33.10 10.23 40.00 8.80 41.50 6.84
35.75 12.00 40.50 17.00 51.75 12.21 33.85 4.75 35.50 1.96
38 FAGERIA AND CARVALHO

Based on the results shown in Table 1, the cultivars were divided into 3 groups.
Cultivars which gave positive linear regression equations were put in one group,
those which gave non-significant regression equations were put in the second
group and those which gave quadratic regression equations were put into the
third group. Two cultivars from each group were selected and values of
concentration and content of aluminum, macro and micro nutrients in the tops
and roots of these cultivars were presented in Tables 2 and 3.
Aluminum concentration and content in tops as well as in the roots increased
as a result of increasing A1 concentrations in the nutrient solution. Aluminum
concentrations in the roots were much higher than the concentrations in the
corresponding plant tops at all treatments levels. Similarly A1 content was higher
in the roots as compared with tops except at the highest level of A1. At 60 p p m A1,
the situation was changed and 67~o, of the cultivars had more AI in the tops as
compared with the roots. The relatively low concentration of A1 in the plant tops
compared with those in the roots indicate that A1 accumulates to a greater extent
in the roots of plants grown in A1 solutions 1~ 24
Before harvest, plant roots were washed thoroughly in distilled water to
remove any elements that might have been adsorbed, on to the root surfaces.

~ ' , o c - %Fernondes

L
E 2r ~
:I CRITICALTOXIC LEVEL 1250pprn) : /CRITICALTOXICLEVEL

i/ o,o , 500 IO00

~ ,oc-N MofSo lpeoco 5 6 2
--oL
t~ 6c

0
~ 4C-- RITICAL TOXICLEVEL (t33 ppm)

0 CRITICALTOXICLEVEL 1160ppm)

!/ ~'o ,2~0 -- 2000 3000

~-~0,oo~ Zrof 2 J'peooo f62

~RITICAL TOXIC LEVEL (|00 ppm)

[ [ / I
I000 1000 200o 4000
ALUMINUM CONCENTRATION Ippm)
Fig. 1. The relationship between dry matter production and concentration of A1in the tops of 21
days old rice plants.
ALUMINUM IN UPLAND RICE CULTIVARS 39

Most of the Al found in the root material would therefore be accumulated

within the root and not merely present on the root surfaces. Aluminum has been
shown to accumulate in the roots of corn 2 ~, potato 19, and barley 27.
Critical toxic level of A1 in the tops of plants of 6 cultivars was determined on
the basis of the relationship between concentration of A1 in plant tops and
production of dry matter (Fig. 1). This was indicated at 90~o of maximum yield by
the interrepted straight lines (Fig. 1). Values of critical toxoc levels varied from
cultivar as follows: IAC 1131, 417 ppm; Fernandes, 250 ppm; Mat~.o, 160 ppm;
IPEACO 562, 133 ppm; IRAT 2, and IPEACO 162, 100 ppm.

Effect of Al on nutrient uptake

The effect of Al treatment on N, P, K, Ca, Mg, S, Na, Zn, Fe, Mn, B, and Cu in
plant roots and tops of 6 cultivars are presented in Tables 2 to 3.

Nitrogen and phosphorus The effects of A1 treatments on N and P concentration

and content in plant tops and roots are presented in Table 2.
Nitrogen concentration was increased or remained constant in the tops of
most of the cultivars up-to 40 ppm of A1 but thereafter it was decreased. In
relation to content, in 33~o of cultivars it was stimulated up to 10 ppm but there
after was a decrease with increasing concentration of AI. In the roots,
concentration and content were not changed significantly up to 40 ppm A1
treatments and then decreased. Concentration and content were much higher in
the tops as compared with roots in all of the cultivars.
Phosphorus, concentration and content were decreased at higher levels of Al in
the tops as well as in the roots. Phosphorus content in the plant tops was
drastically reduced at 40 and 60 ppm A1. In all the cultivars P content was higher
in the tops as compared with roots but concentration was more or less equal in
the roots at 60 ppm A1 in most of the cultivars.
Nutrient inhibition was calculated taking into consideration the control and
the highest concentration of aluminum (Table 4). Inhibition of 76 to 100~o was
considered high, 51 to 75~o was considered medium and less than 50~o was
considered low. Inhibition of P was more than N. For N, 33~o of the cultivars
were fall in the inhibition range of 76 to 100~o, 63~o of the cultivars were inhibited
51 to 75~o and the rest were in the group of less than 50~o inhibition. For P 57~o of
the cultivars were in the inhibition range of 76 to 100~o, 40~o were inhibited 51 to
75~o and 3~o were in the group of less than 50~o inhibition.
Reduction in phosphorus absorption by plant roots and tops by A1 was
explained by Andrew et al. 2, Rorison 26, and Clarkson 3. Inhibition data
presented by them suggested that most of the Ai assimilated by roots became
strongly bound to absorption sites in the cell wall. Positively charged surfaces of
amorphos hydrated aluminum oxides are known to absorb and precipitate P for
solutions 16. Cell surfaces on which hydrated aluminum oxide is present would be
expected to absorb P and, to this extent, they may be regarded as further site for
phosphate fixation 3.
 40 FAGERIA AND CARVALHO

Table 4. Inhibition of nutrients content by A1 treatments in the rice cultivars tops

Cultivars Plant tops

N P K Ca Mg Na S Zn Fe Mn B Cu

IAC 1246 62 75 68 83 76 60 49 77 77 92 40 61
IAC 47 62 72 70 85 81 60 45 80 82 93 35 53
IAC 5544 61 75 70 81 83 60 45 85 84 93 48 56
IAC 5100 66 78 75 88 84 70 65 89 87 96 69 74
IAC 1131 60 70 72 82 79 67 61 78 66 94 32 57
IAC 25 72 77 79 87 82 70 66 84 79 95 70 74
MatZo 68 66 74 77 78 68 69 81 73 91 44 61
Pratao 63 63 70 79 79 55 63 86 83 94 52 67
Dourado precoce 74 75 77 76 82 71 70 86 83 82 71 85
Prat~o precoce 64 67 75 86 83 59 64 92 86 97 51 82
M o n t a n h a liso 67 68 79 82 83 63 67 91 88 97 78 80
Batatais 69 65 77 81 80 69 64 90 85 95 74 79
Quatro meses 71 88 73 81 82 90 52 92 88 95 76 85
P6rola 63 85 65 74 74 16 43 91 85 93 56 84
Ponta Preta 76 92 78 84 87 50 67 93 91 96 77 84
I P E A C O 163 69 87 74 77 76 01 50 93 80 96 64 82
Catal5o 83 94 70 88 85 - 72 93 88 97 54 90
IPEACO-462 60 85 63 74 76 - 41 88 83 94 48 63
IRAT 8 92 98 97 95 93 22 88 95 96 98 70 95
IRAT 12 91 98 96 95 94 15 85 94 93 98 85 93
IRAT 2 89 97 95 92 91 36 86 93 94 98 75 92
IPEACO-162 85 95 98 84 81 43 75 93 91 96 34 84
IPEACO-562 95 98 98 95 97 96 93 97 96 98 78 95
Paulistinha 94 98 95 94 95 66 88 96 96 98 81 93
Fernandes 32 49 45 54 61 - 30 70 69 87 31 23
IRAT 11 64 73 59 73 77 - 55 86 82 95 55 65
IRAT 13 68 74 70 73 76 23 60 80 70 92 54 65
Esa6 68 88 74 81 79 50 62 89 87 94 47 77
Bico ganga 78 87 75 80 80 15 66 82 87 94 56 74
M R C 172-9 91 96 93 94 96 28 84 94 92 97 72 92

content at 0 p p m AI - content at 60 p p m
Inhibition x 100
content at 0 p p m AI

This inhibitory effect of AI, on P has practical implications in acidic soils in

tropical climates. For example, in the 'cerrado' region of Brazil, which represents
a b o u t 180 million hectares (approximately 22~ of the National territory),
phosphorus deficiency is the one of the important limiting factors in upland rice
productiona. One reason for this deficiency is the fixation of phosphorus by
aluminum.
ALUMINUM IN UPLAND RICE CULTIVARS 41

Potassium, calcium, magnesium, sulfur and sodium Results related to

concentration and content of K, Ca, Mg, S and Na in the roots and tops of 6
cultivars of rice are presented in Table 2. In 5 0 ~ of the cultivars, K concentration
was increased up to 10 p p m of A1 in the tops and then decreased. Most drastic
decreases occurred at 60 p p m of A1. With respect to K content, there was a
tendency to decrease with increasing levels of A1 and the most drastic change
occurred at 40 and 60 ppm of AI. In the roots similar trends were observed but
decrease in concentration and content at higher A1 concentrations was not as
sharp as in the case of the tops. Concentration as well as potassium content was
lower in the roots as compared with tops. With respect to inhibition, 40~o of the
cultivars were in the group of 76 to 100~, 57~o in the group of 51 to 75~ and rest
were in the group of less than 50~ inhibition.
Ca concentration and content in the tops decreased with increasing
concentration of A1 but in the roots concentration was more or less constant at all
AI levels. Content was decreased at 40 and 60 p p m A1. The plant tops had more
Ca as compared with the roots. Calcium inhibition was higher than P, K, N, S
and Na. Eighty-three percent of the cultivars were in the range of 76 to I 0 0 ~
inhibition, and 17~ were in the range of 51 to 75~.
A similar trend was observed with respect to Mg concentration and content in
the tops and roots of 6 cultivars of rice. Aluminum completely inhibited uptake of
Mg. Among the cultivars, 93~ were in the group of 76 to 100~ and the rest were
in the range of 51 to 75~.
Sulfur uptake was only determined in the plant tops. Concentration of this
element was either increased or constant at lower A1 concentrations.
Sulfur content was decreased only at higher levels of A1 (40 and 60 ppm). Sulfur
inhibition by A1 in 2 0 ~ of the cultivars in the tops occurred in the range of 76 to
100~, 57yo in the group of 51 to 75% and 23~o of the cultivars had less than 50~
sulfur inhibition.
Sodium content in the tops was affected at a higher concentration of A1.
Decrease in the content was higher at 40 and 60 p p m of A1. In general,
concentration with A1 treatments was neither affected in tops nor in the roots.
Sodium concentration in the roots was higher as compared with the tops but
content was higher in the tops.
Among macronutrients N a was least affected by A1 treatment. At 60 ppm AI
only 3~o of the cultivars were in the group of 76 to 100~o inhibition, 47~o in the
group of 51 to 75~o and rest were less than 50~ inhibited (Table 4).

Zinc, iron, manganese, boron and copper Zinc concentration and content was
decreased with increasing concentrations of A1 in the tops. Similary, content, in
the roots was also decreased with AI treatments, but concentration was decreased
only at 60 ppm A1 concentration.
Zinc content was higher in the tops as compared with roots, but concentration
in the roots was higher especially at higher concentrations of aluminum.
42 FAGERIA AND CARVALHO

Concentration and content of iron in tops as well as in the roots was decreased
with increasing concentrations of A1. Iron concentration and content was higher
in the roots as compared with the tops. These results indicates that even though
Fe enters plant roots that are growing in A1 solutions, Fe accumulates in the roots
with no increased translocation into plant tops.
Concentration and content of Mn was decreased with increasing A1
concentration in the tops as well as in the roots. These values were higher in the
tops as compared with roots.
Boron concentration and content in the tops was increased up to 10 ppm of A1.
Above 10 ppm content decreased with increasing A1 concentration in most of the
cultivars but concentration in most of the cultivars either increasing or remained
constant. A similar trend was observed in the roots. Concentration of boron was
higher in the roots as compared with tops but tops had a higher content as
compared with the roots.
Copper concentration up to 20 ppm of A1 was not affected much but decreased
at 40 and 60 ppm of A1 in the tops and the roots in most of the cultivars. Content
was higher in the tops while concentration was higher in the roots. These results
suggest that Cu may compete effectively with A1 for root absorption sites.
Among micronutrients Mn uptake was affected most while boron was affected
least (Table 4). Among the 30 cultivars tested. 100~o fell into the group of 76 to
100~o inhibition of Mn. The second most inhibited micronutrient was Zn.
Ninety-seven percent of the cultivars fell into the group of 76 to 100~ inhibition
and the remaining 3~o were in the group of 51 to 76~o inhibition.
As far as Fe is concerned, 87~o of the cultivars were in the group of 76 to 100%
inhibition and rest fell in to the range of 51 to 71~o inhibition.
In regards to copper, 57~ of the cultivars were 76 to 100~o inhibition, 4 0 ~ were
in the range of 51 to 75~o, and the remaining 3~o were in the lowest group, Boron
uptake was, least affected among the, micronutrients tested with A1 treatment.
With this element no cultivars fell in to the inhibition range of 76 to 100~o, 70~o of
the cultivars were in the range of 51 to 75~o inhibition and the remaining 30~
were in the last range < 50~o inhibition.

Reasons for decreasing nutrients uptake by aluminum

As discussed above the results demonstrated higher concentrations, of A1 (40
and 60 ppm) decreased uptake of most nutrients. Decreases in uptake of these
elements was mostly related to morphological, physiological and biochemical
effects of aluminum.
The following effects occurred:
1. Aluminum inhibited, root growth 6'~, thereby causing the uptake of these
nutrients to be reduced 22.
2. Aluminum reduced cellular respiration in plants causing an inhibition in
the uptake of all ions 1,8.
A L U M I N U M IN U P L A N D RICE CULTIVARS 43

3. Many investigators have shown that A1 increases the viscosity of

protoplasm in plant root cells and decrease overall permeability to salts 1,22.
4. Aluminum blocks, neutralizes, or reverses the negative charge on the pores
of the free space and thereby reduced the abilities of such pores to bind C a 4.
5. Aluminum may compete for common binding sites at or near the root
surface and thereby reduce uptake of K, Ca, Mg and Cu ~4' 15
6. A hypothesis advanced to explain the A1 reduction of Ca uptake is that a
portion of the Ca accumulation mechanism was completely inactivated by the A1
concentration 17.
7. In general, A1 has been shown to interfere with cell division in plants roots,
decrease root respiration, interfere with certain enzymes governing the
deposition of polysaccharides in cell walls, increase cell wall rigidity (by cross-
linking pectins) and interfere with the uptake, transport and use of several
elements, such as K, Ca, and Mg a2.
8. Aluminum injured plant roots and reduced Ca uptake 18.
9. Aluminum decreased the sugar content, increased the ratio of non-protein
to protein N, and decreased P contents of leaves from several plants grown on
acidic soils lz.

Acknowledgements The authors are grateful to Dr. Almiro Blumenschein, Director of the National
Rice and Bean Research Center for providing necessary facilites for this investigation. The authors
also acknowledge with thanks the assistance of Mr. Vand61io Afonso de Oliveira, Miss Nilva Abadia
Rabelo and Miss Osmira Ffitima da Silva, all Laboratory Technicians, for tabulating the data of this
work. Thanks are due to Dr. W. R. Kussow, Professor of Soil Science, University of Wisconsin,
Madison and Dr. C. A. Jones, Plant Physiologist, A. R. S. Temple, Texas for their help in getting the
Plant Samples analysed.

Received 8 June 1982. Revised August 1982

References

1 Aimi R and Murakami T 1964 Cell Physiologycal studies on the effect of aluminum on the
growth of crop plants. Nat. Inst. Agric. Sci. Bull. Ser. D. 11, 331-396.
2 Andrew C S, Johnson A D and Sandland R L 1973 Effect of aluminum on the growth and
chemical composition of some tropical and temperate pasture legumes. Aust. J. Agric. Res. 24,
325-339.
3 Clarkson D T 1967 Interactions between aluminum and phosphorus on root surfaces and cell
wall materials. Plant and Soil 27, 347 356.
4 Clarkson D T 1971 Inhibition of the uptake and long distance transport of calcium by
aluminum and other polyvalent cations. J. Exp. Bot. 23, 837 851.
5 Fageria N K 1980 Defici~ncia hidrica em arroz de cerrado e resposta ao f6sforo. Pesq. agropec.
bras. Brasilia 15, 259-265.
6 Fageria N K 1982 Toler~.ncia diferencial de cultivares de arroz ao Aluminio em solu~;,~o
nutritiva. Pesq. agropec, bras. Brasilia 1, 1-9.
7 Fageria N K and Zimmennann F J P 1979 Selegfio de cultivares de arroz para tolerfincia a
toxidez de aluminio em solu~;fio nutritiva, Pesq. agropec, bras., Brasilia 14, 141-157.
8 Fageria N K, Barbosa Filho M P and Carvalho J R P 1982 Response of upland rice to
phosphorus fertilization on an Oxisol of a Central Brazil. Agron. J. 74, 51-56.
44 ALUMINUM IN UPLAND RICE CULTIVARS

9 Farhoomand M B and Peterson L A 1968 Concentration and content. Agron. J. 60, 708-709.
10 Foy C D and Brown J C 1963 Toxic factors in acid soils I. Characterization of aluminum
toxicity in cotton. Soil Sci. Soc. Am. Proc. 27, 403-407.
11 Foy C D and Brown J C 1964 Toxic factors in acid soils II. Differential aluminum tolerance of
plant species. Soil Sci. Soc. Am. Proc. 28, 27-32.
12 Foy C D 1974 Effects of aluminum on plant growth. In The Plant Root and its Environment,
Ed. E W Carson. Charlottesville. The University Press of Virginia, USA pp 601-642.
13 Gregg D C 1965 Principles of Chemistry. Allyn and Bacon. Inc., Boston.
14 HarwardME, JacksonWA, L o t t W L a n d M a s o n D D 1955 Effects of AI, Fe, a n d M n u p o n
the growth and composition of lettuce. Proc. Am. Soc. Hortic. Sci. 66, 281-286.
15 Hiatt A J, Amos D F and Massey H F 1963 Effect of aluminum on copper sorption by wheat.
Agron. J. 55, 284-287.
16 Hsu P H and Rennie D A 1962 Reactions of phosphorus in aluminum systems I. Absorption of
phosphorus by X-Ray amorphos aluminum hydroxide. Can. J. Soil Sci. 42, 197.
17 Johnson R E and Jackson W A 1964 Calcium uptake and transport by wheat seedlings as
affected by aluminum. Soil Sci. Soc. Am. Proc. 28, 381-386.
18 Lance J C and Pearson R W 1969 Effect of low concentrations of aluminum on growth and
water and nutrient uptake by cotton roots. Soil Sci. Soc. Am. Proc. 33, 95-98.
19 Lee C R 1971 Influence of aluminum on plant growth and mineral nutrition of potatoes.
Agron. J. 63, 604-608.
20 Liegel E A, Simson C R and Schutte E E 1980 Wisconsin procedure for soil testing, plant
analysis and feed forage analysis. Madison, Univ. of Wisconsin., Dept. of soil Sci. Soil fertility
series, 6, WI,. Revised.
21 Lopes A S and Cox F R 1977 A survery of the fertility status of surface soils under 'cerrado'
vegetation in Brazil. Soil Sci. Soc. Am. J. 41,742-747.
22 Mclean F T and Gilbert B E 1927 The relative aluminum tolerance of crop plants. Soil Sci. 24,
163-175.
23 Olmos I L J and Camargo M N 1976 Ocorr6ncia de aluminio t6xicos nos solos do Brasil, sua
caracterizagSo e distribui~5.o Ci e Cult. S5o Paulo 28, 171-80.
24 Randall P J and Vose P B 1963 Effect of aluminum on uptake and translocation of
phosphorus 32 by perennial ryegrass. Plant Physiol. 38, 403-409.
25 Rasmussen H P 1968 Entry and distribution of aluminum in Zea mays: Electron microbe X-
Ray analysis. Planta Berlin 81, 28-37.
26 Rorison I H 1965 The effect of aluminum on the uptake and incorporation of phosphate by
excised sainfoin roots. New Phytol. 64, 23-27.
27 Wright K E and Donahue B A 1953 Aluminum toxicity studies with radioactive phosphorus.
Plant Physiol. 28, 674-680.
28 Yoshida S, Forno D A, Cock J H and Gomez K A 1976 Laboratory manual for physiological
studies of rice. Los Bafios, Philippines. The International Rice Research Institute, pp 54--57.