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African Crop Science Journal
African Crop Science Society
ISSN: 1021-9730 EISSN: 2072-6589
Vol. 9, Num. 2, 2001, pp. 393-400
African Crop Science Journal, Vol. 9. No. 2, pp. 393-400

African Crop Science Journal, Vol. 9. No. 2, pp. 393-400

Calcium tolerance and ion uptake of Egyptian lupin landraces on calcareous soils

S. Raza, A. Abdel-Wahab1, B. Jørnsgård and J. L. Christiansen
The Royal Veterinary and Agricultural University, Department of Agricultural Sciences, Agrovej 10, DK- 2630 Taastrup, Denmark
1Soil, Water and Environment Research Institute, Agricultural Research Centre, Egypt

(Received 23 February, 2000; accepted 20 September, 2000)

Code Number: CS01021

INTRODUCTION

White lupin (Lupinus albus L.) is a calcifuge susceptible to iron chlorosis making it grow poorly on calcareous soils (Tang et al., 1995b). In general cultivation is only possible on acid to neutral soils with moderate calcium content. This has been a major constraint to development of the lupin crop in Europe (Duthion, 1992; Siddons et al.,1994), and Australia (Tang et al.,1995a). The growth reduction may be caused by several factors such as high pH, high bicarbonate content, Fe deficiency or Ca toxicity. Thus, both root and shoot growth (Tang and Thomson, 1996) were reduced in L. albus at pH 7.5 compared to 6.0. Also the nodulation was negatively affected by increasing pH above 6.0 (Tang and Robson, 1993). The poor growth may also be caused by iron deficiency related to low uptake on alkaline soil (White and Robson, 1989). The severity of iron chlorosis correlates, however, poorly with shoot growth and seed yield in the field (Tang et al., 1995b), especially for plants showing iron chlorosis at early growth stages (White and Robson, 1989; Tang et al., 1995b).

The cause of poor growth in alkaline soil may also be Ca toxicity. Thus, Duthion (1992) showed that plant growth was related to the CaCO3 level but not to extractable Fe or soil pH. Similarly, Jessop et al. (1990) found that the weight of shoot and roots decreased when the concentration of calcium in the soil increased. De Silva et al. (1994) has explained the adverse effect of high soil Ca found in L. luteus as a result of uncontrolled Ca uptake resulting in poor growth and reduced CO2 uptake, but it is not known if a similar mechanism exists in L. albus. Because of its susceptibility to calcareous soils white lupin is generally not grown on soils with a pH above 7.5. However, white lupin is an ancient crop in Egyptian agriculture, where soils generally have a high calcium content with pH ranging from 7.4 to 9.0 (Raza et al., 1999). It could therefore be anticipated that germplasm of lupin adapted by Egyptian farmers would present a source of calcium tolerant genotypes. A collection of Egyptian landraces was undertaken in 1996 (Christiansen et al., 1999). The main objective of the present experiment was to screen this collection to identify potential tolerant accessions and access the effect of a high content of calcium on growth and ion uptake in white lupin.

MATERIALS AND METHODS

Field experiment. A collection of 92 landraces of white lupin from Egypt (Raza et al., 1999), six lines from Australia and the Egyptian cultivars Giza 1 and Giza 2 (Table 1), developed by The Agricultural Research Centre, Egypt about 30 years ago (Anon., 1994) were tested in a triple lattice design in Nubaria (West Delta). This location has a calcareous soil (20-26 % CaCO3). Eight randomly selected soil samples from the experimental plot were analysed for chemical and physical characteristics (Table 2). The experiment was sown on 22 November 1997. Each plot was composed of two rows of 3 m with inter-row spacing of 0.6 m (i.e., plot size 3.6 m2). Seeds were sown in hills 15 cm apart with 2 seeds per hill. Basal doses of N, P and K were added at a rate of 30 kg of N (as urea 46%N), 60 kg of P (15.5% P2O5) and 40 kg of K (48% K2O). The experiment was irrigated twice. The first irrigation took place immediately after sowing and the second just before flowering. Harvest took place at full maturity. Ion uptake of N, Fe, Mn and Ca were determined 60 days after sowing in five tolerant and five susceptible accessions among the 43 surviving accessions. N content was measured by the micro-Kjeldahl method using acid digest of sulphuric-perchloric solution, (Eastin, 1978; Page, 1982). Fe, Mn and Ca were measured by atomic absorption spectrophotometer using ascorbic acid extraction (Watanabe and Olsen, 1965). Analysis were based on a sample of five plants from each replication. At harvest 5 plants in each plot were taken at random to record plant height, number of branches per plant, number of pods per plant, number of seeds per plant, and seed weight per plant. Seed yield per plot was also recorded.

Table 1. Source of different lupin cultivars
Number Accession number* Origin or source Number Accession number Origin or source
1 1 Belbies-Sharkia 51 G7 Giza 7
2 2 Belbies-Sharkia 52 G8 Giza 8
3 3 Belbies-Sharkia 53 G9 Giza 9
4 4 Belbies-Sharkia 54 G10 Giza 10
5 5 Belbies-Sharkia 55 G11 Giza 11
6 6 Belbies-Sharkia 56 G12 Giza 12
7 7 Belbies-Sharkia 57 G13 Giza 13
8 8 Belbies-Sharkia 58 G14 Giza 14
9 9 Abo hammad- 59 L7 Local 7
10 10 Abo hammad- 60 L12 Local 12
11 11 Abo hammad- 61 L20 Local 20
12 12 Fakous-Sharkia 62 L23 Local 23
13 13 Fakous-Sharkia 63 A6 Ac.6
14 14 Fakous-Sharkia 64 A15 Ac.15
15 15 Fakous-Sharkia 65 A21 Ac.21
16 16 Fakous-Sharkia 66 H2 H1-90-72
17 18 Ismailia 67 S1 Saflta Sohag
18 19 Ismailia 68 K1 Kana
19 20 Fayed-Ismailia 69 A1 Adfor Aswan
20 21 Fayed-Ismailia 70 E1 Esna Kana
21 22 Fayed-Ismailia 71 E2 Esna Kana
22 23 Kantra-Ismailia 72 E3 Esna Kana
23 24 Kantra-Ismailia 73 E4 Esna Kana
24 25 Ismailia 74 K2 Kaos Kana
25 26 Abo Soeir-Isma. 78 K3 Kaos Kana
26 27 Abo Soeir-Isma. 79 B1 Belbies
27 28 Algarb-Ismailia 80 K4 Kana
28 29 Algarb-Ismailia 81 K5 Kana
29 30 Meet Gamer-Dak. 82 K6 Kana
30 31 El-Badrashein-Giza 83 K7 Kana
31 32 El-Aiat-Giza 84 N1 N-Hamadi
32 34 Beni Salh-Fayoum 85 N2 N-Hamadi
33 35 Beni Suef 86 K8 Abotashet Ka
34 36 Beni Suef 87 K9 Abotashet Ka
35 39 Beni Suef 88 G1 Grga Sohag
36 40 El-Minia 89 G2 Grga Sohag
37 41 Aswan 90 M1 M-Sohag
38 42 Aswan 91 M2 M-Sohag
39 43 Sohag 92 M3 M-Sohag
40 44 Assiut 93 S2 Sohag 1
41 Kiev Mutant Kiev Mutant 94 S3 Sohag 2
42 Buttercup Buttercup 95 T1 Tama Sohag
43 Piscevoj Piscevoj 96 T2 Assuit
44 75 B15.17-Aus. 75 B15.17-Aus. 97 G15 Elaiat-Giza
45 75 B9.10-Aus. 75 B9.10-Aus. 98 K8 Kalubia
46 P 20950-Aus. P 20950-Aus. 99 Giza 1 Giza 1
47 ARC 1 Giza 100 Giza 2 Giza 2
48 ARC 2 Giza 4      
49 ARC 3 Giza 5      
50 ARC 4 Giza 6      
* collection number in the Egyptian gene bank

Table 2. The main physical and chemical properties of the Egyptian calcareous soil samples
Determinations 1* 2 3 4 5 6 7 8
CaCO3 26.6 27.0 23.3 25.9 24.6 24.9 24.9 20.0  
PH 7.9 7.8 8.3 7.8 8.3 8.1 7.8 8.2
EC 5.14 3.92 3.16 3.72 2.56 2.81 4.38 3.16
Ca2+ mg/L 19.06 19.06 12.38 12.36 3.09 2.09 6.18 5.97
Mg2+ mg/L 15.15 14.88 11.62 11.71 12.32 14.88 13.61 9.44
Na+ mg/L 16.85 15.40 13.72 10.56 10.12 10.12 14.89 11.80
K+ mg/L 2.89 2.75 2.227 1.92 1.50 1.22 2.09 2.00
HCO3- mg/L 4.8 4.8 3.4 4.5 4.8 4.2 4.0 4.5
Cl- mg/L 20.0 19.9 14.8 13.6 10.0 11.0 12.0 9.6
SO42- mg/L 29.12 37.39 18.89 18.45 12.23 13.08 20.77 15.08
N (ppm) 140 125 100 80 150 130 90 140
P (ppm) 3.9 5.2 5.2 3.9 2.1 20.8 2.1 5.2
K (ppm) 18 18 19 17 14 54 36 16
O.M % 3.5 2.5 2.4 3.4 1.4 2.3 2.4 2.5
Sand % 80 75 78 72 77 75 82 81
Silt % 15 20 16 18 15 15 10 11
Clay % 5 5 6 10 8 10 8 8
Texture Loamy sand Loamy sand Loamy sand Loamy sand Loamy sand Loamy sand Loamy sand Loamy sand
1* Soil samples

Glasshouse experiment. Three tolerant accessions from the previous experiment (Giza 1, accession 1 and 24) and three French varieties (CH30470, La625 and Lublanch) were grown in pots at nine different levels of calcium (0, 3, 5, 7, 10, 13, 15, 17 and 20% CaCO3). The calcium level was adjusted by adding CaCO3 to a sandy loamy soil. Seeds were inoculated with a suspension of Bradyrhizobium (Lupin). Irrigation from below was done to maintain 100% water holding capacity. Chlorosis was recorded on a 1-5 scale described by White and Robson (1989). Plants were harvested at flowering stage, 70 days after planting. Dry weight of the biomass as well as nodule number were recorded.

Statistical design and analysis. The field experiment was planted in a triple (10x10) lattice design and the pot experiment was laid in a randomised complete block design with three replicates. The proc LATTICE and the proc GLM procedures, respectively were used in the statistical analysis of the SAS (1988) programme.

RESULTS AND DISCUSSION

Field experiment. Germination for all accessions in the trial were successful, but 57 of the 100 accessions died within three weeks after planting. The surviving 43 accessions, all landraces, developed well indicating superior tolerance to calcareous soil conditions (Table 3). Some accessions, however, showed symptoms of iron chlorosis. The trial error in the experiment was rather high and the lattice design was only slightly more efficient (103.4 % - 106.9 %) than the randomised complete block design in the combined analyses.

Table 3. Means for the morphological characters, yield and yield components of 43 surviving accessions (Average of random sample of 5 plants)
Accession number* Plant
height (cm)		 Number of branches Number of pots/plant Number of seeds/plant Yield per plant (g) Yield kg ha-1
1 58.6 4.1 15.2 60.9 16.6 1124
2 58.1 3.9 15.0 60.9 16.7 1119
67 55.9 3.9 15.1 60.0 16.2 1111
4 55.3 4.0 15.0 59.4 15.6 1086
5 57.5 3.9 14.9 59.2 16.0 1069
69 57.5 3.9 14.9 58.2 15.2 1055
7 55.1 3.9 14.9 57.9 16.1 924
20 54.1 3.6 13.9 56.2 14.9 888
10 45.2 3.6 13.4 55.1 14.2 883
9 52.1 3.8 14.0 55.0 15.5 876
13 51.6 3.2 12.6 53.1 11.7 875
30 52.1 3.5 13.0 54.0 14.0 875
75 52.1 3.2 12.8 53.9 11.9 869
14 50.8 3.3 11.6 52.8 11.7 861
16 51.1 3.1 11.6 46.0 9.0 722
23 51.2 3.1 11.5 49.4 9.5 722
55 51.0 3.1 11.3 44.3 8.3 708
24 49.5 3.0 11.2 44.2 8.1 525
19 49.4 3.0 11.1 44.0 8.0 472
50 48.0 3.0 9.2 35.2 7.2 361
32 47.3 2.9 8.1 32.8 6.8 347
49 40.1 2.8 5.4 22.2 6.0 338
22 45.5 2.9 7.5 25.0 6.3 333
12 40.0 2.9 5.1 19.4 5.3 313
25 39.5 2.8 5.2 20.0 5.5 300
Giza1 37.5 2.8 4.8 16.2 4.2 277
Giza2 37.5 2.5 4.7 16.0 4.3 236
78 35.9 2.2 4.0 15.0 4.1 219
29 35.0 2.2 4.0 14.3 4.0 211
3 30.9 2.0 3.6 12.3 3.9 208
31 28.9 2.1 3.4 11.3 3.7 194
34 28.9 2.0 3.2 10.2 3.6 180
33 25.3 2.0 3.0 9.6 3.0 166
80 23.6 1.9 2.9 9.4 3.0 155
35 25.4 1.9 2.6 9.2 3.0 152
36 23.1 2.0 2.5 9.1 2.7 138
37 23.1 1.8 2.2 8.5 2.9 136
38 19.8 1.6 1.9 8.3 2.8 133
39 18.7 1.5 2.0 2.7 2.7 125
45 18.0 1.0 1.7 2.8 2.7 125
41 17.4 1.0 1.3 2.7 2.9 111
42 18.2 1.0 1.0 2.8 2.8 111
40 19.9 1.0 1.0 2.6 2.6 105
LSD(0.05) 6.4 0.5 1.0 2.6 1.6 33.0
* Collection number in the Egyptian gene bank

There were significant differences (P<0.001) among genotypes for plant height and number of branches. The highest plants were obtained from the most tolerant accession, and these also produced the highest number of branches. Thus, the most tolerant accession, accession number 1, was 58.6 cm tall with 4 branches per plant and the most susceptible, accession number 40 only 19.9 cm tall with 1 branch (Table 3).

The results of the yield components analysis is presented in Table 3. There was a significant variation in grain yield of the surviving accessions varying from 1124 kg ha-1 for accession 1 to 105 kg ha-1 for accession 40 (Table 3). Seed number per pod varied from 4 to 1 and pod number per plant from 15 to 1. Variation in seed yield varied from 15 to 2.6 grams per plant. Plant height and branch number affected the yield components significantly (P<0.001). The highest seed yield per plant was obtained from the plants with the highest pod number per plant. The ranking of accessions based on the yield component analysis was in good agreement with yields from the whole plot harvest (Table 3). Giza 1 and Giza 2, the two local varieties were tolerant compared to the 57 non surviving accessions and varieties, but results show that a number of the collected landraces were better adapted to the newly reclaimed area with high soil calcium.

Ion concentration and uptake. Table 5 reveals a significant difference in the uptake of N between landraces. The highest N concentration was recorded in accessions with the largest values of dry weight (Table 4).

Table 4. Means of dry weight of shoots and ion uptake (N, Fe, Mn and Ca) (mg g-1 plant-1 dry weight) of the five most tolerant accessions (1, 20, 23, 24 and 12) and the five most susceptible accessions (3, 34, 33, Giza 1 and 40) at 60 days after planting (average of random sample of 5 plants)
Accessions number Dry weight of shoots(g) N Fe Mn Ca
1 3.7 26 1.8 0.5 11.0
20 3.8 30 1.5 0.3 11.5
23 3.6 23 1.8 0.7 11.0
24 3.6 23 2.6 0.6 10.0
12 3.6 28 2.0 0.7 13.0
3 2.9 20 1.9 0.8 13.0
34 2.8 18 2.0 0.5 15.0
33 2.8 17 2.0 0.5 15.0
Giza1 2.6 17 1.9 0.7 10.0
40 2.1 15 2.5 0.4 26.0
LSD(0.05) 0.6 7.3 0.5 0.2 4.3

Table 5. Mean squares of the analysis of variance for ion uptake (N, Fe, Mn and Ca) at 60 days after planting of the five most tolerant accessions (1, 20, 23, 24 and 12) and the five most susceptible accessions (3, 34, 33, Giza 1 and 40)
Determinations Genotype Error Mean C.V.%
DF MS DF MS
N at 60 (DAP) 9 78.70** 18 18.52 21.70 19.83
Fe at 60 (DAP) 9 0.28 18 0.109 2.100 15.78
Mn at 60 (DAP) 9 0.07 18 0.025 0.570 27.80
Ca at 60 (DAP) 9 65.86*** 18 6.433 13.80 18.37
DAP = days after planting.
D.F = degrees of freedom.
M.S = mean squares.
*** indicates significant at 0.1%

The ranking in dry weight was similar to that of plant height. Results suggest a better nitrogen fixation in the calcium tolerant accessions. The susceptible accessions had the highest Ca concentration in the tissues, ranging from 26 mg g-1 plant-1 in the most susceptible accession (number 40) to 11 mg g-1 plant-1 in the resistant accessions number 1 and 23 (Table 4). The results indicate that susceptible accessions had an uncontrolled uptake of Ca as earlier found in L. luteus ( De Silva et al., 1994). There was a non significant variation in the uptake of Fe (1.8-2.5 mg g-1 plant-1) and Mn (0.3-0.8 mg g-1 plant-1), but the rather high concentrations of both elements may be explained by activity of proteoid roots (George et al., 1997). The high Fe concentration also in the most susceptible accessions suggests that tolerance is not caused by low Fe uptake but may be related to translocation in the plant (White and Robson, 1989).

Glasshouse experiment. The data show a large variation in chlorosis. Scores ranged from 0 (leaves green) in tolerant accessions to 5.0 (leaves bright yellow) in susceptible lines. Accessions from Egypt were the most resistant to chlorosis compared to the French varieties (Fig. 1a). In the French varieties chlorosis sensitivity was closely related to the Ca content in the soil. Nodulation were significantly reduced with increasing Ca content in the soil, strongest for the sensitive French varieties( Fig. 1c). Biomass production for the Egyptian accessions tended to be higher than for the French varieties, especially for Giza 1 ( Fig. 1b). The reaction of the tolerant Egyptian accessions compared to the French varieties confirms the results from the field experiment that calcium tolerant genotypes are available among local landraces.

CONCLUSIONS

From the current field and glasshouse experiments it can be concluded that material tolerant to alkaline soil conditions is available in Egyptian landraces. These tolerant accessions can provide a basis for breeding new lupin varieties for new reclaimed areas in Egypt as well as for other parts of the world, where lupin has not yet been adopted due to high soil calcium content. The mechanisms behind the calcium tolerance is still unclear, but a reduced Ca uptake in tolerant plants may play a role. In view of the large trial error found in the field experiment a reliable laboratory test to identify tolerant material should be developed for screening in breeding programmes.

ACKNOWLEDGEMENTS

Germplasm collection of the Egyptian accessions was carried out as part of a joint research project between the Agricultural Research Centre (ARC) in Egypt and the Royal Veterinary and Agricultural University (KVL), Denmark, with financial support from the Danish International Development Assistance (Danida).

REFERENCES

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