Two wheat, Triticum aestivum, cultivars that differed in their ability to tiller were infested by ovipositing Hessian flies, Mayetiola destructor, under similar controlled conditions. Since a larva typically stunts and kills the stem where it feeds and develops, tiller development of fly infested-wheat seedlings is an important plant trait relative to grain yield. \u27Monon\u27 tillered more than \u27Newton at the 0 infestation level (control). \u27Monon\u27 had about the same number of tillers at 0, 1, 2, and 3 puparia (indicative of the number of feeding larvae) per plant; and \u27Newton\u27 had fewer tillers at 0 than 1, 2, or 3 puparia per stem. However, tillering of both cultivars was less at 4 or more puparia per stem, perhaps due to the depletion of plant nutrients. In general, for both cultivars there was a decrease in leaf length, number and wet weight as the number of puparia increased per tiller

Hoxie, Robert P

Wellso, Stanley G

English

Valparaiso University

The Great Lakes Entomologist Volume 27 Number 4 - Winter 1995 Number 4 - Winter 1995 Article 8 December 1995 Tillering Response of 'Monon' And 'Newton' Winter Wheats Infested With Biotype L Hessian Fly (Diptera: Cecidomyiidae) Larvae Stanley G. Wellso Purdue University Robert P. Hoxie Purdue University Follow this and additional works at: https://scholar.valpo.edu/tgle  Part of the Entomology Commons Recommended Citation Wellso, Stanley G. and Hoxie, Robert P. 1995. "Tillering Response of 'Monon' And 'Newton' Winter Wheats Infested With Biotype L Hessian Fly (Diptera: Cecidomyiidae) Larvae," The Great Lakes Entomologist, vol 27 (4) Available at: https://scholar.valpo.edu/tgle/vol27/iss4/8 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. 1994 THE GREAT LAKES ENTOMOLOGIST 235 TILLERING RESPONSE OF 'MONON' AI\ID 'NEWTON' WINTER WHEATS INFESTED WITH BIOTYPE L HESSIAN FLY (DIPTERA: CECIDOMYIIDAE) LARVAE Stanley G. Wellso1 and Robert P. Hoxie2 ABSTRACT Two wheat, Triticum aestivum, cultivars that differed in their ability otiller were infested by ovipositing Hessian flies, Mayetiola destructor, under similar controlled conditions. Since a larva typically stunts and kills the stem where it feeds and develops, tiller development of fly infested-wheat seedlings is an important plant trait relative to grain yield. 'Monon' tillered more than 'Newton at the 0 infestation level (control). 'Monon' had about the same num­ber of tillers at 0, 1, 2, and 3 puparia (indicative of the number of feeding larvae) per plant; and 'Newton' had fewer tillers at 0 than 1, 2, or 3 puparia per stem. However, tillering of both cultivars was less at 4 or more puparia per stem, perhaps due to the depletion of pla t nutrients. In general, for both cultivars there was a decrease in leaf length, number and wet weight as the number of puparia increased per tiller. Wheat, Triticum aestivum, is not as susceptible to damage by phytophag­ous arthropods as many other crops. It is a very resilient crop, and usuallyproduces secondary tillers (shoots r stems). Wheat usua1ll.' has eight tiller buds, but typically only three or four develop int  full sized tIllers (Williams et al. 1975). A few winter heat tillers develop in the autumn or winter, bu  more tillers appear under warm spring temperatures (Simmonds 1987). Kirby (1983) noted that the main shoot and early formed tillers (those formed when leaves 4 to 6 emerge on the main shoot) are most likely to complete development and form grain. An arthropod may feed on the primary tiller and destroy it, while later­developing tillers may be undamaged and produce seeds. Like other grasses, wheat can compensate for damage or injury by producing more stems per unit of area (tillering), seeds and/or heavier seeds per head, and heads per plant (Schlehuber & Tucker 1967). Wheat is thus very adaptable and tolerant of insect attack and rarely requires insecticide treatment. In dollar value in the U ited States in 1984 it ranked fourth among crops in acreage (Anon. 1984), while in insecticidal usage it ranked tenth (Anon. 1988). Both biotic and abiotic conditions influence tillering. Studies of th  tiller­ing process or the production of additional culms in the Gramineae have focused primarily on cultivar differences and the effects of a wide range of 1Insect and Weed Control Research Unit, USDA·ARS, Department of Entomology, Purdue University, W. Lafayette, IN 47907. Current address: 1083 Kona Lane, Bastrop, TX 78602. 2Insect and Weed Control Research Unit, USDA·ARS, Department of Botany and Plant Pathology, Purdue University, W. Lafayette. IN 47907.1Wellso and Hoxie: Tillering Response of 'Monon' And 'Newton' Winter Wheats InfestedPublished by ValpoScholar, 1995236 THE GREAT LAKES ENTOMOLOGIST Vol. 27, No.4 environmental factors. The physiology of tillering has been investigated by studying the effects of various growth substances and inhibitors (Williams et al. 1975, Williams & Langer 1975). In barley, Hordeum vulgare, a reduction of auxin depressed tillering; however, an application of auxin naphthaleneaceti­cacid (NAA) to plants with destroyed apexes increased tillering (Leopold 1964). Little information is available about the interaction of insect numbers and wheat tillering. Multi-tillering wheat varieties may tolerate heavier infesta­tions of the wheat bulb fly, Delia coarctata (Fallin), but this represents g at r pest survival in the following season (Oakley 1980). Successful Hessian fly, Mayetiola destructor (Say), larval infestation of the main stem usually results in s em death, and may result in the productionof tillers. The economic threshold values for the Hessian fly in Central Europe are 1 to 6 larvae per plant (Wetzel and Freier 1981) and 20% larval infestation of stems in North America (Hill et al. 1943). If the infestation is severe, young :elants do ot tiller, but wheat cultivars that tiller freely survived bette  (Barnes 1956). The main resistance mechanism of wheat to the Hessian fly is larval antibiosis, resulting in the death of young larvae due to their inability to maintain sustained feeding (Gallun 1965, Shukle et al. 1990); the resistant plant continued to grow with little evidence of the previous insect infestation. However, Hessian fly biotypes able to overcome host plant resistance are becoming common. Two cultivars that differed in tillering were evaluated under known Hes­sian fly pup ria (larvae) numbers under controlled growth chamber conditions to evaluate the relationship between Hessian fly numbers and wheat tillering, leaf length, fresh weight, and leaf numbers. MATERIALS AND METHODS Test Plants. 'Newton' (less tillering) and 'Monon' (greater tillering com­pared to 'Newton') were selected for study, as they were found previously to differ in their tillering response under controlled conditions (authors unpub­lished data). 'Monon' (H3 gene for resistance, but susceptible to biotype L) and 'Newton' (HO, susceptible) seeds were germinated in moist vermiculite in sin­gle seed containers. Seedlings of each cultivar were transplant d after 5 to 7 d to soil in 24 pots (10 cm diameter), three plan s per pot, and held at 15°C at 14:10 (L:D) photoperiod. The plants were provided with Hoagland's solution once weekly and watered when needed. The experiment was replicated four times. Test Insects. Biotype L Hessian flies virulent to all Hessian fly resistant commercial cultivars currently deployed were used in this study. Hessian flies were originally collected from Indiana wheat fields and maintained by the USDA, ARS Insect and Weed Control Research Laboratory, Purdue Univer­sity. In general, 26 pots per cultivar, 3 plan s per pot were planted and 24 pots with plants 7 days in age were infested with biotype L Hessian flies. This design was replicated four times. Seven days after planting, virulent biotype L Hessian flies were placed to oviposit for 2-7 h on 'Monon' a d 'Newton' caged wheat seedlings with the duration of oviposition dependent upon the the number of eggs observed per plant. The number of eggs per plant were recorded the day after oviposition, and if more eggs than 5-10 were found per plant, the excess eggs were removed with a brush, so that the eggs laid on plants of the two cultivars were about equal. The numbers of puparia (indica­tive of the previous larval infestation), tillers, and the total plant length from 2The Great Lakes Entomologist, Vol. 27, No. 4 [1995], Art. 8https://scholar.valpo.edu/tgle/vol27/iss4/8237 J994 THE GREAT LAKES ENTOMOLOGIST Table 1. Tillering and other plant responses of 'Newton' and 'Monon' winter wheats to bi type L Hessian fly larval feeding No. Puparia Aerial plant responses No. of of per Egg No. of Tillers Length Weight Leaves Puparia Eggs (%) Plants no. mm. mg. no. 0 0 d 0 f 24 3.8ab 343a 2056a 10.7a 1 4.7c 31.7d 57 3.8a 280b I457b 10.2a 2 6.0b 42.5c 32 3.8a 271bc 1448b 9.9a 3 6.3b 52.2c 27 3.4ab 252cd 1113c 8.2b 4 6.7b 65.8ab 26 3.3bc 22ge 834d 7.3c 5 8.9a 58.0bc 10 2.7c 238de 959cd RObc 6_8a F-ratiosb 9.3a 41" 79.6a 14" 12 3.1bc 3' 251cde 32" 1043cd 27** 8.3bc IS" 'Newton' 0 0 e 0 e 24 2.6b 336a 1568a 9.0a 1 4.6d 35.4d 55 3.3a 268b 1091b 8.7a 2 5.5cd 46.9c 31 3.4a 251bc 909bc 7.8b 3 5.8c 56.7bc 45 3.2a 243ed 784cd 7.7b 4 7.1b 59.8b 24 2.7b 227e 637e 6.4c 5 6.9b 75.8a 20 2.4b 228de 680de 6.4c 6-13 8.2a 85.7a 31 2.4b 217e 597e 6.3e F·ratios 43" 26" 7** 50" 32" 20" Puparia (includes 3rd instar8) were removed from the stem 21 d after oviposition. aMonon: 6-8 puparia (mean=6.75), Newton: 6-13 puparia (mean=6.90), no significant differ­gnce (t-te8t). F-ratios from a oneway ANOVA of larval·damage levels within each cultivar; significance: *, P ,,; 0.05; **, P ,,; 0.01. the crown to apex of the longest leaf per plant were recorded 21 dafter oviposition. Statistical Analysis. A one-way analysis of variance (SPSS 1988) was used to test the effects of different leve s of Hessian fly infestation on the two c ltivars on a per plant basis relative to number of tillers, number of leaves, plant length and weight. Differences in significance were identified at the P < 0.05 and P < 0.01 levels and separated by Least Significant Difference Tests. RESULTS AND DISCUSSION Based upon the greater values above ground of the four control plant parameters (leaf and tiller numbers, and plant length and weight) measured under controlled conditions, 'Monon' is a more robust cultivar than 'Newton' (Table 1). Both cultivars were susceptible to virulent biotype L larvae; within each cultivar, the number of tillers and leaves, and plant le gth in general decreased as the infestation level increased. An exception occurred with the plant weight of 'Monon,' where the greatest weight loss occurred at the four puparia per plant level. The cultivars differed in their tillering response to similar Hessian fly levels, which are related to the genetics and physiology of the wheat cultivars. The numbers of tillers of 'Newton' were lower at 0, and 3 or more pupariainfestation level than at 1-2 puparia per plant, indicating th t Hessian fly at the 1 or 2 puparia per plant level promoted tillering in th s cultivar. This was different than th  tillering response of 'Monon'. For 'Monon' within the 0 to 3 3Wellso and Hoxie: Tillering Response of 'Monon' And 'Newton' Winter Wheats InfestedPublished by ValpoScholar, 1995238 THE GREAT LAKES ENTOMOLOGIST Vol. 27, No.4 Table 2. Analysis of variance of tillering and other plant responses to winter wheat cultivars 'Newton' and 'Monon' to biotype L Hessian fly larval feeding. No. of No. of Length Weight No. of Sources of Eggs Tillers mm mg Leaves Variation df F P P df F P F P F P F P 1 6.5 Puparia 5 29.6 ** ** ** ** ** ** Cultivar-Puparia Interaction 5 1.3 NS 1.1 NS 6 0.3 NS 1.0 NS 1.7 NS 1.9 NS Replicate (Date) 3 7.7 ** 8.2 ** 3 7.1 ** 33.9 ** 69.1 ** 33.9 ** Puparia levels: 0, control; 1,2,3,4, 5-number of puparia; 6, 2: 6 puparia. For variables no. of eggs and % puparia! egg, only puparia levels 1 through 6 were used in the ANOVA, because controls were constant zeros. Plant weight was transformed for homogeneity. puparia levels, tillering remained constant, but at greater than 3 puparia, there was less tillering. Although the number of eggs and puparia for b th cultivars within each replicate were similar, the percentages of puparia per egg oviposited were greater for aU infestation levels on 'Newton' than on 'Monon' (Table 1). The resistance associated with the H3 gene of 'Monon' may have adversely affected virulent biotype L larvae or the 'Monon' plants may have been more tolerant to Hessian fly feeding damage. This may also reflect infestation with an impure Hessian fly biotype. The percentages of puparia per egg were lower than expected, especially at the lower infestation levels. Stunting of the main stem due to Hessian fly larval feeding mi ht have caused more nutrients to be available for tillering. Thus, Hessian fly feeding not only stopped elongation of both cultivars, but may have also promoted some tillering. Perhaps as the number of larvae (puparia) pe  plant increased more nutrients were removed (WeUso et al. 1989), ana eventually the ability to tiller decreased. The multiple analysis of variance (Table 2) shows that the cultivars and puparia factors significantly affected all of the parameters examined: The greatest effect of Hessian fly infestation on both cultivars was on plant wet weight, portraying the detrimental effect of this insect on the growth of infested wheat. There was no significant interaction between cultivar-puparia levels; however, the replications differed significantly due to different egg deposition ranges. Sosa & Foster (1976) evaluated the resistance of 'Arthur 71', 'Knox 62', 'Monon', and 'Seneca' (H5, H6, H3, and H7H8 genes for resistance, respec­tively (GaUun 1977) to Hessian fly biotypes GP (Great Plains), B, C, and D at 15 to 27°C. They noted that tillering varied with the Hessian fly biotype, cultivar, and temperature, with most cultivars exhibiting a grea er fly infesta­tion and tillering at higher temperatures. Greater tillering could be a poten­tially important form of tolerance in some cultivars, offsetting fly induced losses. Increased tillering might be particularly beneficial, if used in conjunc­tion with antibiosis. To use Hessian fly infested wheat as a forage crop because it may have more tillers may not be a valid option, s Bun n & Raymer (1989) noted that low to moderate levels of Hessian fly damage reduced wheat forage yield primarily by reducing tiller size and weight rather than tiller density. This experiment establishes hat the response of wheat to virulent Hes­4The Great Lakes Entomologist, Vol. 27, No. 4 [1995], Art. 8https://scholar.valpo.edu/tgle/vol27/iss4/81994 THE GREAT LAKES ENTOMOLOGIST 239 sian fly damage varied relative to the capacity of the cultivar to tiller. If an insect such as the Hessian fly infests and destroys the main stem, tillering becomes an important response for wheat survival. Tillering of these two cultivars varied in response to infestation of similar number  of Hessian fly puparia (larvae), and plants that tiller more at lower infestation levels may have a survival and yield advantage.LITERATURE CITED Anon. 1988. Division of Contaminants Chemistry, FDA. Anon. 1984. Agricultural statistics 1984. U. S. Government Printing Office, Washing­ton, D. C. Barnes, H. F. 1956. Gall midges of cereal crops, the Hessian fly, pp. 95-140. In: H. F. Barnes (ed.), Gall midges of economic importance, Vol. VII. Crosby Lockwood & Son LTD, London. Buntin, G. D. & P. L. Raymer. 1989. Hessian fly (Diptera: Cecidomyiidae) damage and forage production of winter wheat. J. Econ. Entomol. 82: 301-306. Gallun, 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. Hill, C. C., E. J. Udine & J. S. Pinckney. 1943. A method of estimating reduction in yield of wheat caused by Hessian fly infestation. USDA Circ. 663. Kirby, E. J. M. 1983. Development of the cereal plant. pp. 1-3. In: D. W. Wright (ed.), The yield of cereals. Royal Agriculture Society of England, London. 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. Schlehuber, A. M. & B.B. 'lUcker. 1967. Culture of wheat, pp.117-179. In: K. S. Quisen­berry & L. P. Reitz (eds.), Wheat and Wheat Improvement, Agron. Monogr. No. 13. Am. Soc. Agron., Madison. Shukle, R. H., P. B. Grov r, Jr., & J  E. Foster. 1990. Feeding of Hessian fly (Diptera: Cecidomyiidae) larvae on resistant and susceptible wheat. Environ. Entomol. 19:494-500. Simmons, . R. 1987. Growth, development and physiology, pp. 77-113. In: E. G. Heyne (ed), Wheat and Wheat Improvement (2nd Edition), Agron. Monogr. No. 13, Am. Soc. Agron., Madison. Sosa, 0., Jr. & J. E. Foster. 1976. Temperature and expression of resistance in wheat to the Hessian fly. Environ. Entomol. 5:333-336. Wellso, S. G., R. P. Hoxie & C. R. Olien. 1989. Effects of Hessian fly (Diptera: Cecidomy­iidae) larvae and plant age on growth and soluble carbohydrates of Winoka winter wheat. Environ. Entomol. 18:1095-1100. Wetzel, T. & B. Freier. 1981. Bekampfungsrichtwerte fur Schadlinge des Getreides. Nachr.-Bl. Pflanzenschutz DDR 35:47-50. Williams, R. F., B. C., Sharman & R. H. M. Langer. 1975. Growth and development of the wheat tiller. 1. Growth and form of the tiller bud. Aust. J. Bot. 23:715-743. Williams, R. F. & R. H. M. Langer. 1975. Growth and development of the wheat tiller. II. The dynamics of tiller growth. Aust. J. Bot. 23:745-759. 5Wellso and Hoxie: Tillering Response of 'Monon' And 'Newton' Winter Wheats InfestedPublished by ValpoScholar, 1995

Tillering Response of \u27Monon\u27 And \u27Newton\u27 Winter Wheats Infested With Biotype L Hessian Fly (Diptera: Cecidomyiidae) Larvae

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Tillering Response of \u27Monon\u27 And \u27Newton\u27 Winter Wheats Infested With Biotype L Hessian Fly (Diptera: Cecidomyiidae) Larvae

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