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Comparative resistance to three species of Callosobruchus in a strain of chickpea (Cicer arietinum L.)
J. stored Prod. Res., 197 1, Vol. 7, pp. 213-216.
Pergamon
Press. Printed
in Great
Britain.
Comparative Resistance to Three Species of Callosobruchus in a Strain of Chickpea (Cicer arietinum L.) A, K. RAINA* United
States
Agency
for
International
Development Regional New Delhi, India
Pulse
Improvement
Project,
(First received 1 June, 1970, and infinulform 14 April, 1971) Abstract-Chickpea is an important source of protein in several countries. In storage, it is infested by Callosobnuhus analis, C. maculatus and C. chinensis. A reportedly resistant strain of the seed was tested with 14 other common varieties. Two methods, selective preference and no choice, were used. The strain named Gl09-1 was significantly better than other varities in being least preferred for egg laying by these bruchids. Gl09-1 has a rough seedcoat which is almost spiny. This spiny character, absent in susceptible varieties, appears to act as a deterent to oviposition.
CHICKPEA or Bengal gram (Cicer arietinum L.) is the most important pulse crop in India. It is grown throughout the country and consumed mostly as whole grain. Among various storage pests, bruchids cause the greatest damage. Three species of Callosobruchus which infest it under storage conditions are C. analis (F.), C. madatus (F.) and C. chinensis L. Chickpeas infested by these are useless for consumption and lose their germination capacity. Control by insecticides or fumigation is costly and assumes the availability of required material and some technical knowledge by the farmer and merchant. Since in developing countries such as India this is not always or consistently the case, incorporating genetic resistance, where possible, in varieties is the best way of preventing serious losses. In the case of grain legumes, PODOLER et al. (1968) did some work on C. chinensis resistance in broad bean (Vi&a faba) varieties. Several people have worked on bruchid resistance in garden peas. The only reference in available literature about chickpea pertains to cutworm resistance and suggests that the internal stem structure in the plants is correlated with resistance to cutworm attack (PAL, 1936). During host preference studies with bruchids, a strain of chickpea, G109-1 (Accession No. 12-069-06629), was found to show resistance to three species of Callosobruchus (SAXENA and RAINA, 1970). The present investigation was undertaken to find the degree of resistance in this strain in comparison with fourteen other common chickpea varieties and to determine the factors responsible for the resistance. + Present address: 58102. S.P.R.7/3--P
Department
of Entomology,
North Dakota
213
State University,
Fargo,
North Dakota
214
A. K. RAINA MATERIALS
AND METHODS
Seeds of G109-1 and 14 other varieties of chickpea were obtained. Only fresh uninfested seeds were selected. Two methods of testing were employed: selective preference (Raina and Gibson, unpublished data) and no choice. In the selective preference method, 5 seeds of each variety were placed in a cavity on a Styrofoam disc which had 15 such cavities along its circumference. The disc was then placed in a desiccator, maintained at 70% r.h and kept inside an incubator at 30°C. Ten freshly mated females were released from a hole in the center of the disc. The females had the opportunity to select among the different varieties for their oviposition. Since these bruchids lay about 95 per cent of their eggs in the first 5 days and these hatch in about 4 days, the seeds were removed on the tenth day and eggs counted on each variety. The seeds were held in vials with muslin cloth covers for the emergence of adults. There were three replicates for each bruchid species. In the no choice method, 25 seeds of each variety were placed in each of the three small plastic jars. This was done for all 15 varieties. Two pairs of freshly emerged adults of a species were released in each of the 45 jars. This experiment was also conducted at 30°C and 70% r.h. for all three species. Eggs laid were counted on the tenth day and the seeds then kept for the emergence of adults. Analysis of variance and Duncan’s multiple range test were conducted on the oviposition data. RESULTS
AND
DISCUSSION
In the selective preference method study, preference was reflected by the number of eggs found on the seed (Table 1). Preference for a particular seed varied among the three species except for G109-1, which was least preferred by all. In the no choice method, if the seed was a preferred one, females laid plenty of eggs on these, as in the case of variety C-104. On the other hand, if they did not prefer it as in G109-1, they either died without laying many eggs or laid them on the sides of the containers. The statistical analysis of the oviposition data showes highly significant F values in all cases. G109-1 was best among the varieties tested as regards negative preference of the bruchids. When comparisons were done, it was found that at least for C. an& and C. chinensis G109-1 did not differ from G62-404 at the 5 per cent probability level. For C. maculatus, the strain G109-1 was outstanding in both methods. Certain large seeded varieties were more susceptible to bruchid damage. There was some similarity in preference exhibited by C. maculatus and C. chine&s in at least 9 varieties of the 15 tested. Since the number of eggs laid by all three species on G109-1 was very small, developmental mortalities could not be compared. The strain G109-1 is therefore quite resistant although not immune to all three species of Callosobruchus tested. Whereas C. maculatus females died without laying eggs on this strain, C. analis and C. chinensis females laid a few eggs, and such eggs showed the normal course of development. Possible factors responsible for the resistance in G 109- 1 were investigated. Examination of the seed coat revealed that it is very rough, almost spiny, compared to that of the other varieties. A similar roughness of the seedcoat but not as marked was also observed in G62-404. In studying relative resistance of some broad bean varieties to C. chinensis, PODOLER et al. (1968) o b served that the thickness of theseedcoat seemed
FI(;. 1. (Top) Vertical section of the seedcoat of G109-1, the resistant strain. I Bottom) Vertical section of the seedcoat of XP-53, a susceptible variety.
[j~zcin~ p. 21 !
18-3cd” 18. 7’d” IO-Ob 28.35 14.7*= IO-Ob 40-00 O-0” 30-O’ 23 - 7de’ 14.3”’ 16-7bcd 3O.Of 25-3” 12.3bc 2.330
11*7*c**t 15.7’6 6-70b 14.O=d 15.3’6 13-o*‘* lo-3b” O-7” 10.7”’ 10*0*= 12.Ob’d 18.3d 18~7~ 15*0Cd 15.Ofd 2.075
c.
maculatus
C. analis
C.
28.3de 28.0dc 10*7’b 36.3’ 34.7’ 15*0bC 55.7” 4.7” 37.oe 35-O” 21.0’d 23 - 7Cd 29.7d= 28 - 7de 22 ’ OCd 2.776
chinensti
17-7bCd 24 * 7de* 8.Ob 24-3def 11*3b 14*0b=* 52.78 0.3” 35.3’ 23~7~~~ 17*3*=‘+ 14*0”‘d 32 - 7”’ 24 - Ode* 12*3b’ 3,369
67.7’ 37 - Ob’d 28.3b 61 .3d’ 39 *3b’d 52.76~de
55 *O=de 67.0” 73.0’ 7.499
35 *76= 8.3” 31*3”C 35.0b’ 80.0f 40 *3bCd
C. madatus
C. analis
No choice method
* n = 3 for all means. t Means with a common letter (within any column) do not differ significantly at 95 per cent level of confidence, $ S = Standard error of means within the column.
a
NP-53 B-98 G-62-484 L-144 NP-58 NP-100 G-736-l G-109-1 Gwalior-2 Chaffa G-24 C-235 c-104 RS-11 BG-482
Variety
Selective preference method
44.7” 26.Ocd 49.3’ 3.947
23-O*’ 13-o* l-3” 31*0Cd 23.0bc 26~3~~ 4.3”
22 ’ ObC 27-3c6 6-7’b lo-7* 37*0d=-
C. chinensis
TABLE 1. AVERAGENUMBER OF EGGSLAIDBY THREESPECIES OF CallosobruchusON FIFTEEN CHICKPEA VARIETIES
-
$ F
=: 0 I. 8 0
2 z
z $ 8 s
T 8.
z. 2
216
A. K. RAINA
to be the only limiting factor for the larvae to penetrate inside the seed. Bruchids are known to prefer smooth seeds for oviposition (SRIVASTAVA and BHATIA, 1959; TEOTIA and SINGH, 1966). Sections of the seedcoats of G109-1 and NP-53 (a susceptible variety) were cut and examined. It was found that the former has sharp protrusions over all the surface (Fig, 1, top). It is assumed that these spiny projections act as a deterrent to oviposition. These projections are markedly absent in susceptible varieties (Fig. 1, bottom). The strain G109- 1 is therefore not preferred by bruchids for oviposition. There may be other factors involved in the mechanism of resistance which need further investigation. author wishes to thank Dr. P. H. VAN SCHAIK,Coordinator, Regional Pulse Improvement Project, for providing facilities and to Dr. M. S. SWAMINATHAN, Director, Indian Agricultural Research Institute, for his interest in the studies. Thanks are also extended to Dr. L. M. JESWANI,Pulse Breeder, IARI, for poviding the seed; to Dr. A. K. KAUL and Dr. R. DHAR of the Genetics Division, IARI, for cutting sections of the seedcoats, and finally to Dr. P. N. SAZXANA, Head Statistician, Agronomy Division, IARI, and Dr. ROBERT B. CARLSON,Entomology Department, North Dakota State University, for their help in the statistical analysis of the data. Acknowledgements-The
REFERENCES PAL, B. P. (1936) A note on the relation between internal stem structure of certain varieties of gram (C&r arietinum L.) and their resistance to cutworm attack. Proc. Indian Acad. Sci. 3,527-534. PODOLER,H. and APPLEBAUM,S. W. (1968) Physiological aspects of host specificity in the BruchidaeV. Varietal differences in the resistance of Vicia faba L. to Callosobruchus chine&s L. J. storedprod. Res. 4, 9-11.
SAXENA,H. P. and RA~NA,A. K. (1970) A bruchid resistant strain of Bengal gram. Curr. Sci. 39, 189-190.
SRIVASTAVA, B. K. and BHATIA,S. K. (1959) The effect of host species on the oviposition of Callosobruchus chinensis
Lin. (Col. Bruchidae). Ann. Zool., Agra 3, 37-42.
TEOTIA,T. P. S. and SINGH,V. S. (1966) The effect of host species on the oviposition, fecundity and development of Callosobruchus chinensis Linn. (Bruchidae: Coleoptera). Bull. Grain Tech&. 4, 3-10.
Pergamon
Press. Printed
in Great
Britain.
Comparative Resistance to Three Species of Callosobruchus in a Strain of Chickpea (Cicer arietinum L.) A, K. RAINA* United
States
Agency
for
International
Development Regional New Delhi, India
Pulse
Improvement
Project,
(First received 1 June, 1970, and infinulform 14 April, 1971) Abstract-Chickpea is an important source of protein in several countries. In storage, it is infested by Callosobnuhus analis, C. maculatus and C. chinensis. A reportedly resistant strain of the seed was tested with 14 other common varieties. Two methods, selective preference and no choice, were used. The strain named Gl09-1 was significantly better than other varities in being least preferred for egg laying by these bruchids. Gl09-1 has a rough seedcoat which is almost spiny. This spiny character, absent in susceptible varieties, appears to act as a deterent to oviposition.
CHICKPEA or Bengal gram (Cicer arietinum L.) is the most important pulse crop in India. It is grown throughout the country and consumed mostly as whole grain. Among various storage pests, bruchids cause the greatest damage. Three species of Callosobruchus which infest it under storage conditions are C. analis (F.), C. madatus (F.) and C. chinensis L. Chickpeas infested by these are useless for consumption and lose their germination capacity. Control by insecticides or fumigation is costly and assumes the availability of required material and some technical knowledge by the farmer and merchant. Since in developing countries such as India this is not always or consistently the case, incorporating genetic resistance, where possible, in varieties is the best way of preventing serious losses. In the case of grain legumes, PODOLER et al. (1968) did some work on C. chinensis resistance in broad bean (Vi&a faba) varieties. Several people have worked on bruchid resistance in garden peas. The only reference in available literature about chickpea pertains to cutworm resistance and suggests that the internal stem structure in the plants is correlated with resistance to cutworm attack (PAL, 1936). During host preference studies with bruchids, a strain of chickpea, G109-1 (Accession No. 12-069-06629), was found to show resistance to three species of Callosobruchus (SAXENA and RAINA, 1970). The present investigation was undertaken to find the degree of resistance in this strain in comparison with fourteen other common chickpea varieties and to determine the factors responsible for the resistance. + Present address: 58102. S.P.R.7/3--P
Department
of Entomology,
North Dakota
213
State University,
Fargo,
North Dakota
214
A. K. RAINA MATERIALS
AND METHODS
Seeds of G109-1 and 14 other varieties of chickpea were obtained. Only fresh uninfested seeds were selected. Two methods of testing were employed: selective preference (Raina and Gibson, unpublished data) and no choice. In the selective preference method, 5 seeds of each variety were placed in a cavity on a Styrofoam disc which had 15 such cavities along its circumference. The disc was then placed in a desiccator, maintained at 70% r.h and kept inside an incubator at 30°C. Ten freshly mated females were released from a hole in the center of the disc. The females had the opportunity to select among the different varieties for their oviposition. Since these bruchids lay about 95 per cent of their eggs in the first 5 days and these hatch in about 4 days, the seeds were removed on the tenth day and eggs counted on each variety. The seeds were held in vials with muslin cloth covers for the emergence of adults. There were three replicates for each bruchid species. In the no choice method, 25 seeds of each variety were placed in each of the three small plastic jars. This was done for all 15 varieties. Two pairs of freshly emerged adults of a species were released in each of the 45 jars. This experiment was also conducted at 30°C and 70% r.h. for all three species. Eggs laid were counted on the tenth day and the seeds then kept for the emergence of adults. Analysis of variance and Duncan’s multiple range test were conducted on the oviposition data. RESULTS
AND
DISCUSSION
In the selective preference method study, preference was reflected by the number of eggs found on the seed (Table 1). Preference for a particular seed varied among the three species except for G109-1, which was least preferred by all. In the no choice method, if the seed was a preferred one, females laid plenty of eggs on these, as in the case of variety C-104. On the other hand, if they did not prefer it as in G109-1, they either died without laying many eggs or laid them on the sides of the containers. The statistical analysis of the oviposition data showes highly significant F values in all cases. G109-1 was best among the varieties tested as regards negative preference of the bruchids. When comparisons were done, it was found that at least for C. an& and C. chinensis G109-1 did not differ from G62-404 at the 5 per cent probability level. For C. maculatus, the strain G109-1 was outstanding in both methods. Certain large seeded varieties were more susceptible to bruchid damage. There was some similarity in preference exhibited by C. maculatus and C. chine&s in at least 9 varieties of the 15 tested. Since the number of eggs laid by all three species on G109-1 was very small, developmental mortalities could not be compared. The strain G109-1 is therefore quite resistant although not immune to all three species of Callosobruchus tested. Whereas C. maculatus females died without laying eggs on this strain, C. analis and C. chinensis females laid a few eggs, and such eggs showed the normal course of development. Possible factors responsible for the resistance in G 109- 1 were investigated. Examination of the seed coat revealed that it is very rough, almost spiny, compared to that of the other varieties. A similar roughness of the seedcoat but not as marked was also observed in G62-404. In studying relative resistance of some broad bean varieties to C. chinensis, PODOLER et al. (1968) o b served that the thickness of theseedcoat seemed
FI(;. 1. (Top) Vertical section of the seedcoat of G109-1, the resistant strain. I Bottom) Vertical section of the seedcoat of XP-53, a susceptible variety.
[j~zcin~ p. 21 !
18-3cd” 18. 7’d” IO-Ob 28.35 14.7*= IO-Ob 40-00 O-0” 30-O’ 23 - 7de’ 14.3”’ 16-7bcd 3O.Of 25-3” 12.3bc 2.330
11*7*c**t 15.7’6 6-70b 14.O=d 15.3’6 13-o*‘* lo-3b” O-7” 10.7”’ 10*0*= 12.Ob’d 18.3d 18~7~ 15*0Cd 15.Ofd 2.075
c.
maculatus
C. analis
C.
28.3de 28.0dc 10*7’b 36.3’ 34.7’ 15*0bC 55.7” 4.7” 37.oe 35-O” 21.0’d 23 - 7Cd 29.7d= 28 - 7de 22 ’ OCd 2.776
chinensti
17-7bCd 24 * 7de* 8.Ob 24-3def 11*3b 14*0b=* 52.78 0.3” 35.3’ 23~7~~~ 17*3*=‘+ 14*0”‘d 32 - 7”’ 24 - Ode* 12*3b’ 3,369
67.7’ 37 - Ob’d 28.3b 61 .3d’ 39 *3b’d 52.76~de
55 *O=de 67.0” 73.0’ 7.499
35 *76= 8.3” 31*3”C 35.0b’ 80.0f 40 *3bCd
C. madatus
C. analis
No choice method
* n = 3 for all means. t Means with a common letter (within any column) do not differ significantly at 95 per cent level of confidence, $ S = Standard error of means within the column.
a
NP-53 B-98 G-62-484 L-144 NP-58 NP-100 G-736-l G-109-1 Gwalior-2 Chaffa G-24 C-235 c-104 RS-11 BG-482
Variety
Selective preference method
44.7” 26.Ocd 49.3’ 3.947
23-O*’ 13-o* l-3” 31*0Cd 23.0bc 26~3~~ 4.3”
22 ’ ObC 27-3c6 6-7’b lo-7* 37*0d=-
C. chinensis
TABLE 1. AVERAGENUMBER OF EGGSLAIDBY THREESPECIES OF CallosobruchusON FIFTEEN CHICKPEA VARIETIES
-
$ F
=: 0 I. 8 0
2 z
z $ 8 s
T 8.
z. 2
216
A. K. RAINA
to be the only limiting factor for the larvae to penetrate inside the seed. Bruchids are known to prefer smooth seeds for oviposition (SRIVASTAVA and BHATIA, 1959; TEOTIA and SINGH, 1966). Sections of the seedcoats of G109-1 and NP-53 (a susceptible variety) were cut and examined. It was found that the former has sharp protrusions over all the surface (Fig, 1, top). It is assumed that these spiny projections act as a deterrent to oviposition. These projections are markedly absent in susceptible varieties (Fig. 1, bottom). The strain G109- 1 is therefore not preferred by bruchids for oviposition. There may be other factors involved in the mechanism of resistance which need further investigation. author wishes to thank Dr. P. H. VAN SCHAIK,Coordinator, Regional Pulse Improvement Project, for providing facilities and to Dr. M. S. SWAMINATHAN, Director, Indian Agricultural Research Institute, for his interest in the studies. Thanks are also extended to Dr. L. M. JESWANI,Pulse Breeder, IARI, for poviding the seed; to Dr. A. K. KAUL and Dr. R. DHAR of the Genetics Division, IARI, for cutting sections of the seedcoats, and finally to Dr. P. N. SAZXANA, Head Statistician, Agronomy Division, IARI, and Dr. ROBERT B. CARLSON,Entomology Department, North Dakota State University, for their help in the statistical analysis of the data. Acknowledgements-The
REFERENCES PAL, B. P. (1936) A note on the relation between internal stem structure of certain varieties of gram (C&r arietinum L.) and their resistance to cutworm attack. Proc. Indian Acad. Sci. 3,527-534. PODOLER,H. and APPLEBAUM,S. W. (1968) Physiological aspects of host specificity in the BruchidaeV. Varietal differences in the resistance of Vicia faba L. to Callosobruchus chine&s L. J. storedprod. Res. 4, 9-11.
SAXENA,H. P. and RA~NA,A. K. (1970) A bruchid resistant strain of Bengal gram. Curr. Sci. 39, 189-190.
SRIVASTAVA, B. K. and BHATIA,S. K. (1959) The effect of host species on the oviposition of Callosobruchus chinensis
Lin. (Col. Bruchidae). Ann. Zool., Agra 3, 37-42.
TEOTIA,T. P. S. and SINGH,V. S. (1966) The effect of host species on the oviposition, fecundity and development of Callosobruchus chinensis Linn. (Bruchidae: Coleoptera). Bull. Grain Tech&. 4, 3-10.