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Differential response of cowpea lines to aluminum and phosphorusapplicationGani Oladejo Kolawole ab; Guanglong Tian ab; Bir Bahadur Singh ab

a International Institute of Tropical Agriculture (IITA), Ibadan, Nigeria b c/o L. W. Lambourn &Company, Croydon, England

To cite this Article Kolawole, Gani Oladejo, Tian, Guanglong and Singh, Bir Bahadur'Differential response of cowpea linesto aluminum and phosphorus application', Journal of Plant Nutrition, 23: 6, 731 — 740To link to this Article: DOI: 10.1080/01904160009382055URL: http://dx.doi.org/10.1080/01904160009382055

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JOURNAL OF PLANT NUTRITION, 23(6), 731-740 (2000)

Differential Response of Cowpea Lines toAluminum and Phosphorus Application

Gani Oladejo Kolawole, Guanglong Tian,1 andBir Bahadur Singh

International Institute of Tropical Agriculture (IITA), PMB 5320, Ibadan,Nigeria, c/o L. W. Lambourn & Company, Carolyn House, 26 Dingwall Road,Croydon CR9 3EE, England

ABSTRACT

Two pot experiments were conducted, one to evaluate the levels of toleranceof fifteen cowpea [Vigna unguiculata (L.) Walp] lines to aluminum (Al)application, and the second to determine the effect of phosphorus (P) additionon the performance of Al-tolerant lines (IT 91K-93-10, IT 93K-2046-1, and IT90K-2 77-2) and Al-sensitive lines (IT 86D-719, IT 90K-284-2, and IT 89KD-349)in an Alfisol with Al amendment. Fourteen of the fifteen lines tested showeddecreased root biomass (between 19 to 81% reduction) with Al addition, butthis effect was significant for eight of them. Fewer lines showed decreasedshoot biomass and grain yield with Al application. Despite little change innodule number following Al application, there was a significant decrease innodule weight (between 24 and 53% reduction) for nearly all lines. Phosphorusfertilization increased shoot and root biomass, grain yield, nodule number,and weight, and nitrogen (N) and P content of nearly all lines. Al-tolerant linesshowed higher response in shoot and root biomass and nodulation to P

1 Corresponding author (fax #: 234-2241 -2221 ; e-mail address: [email protected]).

731

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732 KOLAWOLE, TIAN, AND SINGH

fertilization than Al-sensitive lines, with the highest response from IT 90K-277-2. Increase in shoot dry weight as a result of P fertilization was from 64 to107% for Al-tolerant lines and from 44 to 48% for the Al-sensitive lines, andincrease in root dry weight was from 46 to 86% for the Al-tolerant lines andfrom 7 to 42% for the Al-sensitive lines. Results of these trials indicated thatlines IT 91K-93-10, IT 93K-2046-1, and IT 90K-277-2 have potential for goodperformance in soil with Al toxicity problems, and that cowpea lines withinherent genetic tolerance to Al will give higher response to P fertilizationwhen grown in soil with Al toxicity problems.

INTRODUCTION

Many well-drained soils in the humid tropics are acid and contain low amountsof basic cations (Kamprath, 1980). Exchangeable Al occupies a large proportion ofthe effective cation exchange capacity in these soils. Concentration of soil solutionAl above 1 cmol (+) kg1 soil often causes direct yield reduction (Sanchez, 1976).Aluminum toxicity is generally considered as the most important potential growth-limiting factor for plants grown in acid soils in many parts of the world (Fageria etal., 1988; Foy, 1996).

There is increasing interest in the inclusion of grain legumes in improvedcropping systems on the acid soils of the lowland humid tropics. The successfulinclusion of grain legumes will depend on the mitigation of the Al toxicity constraint.One of the low input approaches to solving this problem is the selection of speciesor varieties with genetic potential for tolerance to the Al stress associated withacid soils (Foy, 1988). Considerable potential tolerance to soil acidity exists withinthe cowpea species and they have high yield potential even in circumstanceswhere lime is not available (Munns and Fox 1977; Spain et al., 1975; Edwards et al.,1981). Wide differences in Al stress tolerance have been documented among andwithin many agronomic plants (Foy, 1992). Cultivar differences in sensitivity to Altoxicity have been reported for soybean (Horst and Klotz, 1990) and snapbean(Foy et al., 1972). To enhance the cultivation of cowpea in acid soil regions,research is needed to test existing germplasm for Al tolerance and select suitablevarieties.

Aluminum toxicity of plants grown in acid soils is closely related to P nutrition(Bollard, 1983). P fixation in acid soils by precipitation of Al phosphate is animportant factor contributing to low P availability (Foy, 1984). Soil P availabilityduring plant seedling development is an important determinant of growth andgrain yield (Hajabbasi and Schumacher, 1994). Aluminum stress may interfere withP metabolism. However, the interaction between Al and P may differ from onespecies to another and varietal differences within species are possible. This paperreports the results of screenhouse trials to evaluate the differential responses offifteen cowpea lines to Al application, and the effect of phosphorus fertilization onthe performance of Al-tolerant and Al-sensitive lines in an Alfisol.


DIFFERENTIAL RESPONSE OF COWPEA LINES TO Al AND P 733

MATERIALS AND METHODS

Experiment 1

A screenhouse trial was carried out at the International Institute of TropicalAgriculture (UTA), using surface (0-15 cm) soil of an Alfisol (Oxic Paleustalf)collected in Fasola village, Oyo state, southwestern Nigeria. The soil has thefollowing properties: pH-ILp, 6.0; organic carbon (C), 6.5 g kg1 ; total N, 0.5 g kg1;Olsen-extractable P, 7.5 mg kg1; exchangeable potassium (K) 0.26, calcium (Ca)3.68, Al 0.07, and magnesium (Mg) 0.96 cmol (+) kg"1 soil.

The experiment was carried out using a factorial in randomized complete blockdesign with four replications. Fifteen cowpea lines namely; IT 89KD-374, IT 88D-363, Danffla, IT 91K-93-10, Kanannado, TVU-12349, IT 86D-719, IT 93K-734, IT90K-284-2, IT 81D-994, IT 93K-2046-1, IT 93K-1140, IT 89KD-349, TVU-4630, andIT 90K-277-2, from the germplasm collection at UTA were grown in soil (3.5 kg perpot) with two levels of Al (Al2Cl3-6H20): 0 and 90 mg Al kg1 soil (equivalent tocommon Al concentration in Ultisol in Nigeria). All the pots received a basaldressing of 50 mg K (KCl), 50 mg Mg (MgSO4- 7H20), 5 mg zinc (Zn) (ZnSO,,), 10 mgmanganese (Mn) (MnC^H^O), 5 mg copper (Cu) (CuSO4-5H2O), 5 mg Mo[(NH4)6Mo7O24-4H2O] and 5 mg P (NaH2PO4-7H2O) per kg soil. Soil moisture in allpots was equilibrated at 70% of field capacity for 14 days before planting.

Four seeds were sown in each pot on 1 November 1996. One week after plantingthe seedlings were thinned to two plants per pot. The plants were grown tomaturity, pods were harvested and grain weight was determined. The plant shootswere cut at ground level. Roots were washed free of soil with water using a screenwith 1 mm meshsize. Nodules were collected from the roots and nodule numbercounted. Plant shoots, roots and nodules were oven dried at 65°C for 48 hours fordry weight determination. Litterfall during the experimental period was collectedas part of the shoot biomass. Data collected were subjected to analysis of varianceusing SAS package (SAS, 1985).

Experiment 2

Six lines (three Al-tolerant and three Al-sensitive) were selected from Experiment1 to determine the P effect. IT 91K-93-10, IT 93K-2046-1, and IT 90K-277-2 are thetolerant lines shown by the non-decreased grain yield with Al application, whereasIT 86D-719, IT 90K-284-2, and IT 89KD-349 are the Al-sensitive lines shown bylarge decreases in grain yield with Al addition. Two P rates, 0 and 13.1 kg P ha1 assingle super phosphate, were applied to the six selected lines using the sameexperimental design and the same soil type as in Experiment 1. In addition to thebasal dressing of nutrients as described in Experiment 1, 90 mg Al kg"1 soil asA12C13-6H2O was applied to all pots. The soil was then incubated for 14 days beforeplanting. All the other procedures were as described for Experiment 1. Plantshoots were ground in a Wiley mill to pass through a 60 meshsize sieve and later



TABLE 1. Effect of aluminum addition on shoot, root and grain production (g pot"') ofcowpea lines.

Lines

IT 89KD-374

IT88D-363

Danilla

IT91K-93-10

Kanannado

TVU-12349

IT86D-719

IT93K-734

IT90K-284-2

IT81D-994

IT93K-2046-1

IT93K-1I40

IT 89KD-349

TVU-4630

IT 90K-277-2

-A!

6.1

4.7

3.6

7.5

8.8

9.0

7.5

4.5

4.2

6.4

6.0

4.3

3.9

9.5

6.9

Shoot

+A1

3.4

3.0

4.0

6.4

5.4

6.4

5.5

5.4

3.8

5.3

5.3

3.3

2.9

8.3

6.7

West

•

NS

NS

NS

•

*

NS

NS

NS

NS

NS

NS

NS

NS

NS

-Al

0.8

0.9

0.3

1.3

0.7

0.9

1.1

0.9

0.6

1.3

1.0

0.6

0.5

1.3

1.3

Root

+A1

0.3

0.3

0.2

1.1

0.1

0.2

0.3

0.6

0.6

0.5

0.8

0.4

0.3

0.6

0.6

t-test

*

*

NS

NS

*

•

*

NS

NS

•

NS

NS

NS

•

-Al

2.0

3.0

2.7

1.2

0.6

0.4

2.5

2.6

1.9

1.9

0.9

1.9

3.0

0.0

1.9

Grain

+A1

1.0

2.2

1.7

1.6

0.6

0.4

0.3

1.4

1.0

1.2

1.2

1.4

1.7

0.0

2.1

t-test

NS

NS

NS

NS

NS

NS

*

*

•

NS

NS

•

NS

NS

»Significantly different at P=5%; NS=not significant.

analyzed for N and P concentrations using the procedure described by Okalebo etal. (1993).

RESULTS

Experiment 1

Following 2 weeks incubation after application of Al, soil pH was reduced from6.0 to 5.6 and extractable P decreased from 7.5 to 4.6 mg kg"1. Exchangeable Alincreased from 0.07 to 0.09 cmol (+) kg"1 soil. Extractable P, pH, and exchangeableAl in the control treatment were not affected by incubation.

The lines differed in their responses to Al application. Shoot dry weights ofDanilla and IT 93K-734 that received 90 mg Al kg"1 soil were, respectively, 11 and20% higher than the control without Al. Other lines had their shoot dry weightsreduced by between 3% (IT 90K-277-2) and 44% (IT 89KD-374) by Al. However,



TABLE 2. Effect of aluminum addition on nodulation of cowpea lines.

Lines

IT 89KD-374

IT88D-363

Danilla

IT91K-93-10

Kanannado

TVU-12349

IT 86D-719

IT93K-734

IT90K-284-2

IT81D-994

IT93K-2046-1

IT93K-n40

IT 89KD-349

TVU-4630

IT90K-277-2

-Al

33

19

19

48

23

30

28

16

19

28

20

19

18

49

33

Nodule number (no por')

+ A1

13

17

18

3320

31

18

17

20

22

17

16

13

24

31

t-test

NS

NS*

NS

NS

NS

NS

NS

NS

NS

NS

NS*

NS

-Al

28

45

39

40

39

38

41

32

31

40

47

45

38

47

42

Nodule weight (mg pot1)

+ A1

15

31

28

30

30

25

24

17

22

34

35

25

22

22

29

t-tes

**

*

*

*

•

*

•

•

NS•

*

•

•

*

*Significantly different at P=5%; NS=not significant.

statistical analysis showed that the decrease in shoot biomass due to Al applicationwas significant only for three lines, namely IT 89KD-374, Kanannado, and TVU-12349 (Table 1).

Aluminum had more adverse effect on root growth than on shoot growth.Reduction by Al in root growth ranged from 19%(IT91K-93-10)to 81% (Kanannado).More than 50% of the lines showed significant reduction in root biomass due to Al(Table 1).

For most lines, Al adversely affected nodulation (Table 2). More lines showedsignificant reduction in nodule weight than in nodule number. Reduction in noduleweight due to Al was between 24% (IT 81D-994) and 53% (TVU-4630). Among thelines that showed negative response, IT 86D-719, IT 93K-734, IT 90K-284-2, IT81D-994, and IT 89KD-349 showed significant reduction in grain weight due to Al(Table 1).

Experiment 2

Phosphorus fertilization increased shoot and root biomass and grain yield ofnearly all the lines (Table 3). Aluminum-tolerant lines showed higher response in



TABLE 3. Effect of phosphorus application on shoot, root and grain production (g pot"1)of cowpea lines.

Lines

Al-tolerant

IT91K-93-10

ITOK-204S-1

IT90K-277-2

AI-sensitive

IT90K-284-2

IT 86D-719

IT 89KD-349

-P

4.7

3.0

4.1

3.1

4.2

3.1

Shoot

+P

7.6

5.8

8.5

4.4

6.1

4.4

t-test -P

• 0.7

• 0.5

• 0.6

• 0.4

• 0.6

• 0.4

Root

+P

1.2

0.8

1.1

0.4

0.9

0.5

t-test

*

*

*

NS

*

NS

-P

0.9

1.1

1.6

1.0

0.7

1.6

Grain

+P t-test

2.6 •

3.0 •

4.0 •

2.4 •

3.1 •

2.7 NS


TABLE 4. Effect of phosphorus application on nodulation of cowpea lines.

Nodule number (no por1) Nodule weight (mg pot"1)

58

27

17

112

76

58

Lines -P +P t-test -P +P t-test

Al-tolerant

IT91K-93-10 23 46 *

IT93K-2046-1 12 13 NS

IT90K-277-2 14 49 * 17 58 NS

Al-sensitive

IT90K-284-2 14 17 NS

IT86D-719 22 38 *

IT89KD-349 10 11 NS

16

52

12

13

100

17

NS

NS

NS




TABLE 5. Effect of phosphorus fertilization on nitrogen and phosphorus contents (gpot') in shoot of cowpea lines.

Lines

AI-tolerant

IT91K-93-10

IT93K-2046-1

IT90K-277-2

Al-sensltive

IT90K-284-2

IT86D-719

IT89KD-349

-P

13.2

9.9

12.5

10.8

13.3

10.3

N

+P

20.6

15.9

24.6

14.3

18.3

13.0

t - test

*

*

*

NS

*

NS

-P

1.1

1.0

1.3

1.3

1.2

1.1

P

+P t - test

2.2 •

2.5 •

2.5 *

2.5 *

2.6 •

1.8 •

•Significantly different at P=5%; NS=not significant.

shoot and root biomass to P fertilization than Al-sensitive lines, with the bestresponse from IT 90K-277-2. Increase in shoot dry weight was from 64 to 107% forAl-tolerant lines and from 44 to 48% for the Al-sensitive lines, and increase in rootdry weight was from 46 to 86% for the Al-tolerant lines and from 7 to 42% for theAl-sensitive lines.

Higher nodule number and weight were observed with P application (Table 4).Like the biomass, response of nodulation to P was clearly higher in Al-tolerantlines. This difference was particularly pronounced for the nodule weight, wheretwo of the three Al-tolerant lines showed significant response to P, but the threeAl-sensitive lines did not.

Phosphorus fertilization significantly enhanced P content of all lines and Ncontent of Al-tolerant lines, whereas only one of the three Al-sensitive lines showedsignificantly higher N content due to P fertilization (Table 5).

DISCUSSION

Reduction in extractable P after application of Al observed in this study couldbe due to immobilization by the precipitation of Al phosphates in the soil. Increasein exchangeable Al content and decrease in pH value was due to the addition of Al.



Differences in tolerance to Al stress were demonstrated among the fifteen cowpealines studied. Krizek and Foy (1988) have reported a differential response of cropplants to Al. The detrimental effects of Al were clearly reflected in root dry weightof eight out of the fifteen cowpea lines examined. Similarly, reduction in rootbiomass due to Al has been reported for various legume species (Baligar et al.,1989; Campbell et al., 1990; Mookherji and Floyd, 1991).

Phosphorus fertilization significantly enhanced the performance of both Al-tolerant and Al-sensitive lines in the second experiment, indicating the modifyingeffect of P on Al toxicity. Significant differences among the cowpea lines were alsonoted for response to P fertilization. Sanginga et al. (1996) and Tian et al. (1998)observed similar significant genetic variation in P uptake, accumulation and use invarious legumes. The Al-tolerant lines gave higher response to P fertilizationcompared to the Al-sensitive lines (except IT 86D-719), thus indicating that genetictolerance to Al toxicity may be paramount for better response to P fertilization. Theimproved line IT 90K-277-2 was not only tolerant to Al, but it also yielded wellwithout P application and gave maximum response when P was applied.

Root growth of the Al-tolerant lines tested in this trial was significantly enhancedby P fertilization, and there were corresponding increases in N and P contents bythe plants. Tan and Keltjens (1990) have noted that increasing the P supplyexerted certain roles in eliminating Al phytotoxicity possibly through improvedroot development and nutrient uptake. Plant tolerance to Al was observed to beclosely related to their ability to absorb and utilize P in the presence of excess Al(Foy and Brown, 1964).

The results imply that choice of adapted lines and application of P fertilizer canenhance cowpea production in soils with Al toxicity problems. However, cautionshould be applied to extrapolate the findings of these trials to Al-toxic acid soils asother nutrient disorders may modify the overall response.

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Campbell, T. A., N.J. Nuernberg, and C.D. Foy. 1990. Differential responses of red clovergermplasms to aluminum stress. J. Plant Nutr. 13:11, 1463-1474.

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Foy, C.D. and J.C. Brown. 1964. Toxic factors in Acid Soils. II. Differential aluminumtolerance of plant species. Soil Sci. Soc. Am. Proc. 28:27-30.

Foy, C.D., A.L. Fleming, and G.C. Gerloff. 1972. Differential aluminum tolerance of twosnapbean varieties. Agron. J. 64:815-818.

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Horst, W.J. and F. Klotz. 1990. Screening soybean for aluminum tolerance and adaptationto acid s o i l s . pp. 355-360. In: N. El Bassam, M. Dambroth, and B.C. Loughman (eds.),Genetic Aspects of Plant Mineral Nutrition. Kluwer Academic Publishers, Dordrecht,The Netherlands.

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