Secondary tiller resistance of \u27Caldwell\u27 wheat, Triticum aestivum, with the H6 gene for larval resistance to Hessian fly, Mayetiola destructor, was maintained, after the primary culm had been previously infested with virulent larvae. Earlier studies showed that a primary culm infested initially with a virulent larva allowed subsequent normally avirulent larvae to survive on that cultivar; however, in our study the resistance of secondary tillers was mainained even though the primary culm was infested earlier with virulent Hessian fly larvae. This gene stability for resistance is important for optimizing wheat yield of those cultivars that possess genes resistant to the Hessian fly that are tillering and infested with different biotypes

Araya, Jaime E

Wellso, Stanley G

English

Valparaiso University

The Great Lakes Entomologist Volume 26 Number 1 - Spring 1993 Number 1 - Spring 1993 Article 8 April 1993 Resistance Stability of the Secondary Tiller of 'Caldwell' Wheat After the Primary Culm Was Infested With Virulent Hessian Fly (Diptera: Cecidomyiidae) Larvae Stanley G. Wellso Purdue University Jaime E. Araya Purdue University Follow this and additional works at: https://scholar.valpo.edu/tgle  Part of the Entomology Commons Recommended Citation Wellso, Stanley G. and Araya, Jaime E. 1993. "Resistance Stability of the Secondary Tiller of 'Caldwell' Wheat After the Primary Culm Was Infested With Virulent Hessian Fly (Diptera: Cecidomyiidae) Larvae," The Great Lakes Entomologist, vol 26 (1) Available at: https://scholar.valpo.edu/tgle/vol26/iss1/8 This Peer-Review Article is brought to you for free and open access by the Department of Biology at ValpoScholar. It has been accepted for inclusion in The Great Lakes Entomologist by an authorized administrator of ValpoScholar. For more information, please contact a ValpoScholar staff member at scholar@valpo.edu. 1993 THE GREAT LAKES ENTOMOLOGIST 71 RESISTANCE STABILITY OF THE SECONDARY TILLER OF 'CALDWELL' WHEAT AFTER THE PRIMARY CULM WAS INFESTED WITH VIRULENT HESSIAN FLY (DIPTERA: CECIDOMYIIDAE) LARVAE Stanley G. Wellso 1 and Jaime E. Araya 2 ABSTRACT Secondary tiller resistance of 'Caldwell' wheat, Triticum aestivum, with the H6 gene for larval resistance to Hessian fly, Mayetio a destructor, was maintained, after the primary culm had been previously infested with virulent larvae. Earlier studies showed that a primary culm infested initially with a virulent larva allowed subsequent normally avirulent larvae to survive on that cultivar; however, in our study the resistance of secondary tillers was main­tained even though the primary culm was infested earlier with virulent Hes­sian fly larvae. This gene stability for resistance is important for optimizing wheat yield of those cultivars that possess genes r sistant to the Hessian fly that are tillering and infested with different biotypes. Wheat. Triticum aestivum. is a very adaptable crop being grown across a wide range of environments. and leads in production and acreage of all crops (Briggle and Curtis 1987). It is also quite tolerant of insect attack due to its ability to produce secondary tillers. Wheat usually has eight tiller buds, but generally only three or four of these develop fully (Williams and Langer 1975).Thus, an insect may feed upon and destroy the primary culm. while later­developing tillers may not be damaged and often produce seeds. Three major components affect wheat yield: density of fertile heads per unit area, number of seeds Iler head. and seed weight (often expressed a  the weight per 1000 seeds) (Schlehuber and Tucker 1967). At maxImum yield levels, a substantial increase in one yield component generally results in a decrease in one or both of the others. Biotic and abiotic conditions influence tiIlering. Most studies of tillering in Grarnineae have focused on cultivar differences and th  effects of a wide range of environmental factors. The physiology of tillering has been investi­gated by studying the effects caused by various growth substances and inhib­itors (Leopold 1964. Wil iams et al. 1975. Williams and Langer 1975). Little information. however. is available about the impact of insect numbers on wheat tillering. Multi-tillering wheat varieties were able to tolerate heavier infestations of the wheat bulb fly, Delia coarctata (Fallen), but this had the dis~dvantage of greater pest survival the following season (Oakley 1980). 1Insect and Weed Control Research Unit. U.S. Department of Agriculture. Agricul­ture Research Service. Department of Entomology, Purdue University. West Lafayette. Ind. 47907. 2Department of Entomology, Purdue University, West Lafayette, Ind. 47907. Cur­rent Address: Departmento de Sanidad Vegetal. Facultad de Ciencias Agrarias y Fores­tales. Universidad de Chile, Casilla 1004, Santiago. Chile. 1Wellso and Araya: Resistance Stability of the Secondary Tiller of 'Caldwell' WheatPublished by ValpoScholar, 199372 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 Successful Hessian fly, Mayetiola destructor (Say), infestation of the pri­mary culm usually kills it, and may result in the production of tillers; however, young plants do not tiller under severe infestation; abundantlytillering wheat cultivars appear to survive Hessian fly infestations be ter (Barnes 1956). Buntin and Raymer (1989) noted that low to moderate Hessian fly damage levels reduced forage yield primarily by reducing tiller size and weight. ra her than tiller density. Tillering of 'Arthur 71' (H3H5 genes for resistance), 'Monon' (H3), and 'Seneca' (H7Hs) increased when infested with Hessian fly biotypes B, D, or GP at higher temp atures (Sosa and Foster 1976). Tillering of 'Knox 62' (Hs and perhaps H7Hs) was relatively stable when infested with biotypes B, C, D, or GP, and biotype C did not cause an increase in tiller numbers in any of the four cultivars. 'Monon' and 'Newton' wheat cultivars differ in their tillering response when infested by Hessian fly (unpublished data, Wellso and Hoxie). Although 'Newton' tillered less than 'Monon' at the 0 infestation level, infestations of 1-3 puparia (indicative of the number of infesting larvae per seedling) per primary culm resulted in a greater number of tillers. 'Monon' had the same number of tillers at 0-3 puparia per rimary culm; however, both cultivars tillered less at 4 or more puparia than at 1-3 pup aria per primary culm. Thus, the Hessian fly mayor may not increase tillering under light infestations; however, higher infestation levels lessened tillering, perhaps due to the deple­tion of soluble carbohydrates (Wellso et al. 19891. The main resistance mechanism of wheat to the Hessian fly is larval antibiosis, resulting in the death of young larvae due to their inability omaintain sustained feeding (Gallun 1965, Shukle et al. 1990) and the resistant plant continues to grow with little evidence of insect feeding. Grover et al. (1989) noted that infestation by a single virulent lar a resulted in an alteration of resistance of the plant, allowin~ survival of all normally avirulent larvae that concurrently or subse ntly mfested the plant. Their research dealt only with the primary culm. used four cultivars ('Abe', Purdue line 6549, 'Caldwell' and 'Monon') with various combination of infestation with four Hes­sian fly biotypes (B, C, D, and L). Little information is available about the effectiveness of Hessian fly resistant wheat genes when the primary culm is infested with a biotype that is either virulent or avirulent and a secondary tiller is subsequently infested with the same or another biotype. The objective of this study was to evaluate the stability of resistance in the primary culm and secondary tiller of 'Caldwell' wheat infested by virulent or avirulent Hessian fly biotype combinations. MATERIALS AND METHODS Test Insects. Hessian fly biotypes Band L, originally isolated in Indiana approximately 21 and 13 years ago, respectively, have subsequently been maintained in culture at the Insects and Weed Control Research Unit, U.S. Dept. of Agriculture, Purdue University, Ind. To maintain biotype cultures for these studies, mated females oviposited on susceptible wheat (e.g., 'Newton' CI 17715, Ho gene for resistance) maintained in a growth chamber at 21.1 °C, a 16:8 (L:D) photoperiod, light intensity of 300 ,uEinsteins/m2/sec, and 65% R.H. Biotype purity was verified periodically with wheat cultivars having known resistance genes (Sosa and Gallun 1973). Test Plants. Three 'Caldwell' wheat (CI 17897; Hs gene; resistant to Hes­sian fly biotype B, but susceptible to biotype L) seeds, obtained from Purdue University's wheat breeding program, were soaked for 1 h, planted in a mix­2The Great Lakes Entomologist, Vol. 26, No. 1 [1993], Art. 8https://scholar.valpo.edu/tgle/vol26/iss1/81993 THE GREAT LAKES ENTOMOLOGIST 	 73 ture of soil-vermiculite in forty 10 cm diameter plastic pots (with 4 replicates), and later thinned to one seedling p r pot (24 seedlings of each cultivar per treatment). Plants were maintained in a growth chamber set at 21.1 °C, 65 ± 10% RH and 14:10 (L:D), and provided with Hoagland's solution biweekly and watered as needed. Treatments. 'Caldwell' seedlings were exposed to ovipositing Hessian fly as follows: • Control, not infested; • 	Primary culm infested 1 wk after planting with biotype B; • 	Primary culm not infested; secondary tiller infested 3 wk after planting with biotype B; • 	Primary culm infested 1 wk after planting with biotype B; secondary tiller of the same seedling infested 3 wk after planting with biotype B; • 	Primary culm infested 1 wk after planting with biotype L; secondary tiller of the same seedling infested 3 wk later with biotype B. Adult Hessian fly of the desired biotype were released in a growth cham­ber to oviposit on the first or second leaf of the primary culm, and ovipositionwas verified within 24 h. Leaves without eggs were exposed to adults until eggs were found on each seedling. Secondary tiller leaves were infested by confining a mated female in a cylindrical screen cage (2.5 x 12 cm) with plastic foam plugs supported by a metal rod enclosing a normal leaf [as opposed to dark green leaves on a primary culm that previously had been infested suc­cessfully by virulent larvate)]. The cage was removed after oviposition was confirmed. The plants were scored 6 wk after planting relative to the number of tillers. plant ngth from the crown t  the apex of the l ngest leaf, above ground fresh weight, live Hessian fly larvae or puparia and dead larvae (usu­ally small red larvae about 0_5 mm in length). The experiment was replicated on four dates. Data Analysis: After analysis of variance (ANOVA, SAS Institute 1985) using a plit-plot design, significantly different means ( P S 0.05) were sepa­rated by Student-Newman-Keuls test (Steel and Tonie 1980). RESULTS AND DISCUSSION There was a significantly higher (P=0.05) level of infestation of primary culms of 'Caldwell' seedlings by virulent biotype L than by avirulent biotype B larvae (Table 1). Secondary tillers also were resistant to biotype B larvae, including those from seedlings on which the primary culm was infested by virulent biotype L larvae. The mean numbers of tillers (stems) per plant was not significantly differ­ent among the Hessian fly treatments (Table 1/. Plant length and fresh weight did not differ significantly between control plants infested by avirulent bio­type B larvae on either the primary or secondary tiller, however, plant ngth was significantly less when both the primary culm and secondary tiller were infested with biotype B larvae. Virulent biotype L larvae had the greatest effect on plant parameters. Infestation of the primary culm by biotype L larvae resulted in a significant decrease in plant weight, length and number of leaves per plant. The greatest impact to the plant by biotype L larvae was reduction in fresh plant weight, which decreased 50% of that of the control. Wellso et al. (1989. 1990) reported that uninfested pla ts of 'Winoka' and 'Monon', respectively, had significantly greater plant length, weight. and num­ber of leaves than plants infested with virulent larvae. The 50% reduction in fresh plant weight of 'Caldwell' was less than the 65 to 89% reduction in 3Wellso and Araya: Resistance Stability of the Secondary Tiller of 'Caldwell' WheatPublished by ValpoScholar, 1993~ Table 1. Effect of virulent (biotype L) and avirulent (biotype B) larval infestation of the Hessian fly (FH) on the primary culm and/or second;ary -I ::J: mtillers of 'Caldwell' winter wheat in the laboratory" GlTreatmentsb % HF infestation±SD HFc per tiller ± SD ;0Stems Leaves Plant Fresh m Primary Primary Secondary Primary Secondary per plant per plant length plant wt ~ culm tiller culm tiller culm tiller ±SD ±SD (cm±SD) (g±SD) B 2L9± 2.6b 3.7±2.0 3.9± 1.1a 16.8±3.3a 46.3±2.3a 4.8± 1.4a ~ mB 18.8±12.5a 5.8±4.5 4.1±1.4a 18.2±4.1a 45.4±2.9a 6.0± 1.5a (/1 B B 15.6± 12.0b 12.5 ± 1O.2a 1l.8±9.6 2.3±1.0 3.7± 1.2a 16.3±3.7ab 43.2±4.1b 4.4± 1.5a m L B 75.0±49.9a 9.4± 1.2a 8.2±5.4 13.7±6.0 3.6± 1.5a 13.6±1.5a 36.0±6.1c 2.4± 1.3b Z 3.6±O.9v. 15.9±3.2b 46.0±3.0a 4.8±1.3a os:aMeans in the same column followed by the same letter are not significantly different (P:5 0.05), according to a Fisher's Protected LSD test. for o% HF infestation of BB and LB (df =1; F =21.7), stems per plant (df =4; F = 1.1), leaves per plant (df = 4,5; F = 5.9), plant length (df = 4; 5F = 38.9), and fresh plant weight (df = 4; F = 28.9). This experiment included four replicates and 8 plants per replicate. GlbHF biotype on primary colm and/or a secondary tiller. cLarvae and puparia per tiller. ~ 'ti ,~ z 9 4The Great Lakes Entomologist, Vol. 26, No. 1 [1993], Art. 8https://scholar.valpo.edu/tgle/vol26/iss1/81993 THE GREAT LAKES ENTOMOLOGIST 75 weight of Winoka' infested with 1 to 3 or 7 to 9 larvae, respectively, for 8 wk old plants (Wellso et al. 1989). The greater loss in weight of 'Winoka' plants may have been due, in part. to environmental differences between th  test parameters and to the longer duration of he study.The adverse impact (stunting) of infestation by virulent biotype L larvae on development of the primary culm of susceptible 'Caldwell' seedlings was clearly demonstrated in this study. Of significance, however, was the fact that secondary tillers which developed from stunted primary culms (susceptible to biotype L), retained resistance to the avirulent biotype B larvae. This is in contrast to the finding that resistance of the primary culm to normally aviru­lent larvae is lost following feeding by a virulent larva (Grover et al1989). The response f tillers observed in this test would be beneficial where different biotypes are present in the same field, because tillers would be capable of surviving and producing grain, even after the primary culm was destroyed b  a virulent biotype. ACKNOWLEDGMENTS We thank Wyman Nyquist and Judith Santini for statistical assistance, and Clifford Papsdorf for technical assistance. Purdue Univ. Agr. Exp. Stn. Journal Paper No. __ . LITERATURE CITED Barnes, H. G. 1956. Gali midges of economic importance. VII. Gall midges of cereal crops, the Hessian fly, pp. 95-140. Crosby Lockwood and Sons, LTD. London. Briggle. L. W. and B. C. Curtis. 1987. Wheat worldwide, pp. 1-32. In: E G. Heyne (ed.), Wheat and wheat improvement. Agron. Monogr. No. 13. ASACCSA-SSA, Madison, Wisconsin. Buntin, G. D. and P. L. Raymer. 1989. Hessian fly (Diptera: Cecidomyiidae) damage and forage production of winter wheat. J. Econ. Entomol. 82:301-306. Galiun, R. L. 1965. The Hessian fly, and how t  control it. USDA Leaflet 533. __. 1977. Genetic basis of Hessian fly epidemics. Ann. N. Y. Acad. Sci. 287:223-229. Grover, P. B., Jr., R. H. Shukle and J. E. Foster. 1989. Interactions of Hessian fly (Diptera: Cecidomyiidae) biotypes on r istant wheat. Environ. Entomol. 18:687-690. Leopold, A. D. 1964. Plant growth and development. McGraw-Hill, New York. Oakley, J. N. 1980. An integrated control strategy for the wheat bulb fly. ADAS Quart. Rev. 38:145-159. SAS Institute. 1985. SAS user's guide: statistics. 5 h ed. SAS Institute, Cary, N.C. Schlehuber, A. M. and B B. Tucker. 1967. Culture of wheat, pp. 117-179. In: K. S. Quisenberry and L. P. Reitz (ed.) Wheat and wheat improvement. Agron. Monogr. No. 13. ASA-CCSA-SSA, Madison, Wisconsin. ShukIe, R. H., P. B. Grover, Jr. and J. E. Foster. 1990. Feeding of Hessian fly (Diptera: Cecidomyiidae) larvae on resistant and susceptible wheat. Environ. Entomol. 19:494-500. Sosa, 0., Jr. and J. E. Foster. 1976. Temperature and the expression of resistance in wheat to the Hessian fly. Environ. Entomol. 5:333-336. Sasa, 0., Jr. and R. L. Galiun. 1973. Purification of races B and C of the Hessian fly bgenetic manipulation. Ann. Entomol. Soc. Am. 66:1065-1070. Steel, R. G. D. and J. H. Tome. 1980. Principles and procedures of statistics, with special reference to the biological sciences. McGraw Hill, New York. Wellso, S. G., R. P. Hoxie and C. R. Olien. 1989. Effects of Hessian fly (Diptera: Cecido­5Wellso and Araya: Resistance Stability of the Secondary Tiller of 'Caldwell' WheatPublished by ValpoScholar, 199376 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 myiidae) larvae and plant age on growth and soluble carbohydrates of Winoka winter wheat. Environ. Entomol. 18:1095-1100. Wellso. S. G .. R. P. Hoxie and P. L. Taylor. 1990. A comparison of plant parameters and soluble carbohydrates of resistant and susceptible wheat infested with biotype E Hessian flies (Diptera: Cecidomyiidae). Environ. Entomol. 19:1698-1701. Williams, R. F. and R. H. M. Langer. 1975. Growth and development of the wheat tiller. II. The dynamics of tiller growth. Aust. J. Bot. 23:745-759. Williams, R. F., B. C. Sharman and R. H. M. Langer. 1975. Growth and development of the wheat tiller. I. Growth and form of the tiller bud. Aust J. Bot. 23:715-743. 6The Great Lakes Entomologist, Vol. 26, No. 1 [1993], Art. 8https://scholar.valpo.edu/tgle/vol26/iss1/8

Resistance Stability of the Secondary Tiller of \u27Caldwell\u27 Wheat After the Primary Culm Was Infested With Virulent Hessian Fly (Diptera: Cecidomyiidae) Larvae

ValpoScholar

The Great Lakes Entomologist 
Volume 26 
Number 1 - Spring 1993 Number 1 - Spring 1993 Article 8 
April 1993 
Resistance Stability of the Secondary Tiller of 'Caldwell' Wheat 
After the Primary Culm Was Infested With Virulent Hessian Fly 
(Diptera: Cecidomyiidae) Larvae 
Stanley G. Wellso 
Purdue University 
Jaime E. Araya 
Purdue University 
Follow this and additional works at: https://scholar.valpo.edu/tgle 
 Part of the Entomology Commons 
Recommended Citation 
Wellso, Stanley G. and Araya, Jaime E. 1993. "Resistance Stability of the Secondary Tiller of 'Caldwell' 
Wheat After the Primary Culm Was Infested With Virulent Hessian Fly (Diptera: Cecidomyiidae) Larvae," 
The Great Lakes Entomologist, vol 26 (1) 
Available at: https://scholar.valpo.edu/tgle/vol26/iss1/8 
This Peer-Review Article is brought to you for free and open access by the Department of Biology at ValpoScholar. 
It has been accepted for inclusion in The Great Lakes Entomologist by an authorized administrator of ValpoScholar. 
For more information, please contact a ValpoScholar staff member at scholar@valpo.edu. 
1993 THE GREAT LAKES ENTOMOLOGIST 71 
RESISTANCE STABILITY OF THE SECONDARY TILLER OF 'CALDWELL' 

WHEAT AFTER THE PRIMARY CULM WAS INFESTED WITH VIRULENT 

HESSIAN FLY (DIPTERA: 
CECIDOMYIIDAE) 
LARVAE 

Stanley G. Wellso 1 and Jaime E. Araya 2 
ABSTRACT 
Secondary tiller resistance of 'Caldwell' wheat, Triticum aestivum, with 
the H6 
gene for larval resistance 
to Hessian fly, Mayetio a destructor, was 
maintained, after the primary culm had been previously infested with virulent 
larvae. Earlier studies showed that a primary culm infested initially with a 
virulent larva allowed subsequent normally avirulent larvae to survive on that 
cultivar; however, in our study the resistance of secondary tillers was main­
tained even though the primary culm was infested earlier with virulent Hes­
sian fly larvae. This gene stability for resistance is important for optimizing 
wheat yield of those cultivars that possess genes r sistant to the Hessian fly 
that 
are tillering and infested with 
different biotypes. 
Wheat. Triticum aestivum. is a very adaptable crop being grown across a 
wide range of environments. and leads in production and acreage of all crops 
(Briggle and Curtis 1987). It is also quite tolerant of insect attack due to its 
ability to produce secondary tillers. Wheat usually has eight tiller buds, but 
generally only three or four of these develop fully (Williams and Langer 1975).
Thus, an insect may feed upon and destroy the primary culm. while later­
developing tillers may not be damaged and often produce seeds. Three major 
components affect wheat yield: density of fertile heads per unit area, number 
of 
seeds Iler head. 
and seed weight (often expressed a  the weight per 1000 
seeds) (Schlehuber and Tucker 1967). At maxImum yield levels, a substantial 
increase in one yield component generally results in a decrease in one or both 
of the others. 
Biotic and abiotic conditions influence tiIlering. Most studies of tillering 
in Grarnineae have focused on cultivar differences and th  effects of a wide 
range of environmental factors. The physiology of tillering has been investi­
gated 
by 
studying the effects caused by various growth substances and inhib­
itors (Leopold 1964. Wil iams et al. 1975. Williams and Langer 1975). Little 
information. however. is available about the impact of insect numbers on 
wheat tillering. Multi-tillering wheat varieties were able to tolerate heavier 
infestations of the wheat bulb fly, Delia coarctata (Fallen), but this had the 
dis~dvantage 
of greater 
pest survival the following season (Oakley 1980). 
1Insect and Weed Control Research Unit. U.S. Department of Agriculture. Agricul­
ture 
Research Service. 
Department of Entomology, Purdue University. West Lafayette. 
Ind. 47907. 
2Department of Entomology, Purdue University, West Lafayette, Ind. 47907. Cur­
rent 
Address: Departmento de Sanidad Vegetal. Facultad de Ciencias Agrarias y Fores­
tales. Universidad de Chile, Casilla 1004, Santiago. Chile. 
1
Wellso and Araya: Resistance Stability of the Secondary Tiller of 'Caldwell' Wheat
Published by ValpoScholar, 1993
72 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 
Successful Hessian fly, Mayetiola destructor (Say), infestation of the pri­
mary 
culm usually kills it, 
and may result in the production of tillers; however, 
young plants do not tiller under severe infestation; abundantlytillering wheat 
cultivars appear to survive Hessian fly infestations be ter (Barnes 1956). 
Buntin and Raymer (1989) noted that low to moderate Hessian fly damage 
levels reduced forage yield primarily by reducing tiller size and weight. ra her 
than 
tiller density. Tillering of 
'Arthur 71' (H3H5 genes for resistance), 'Monon' 
(H
3
), and 'Seneca' (H7Hs) increased when infested with Hessian fly biotypes B, 
D, 
or 
GP at higher temp atures (Sosa and Foster 1976). Tillering of 'Knox 62' 
(Hs and perhaps H7Hs) was relatively stable when infested with biotypes B, C, 
D, or GP, and biotype C did not cause an increase in tiller numbers in any of 
the 
four cultivars. 
'Monon' 
and 'Newton' wheat cultivars differ in their tillering response 
when infested by Hessian fly (unpublished data, Wellso and Hoxie). Although 
'Newton' tillered less than 'Monon' at the 0 infestation level, infestations of 
1-3 
puparia (indicative of the 
number of infesting larvae per seedling) per 
primary culm resulted in a greater number of tillers. 'Monon' had the same 
number of tillers at 0-3 puparia per rimary culm; however, both cultivars 
tillered less at 4 or more puparia than at 1-3 pup aria per primary culm. Thus, 
the 
Hessian fly 
mayor may not increase tillering under light infestations; 
however, higher infestation levels lessened tillering, perhaps due to the deple­
tion of soluble carbohydrates (Wellso et al. 19891. 
The main resistance mechanism of wheat to the Hessian fly is larval 
antibiosis, resulting in the death of young larvae due to their inability o
maintain sustained 
feeding (Gallun 
1965, Shukle et al. 1990) and the resistant 
plant 
continues 
to grow with little evidence of insect feeding. Grover et al. 
(1989) noted that infestation by a single virulent lar a resulted in an alteration 
of resistance of the plant, allowin~ survival of all normally avirulent larvae 
that 
concurrently or subse 
ntly mfested the plant. Their research dealt only 
with the primary culm. used four cultivars ('Abe', Purdue line 6549, 
'Caldwell' and 'Monon') with various combination of infestation with four Hes­
sian fly biotypes (B, C, D, and L). 
Little information is available about the effectiveness of Hessian fly 
resistant 
wheat genes when 
the primary culm is infested with a biotype that is 
either 
virulent or avirulent 
and a secondary tiller is subsequently infested 
with the same or another biotype. The objective of this study was to evaluate 
the stability 
of resistance in 
the primary culm and secondary tiller of 
'Caldwell' wheat infested by virulent or avirulent Hessian fly biotype 
combinations. 
MATERIALS 
AND 
METHODS 
Test Insects. Hessian fly biotypes Band L, originally isolated in Indiana 
approximately 21 and 13 years ago, respectively, have subsequently been 
maintained in culture at the Insects and Weed Control Research Unit, U.S. 
Dept. of Agriculture, Purdue University, Ind. To maintain biotype cultures for 
these studies, mated females oviposited on susceptible wheat (e.g., 'Newton' 
CI 17715, Ho gene for resistance) maintained in a growth chamber at 21.1 °C, a 
16:8 (L:D) photoperiod, light intensity of 300 ,uEinsteins/m2/sec, and 65% R.H. 
Biotype purity was verified periodically with wheat cultivars having known 
resistance genes (Sosa and Gallun 1973). 
Test Plants. Three 'Caldwell' wheat (CI 17897; Hs gene; resistant to Hes­
sian fly biotype B, but susceptible to biotype L) seeds, obtained from Purdue 
University's wheat breeding program, were soaked for 1 h, planted in a mix­
2
The Great Lakes Entomologist, Vol. 26, No. 1 [1993], Art. 8
https://scholar.valpo.edu/tgle/vol26/iss1/8
1993 THE GREAT LAKES ENTOMOLOGIST 	 73 
ture of soil-vermiculite in forty 10 cm diameter plastic pots (with 4 replicates), 
and later 
thinned 
to one seedling p r pot (24 seedlings of each cultivar per 
treatment). Plants were maintained in a growth chamber set at 21.1 °C, 65 ± 
10% RH and 14:10 (L:D), and provided with Hoagland's solution biweekly and 
watered as needed. 
Treatments. 'Caldwell' seedlings were exposed to ovipositing Hessian fly 
as 
follows: 
• Control, 
not infested; 
• 	Primary culm infested 1 wk after planting with biotype B; 
• 	Primary culm not infested; secondary tiller infested 3 wk after planting with 
biotype B; 
• 	Primary culm infested 1 wk after planting with biotype B; secondary tiller 
of the same seedling infested 3 wk after planting with biotype B; 
• 	Primary culm infested 1 wk after planting with biotype L; secondary tiller 
of the same seedling infested 3 wk later with biotype B. 
Adult 
Hessian fly of 
the desired biotype were released in a growth cham­
ber to 
oviposit on 
the first or second leaf of the primary culm, and oviposition
was verified within 24 h. Leaves without eggs were exposed to adults until 
eggs 
were found on each seedling. Secondary tiller leaves were infested 
by 
confining a mated female in a cylindrical screen cage (2.5 x 12 cm) with plastic 
foam plugs supported by a metal rod enclosing a normal leaf [as opposed to 
dark 
green leaves on a 
primary culm that previously had been infested suc­
cessfully by virulent larvate)]. The cage was removed after oviposition was 
confirmed. The plants were scored 6 wk after planting relative to the number 
of tillers. plant ngth from the crown t  the apex of the l ngest leaf, above 
ground fresh weight, live Hessian fly larvae or puparia and dead larvae (usu­
ally small red larvae about 0_5 mm in length). The experiment was replicated 
on four dates. 
Data 
Analysis: 
After analysis of variance (ANOVA, SAS Institute 1985) 
using a plit-plot design, significantly different means ( P S 0.05) were sepa­
rated by 
Student-Newman-Keuls 
test (Steel and Tonie 1980). 
RESULTS AND DISCUSSION 
There was a significantly higher (P=0.05) level of infestation of primary 
culms of 'Caldwell' seedlings by virulent biotype L than by avirulent biotype B 
larvae (Table 1). Secondary tillers also were resistant to biotype B larvae, 
including those from seedlings on which the primary culm was infested by 
virulent biotype L larvae. 
The mean numbers of tillers (stems) per plant was not significantly differ­
ent 
among 
the Hessian fly treatments (Table 1/. Plant length and fresh weight 
did not differ significantly between control plants infested by avirulent bio­
type 
B larvae on either 
the primary or secondary tiller, however, plant ngth 
was significantly less when both the primary culm and secondary tiller were 
infested with biotype B larvae. Virulent biotype L larvae had the greatest 
effect on plant parameters. Infestation of the primary culm by biotype L 
larvae resulted in a significant decrease in plant weight, length and number of 
leaves per plant. The greatest impact to the plant by biotype L larvae was 
reduction in fresh plant weight, which decreased 50% of that of the control. 
Wellso et al. (1989. 1990) reported that uninfested pla ts of 'Winoka' and 
'Monon', respectively, had significantly greater plant length, weight. and num­
ber of leaves than plants infested with virulent larvae. The 50% reduction in 
fresh plant weight of 'Caldwell' was less than the 65 to 89% reduction in 
3
Wellso and Araya: Resistance Stability of the Secondary Tiller of 'Caldwell' Wheat
Published by ValpoScholar, 1993
~ 
Table 1. Effect of virulent (biotype L) and avirulent (biotype B) larval infestation of the Hessian fly (FH) on the primary culm and/or second;ary -I ::J: 
mtillers of 'Caldwell' winter wheat in the laboratory" 
Gl
Treatmentsb 
% HF infestation±SD HFc per tiller ± SD ;0Stems Leaves Plant Fresh m Primary Primary Secondary Primary Secondary per plant per plant length plant wt ~ 
culm tiller culm tiller culm tiller ±SD ±SD (cm±SD) (g±SD) 
B 2L9± 2.6b 3.7±2.0 3.9± 1.1a 16.8±3.3a 46.3±2.3a 4.8± 1.4a ~ 
mB 18.8±12.5a 5.8±4.5 4.1±1.4a 18.2±4.1a 45.4±2.9a 6.0± 1.5a (/1 
B B 15.6± 12.0b 12.5 ± 1O.2a 1l.8±9.6 2.3±1.0 3.7± 1.2a 16.3±3.7ab 43.2±4.1b 4.4± 1.5a m 
L B 75.0±49.9a 9.4± 1.2a 8.2±5.4 13.7±6.0 3.6± 1.5a 13.6±1.5a 36.0±6.1c 2.4± 1.3b Z 
3.6±O.9v. 15.9±3.2b 46.0±3.0a 4.8±1.3a o
s:aMeans in the same column followed by the same letter are not significantly different (P:5 0.05), according to a Fisher's Protected LSD test. for o% HF infestation of BB and LB (df =1; F =21.7), stems per plant (df =4; F = 1.1), leaves per plant (df = 4,5; F = 5.9), plant length (df = 4; 5F = 38.9), and fresh plant weight (df = 4; F = 28.9). This experiment included four replicates and 8 plants per replicate. Gl
bHF biotype on primary colm and/or a secondary tiller. 
cLarvae and puparia per tiller. ~ 
'ti 
,~ 
z 
9 
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1993 THE GREAT LAKES ENTOMOLOGIST 75 
weight of Winoka' infested with 1 to 3 or 7 to 9 larvae, respectively, for 8 wk 
old plants (Wellso et al. 1989). The greater loss in weight of 'Winoka' plants 
may have been due, in part. to environmental differences between th  test 
parameters and to the longer duration of he study.
The adverse impact (stunting) of infestation by virulent biotype L larvae 
on development of the primary culm of susceptible 'Caldwell' seedlings was 
clearly demonstrated in this study. Of significance, however, was the fact that 
secondary tillers which developed from stunted primary culms (susceptible to 
biotype L), retained resistance to the avirulent biotype B larvae. This is in 
contrast to 
the finding 
that resistance of the primary culm to normally aviru­
lent larvae is lost following feeding by a virulent larva (Grover et al1989). The 
response f tillers observed in this test would be beneficial where different 
biotypes are present in the same field, because tillers would be capable of 
surviving and producing grain, even after the primary culm was destroyed b  
a virulent biotype. 
ACKNOWLEDGMENTS 
We thank 
Wyman Nyquist and 
Judith Santini for statistical assistance, 
and Clifford Papsdorf for technical assistance. Purdue Univ. Agr. Exp. Stn. 
Journal 
Paper No. 
__ . 
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Wellso and Araya: Resistance Stability of the Secondary Tiller of 'Caldwell' Wheat
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76 THE GREAT LAKES ENTOMOLOGIST Vol. 26, No.1 
myiidae) larvae and plant age on growth and soluble carbohydrates of Winoka winter 
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Resistance Stability of the Secondary Tiller of \u27Caldwell\u27 Wheat After the Primary Culm Was Infested With Virulent Hessian Fly (Diptera: Cecidomyiidae) Larvae

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