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a P.O Box 262, Soil-Water-Plant Laboratory, Sotuba, Mali
b Dep. of Agronomy, Iowa State Univ., Ames, IA 50011 USA
c Dep. of Entomology, Iowa State Univ., Ames, IA 50011 USA
richard{at}iastate.edu
| ABSTRACT |
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Abbreviations: WD, water deficit treatment Bt, Bacillus thuringiensis ECB, European corn borer GTI, general thermal index LAI, leaf area index
| INTRODUCTION |
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1996, several seed companies have commercialized new transgenic maize hybrids that are resistant to European corn borer (ECB). These new hybrids, commonly known as Bt corn, have been genetically engineered to incorporate genes of Bacillus thuringiensis (Bt) (Koziel et al., 1993; Armstrong et al., 1995). The toxic Bt protein is effective against larvae from both first and second ECB generations. Drought stress is known to have both positive and negative effects on protein synthesis in plants, depending on the duration and intensity of the stress (Shiralipour and West, 1968; Gershenzon, 1984; Mattson and Haack, 1987; Lilburn et al., 1991). It may therefore affect a plant's ability to tolerate stress, and therefore, affect levels of resistance to ECB.
The effect of water deficit on maize growth and development has been studied extensively. The results indicate that water deficit during the vegetative period (before tasseling) can result in shorter plants and smaller leaf area (Denmead and Shaw, 1960; NeSmith and Ritchie, 1992; Abrecht and Carberry, 1993), decreased water use due to the reduction in plant size (Robins and Domingo, 1953), decreased vegetative dry matter (Claassen and Shaw, 1970a), and delayed leaf tip emergence, tassel emergence, silking, and onset of grain filling (NeSmith and Ritchie, 1992; Abrecht and Carberry, 1993). Water deficit during the reproductive period (after tasseling) can increase the interval from silking to pollen shed (Herrero and Johnson, 1981) and shorten the grain filling period (Westgate, 1994). There also is a large amount of literature on the effect of water deficit on different maize yield components. The numerous studies indicate that grain yield can be drastically reduced as a result of water deficit during the reproductive period (Robins and Domingo, 1953; Denmead and Shaw, 1960; Harder et al., 1982; Bennett et al., 1989). This grain yield reduction has been attributed to reduced kernel number, kernel weight, or both (Claassen and Shaw, 1970b; Harder et al., 1982; Grant et al., 1989). Abrecht and Carberry (1993) mentioned, however, that nonlethal water deficit at the beginning of the season did not significantly affect grain yield and the number of kernels per plant. Sinclair et al. (1990) attributed the greater sensitivity of grain yield to water deficit at anthesis to this stage also being the period of maximum biomass accumulation and water use.
The objective of this paper is to report on the growth and development of Bt and non-Bt maize plants subjected to soil water deficit, how these measurements relate to thermal time, and analyze the effect of soil water deficit on the yield components of the two types of maize hybrids. On the same experiment, water relations were measured by infrared thermometry, porometry, and sap flow.
| Materials and methods |
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Irrigation was scheduled so that the plants experienced water deficit 3 to 4 d before the peak first and second ECB egg laying periods. The plants not subjected to water deficit (WD3) were irrigated regularly with 7.2 L (l.9 gallon, 28.8 mm) of water per container. At the time of stress imposition, irrigation was withheld until visual signs of water stress (leaf curling and discoloration) persisted for 3 consecutive days. Then, approximately half a normal watering [3.0 L (0.8 gallon, 11.5 mm)] was supplied to the stressed plants to keep them alive. This was considered a survival irrigation. This procedure was carried out for each of the ECB generations and considered separate water deficit treatments. Figure 1 illustrates the cumulative irrigation water for treatments WD1, WD2, and WD3 in 1997 and 1998. Water deficit also was imposed during other critical periods to get a total number of 24 treatments replicated four times. Treatments consisted of five periods of water deficit imposition and a control and four maize hybrids. All are listed in Table 1 . The four hybrids were [MAXIMIZER 454, CrylAb, event 176, and CIBA 4490 (non-Bt) and NK 7333 Bt, Cryl Ab, event Bt 11, and NK 7333 (non-Bt)]. The treatments were arranged in a randomized complete block design in split plots, with water deficit treatments (WD) as main plots and hybrids (HYB) as subplots. Growth and development were monitored following Ritchie et al. (1993). The monitoring consisted mainly in marking the dates of appearance of each leaf collar and the dates when each leaf was >50% senescent. Observations of tasseling and silking were made on experimental plants, but physiological maturity was observed on five border plants every other day. The border plants were used to avoid destructive sampling from the limited experimental area.
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More recently, Stewart et al. (1998) proposed the general thermal index (GTI) method of calculating thermal time. This method uses a cubic polynomial function instead of a linear one to relate development rate and temperature, thereby avoiding the temperature thresholds characteristic of the previous methods. The polynomial temperature function (FT) is:
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The general thermal index (GTI) is calculated by summing daily values of the FT with time:
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t is the time step in days and n is the number of days in the period of summation (planting to silking, or silking to maturity) (Stewart et al., 1998). The length and maximum width of each leaf were measured after full expansion to estimate leaf area by the Mckee (1964) method. Leaf area index (LAI) was obtained by multiplying the total leaf area by plant density. Crop height was measured approximately every 10 d on each plant by stretching a measuring tape from the soil surface to the collar of the top most fully developed leaf, or the base of the tassel.
The dates of peak egg laying of both first and second ECB generations were determined using observations in surrounding fields. All naturally deposited egg masses were then systematically removed from the maize plants and plants were subsequently infested with approximately 50 newly hatched ECB larvae per plant placed inside leaf sheaths (24 June 1997 and 30 June 1998 for the first generation ECB larvae, 25 July 1997 and 24 July 1998 for the second generation ECB larvae). Fifty is a common number of insects entomologists use to infect their plants (Guthrie and Russell, 1989). At harvest, each plant was split longitudinally, the length of individual tunnels measured (cm), and the total tunneling calculated by summing each. Tunneling by first generation larvae was differentiated from that by second generation larvae based on the darker color of the former.
Growth and development observations were limited to the CIBA Bt and non-Bt hybrids, and water deficit treatments imposed during the first and second ECB generations (WD1 and WD2), respectively, and the controls (WD3) (Table 1). At harvest, the yield components (grain and biomass yield, number of kernels per ear, and 100-seed weight) and ECB tunneling were measured on every plant of all treatments.
Grain was harvested on 11 Sept. 1997 and 7 Sept. 1998, respectively. Grain yield was calculated based on the adjustment to a grain moisture content of 155 g kg-1. Leaves and stalks were harvested separately, and biomass yield was determined by adding grain yield at 155 g kg-1 moisture and the oven-dried weights of leaves and stalks. Statistical analysis of treatment effects was performed using the GLM procedure of SAS (SAS Inst., 1985), and differences among treatment means were considered significant at the 0.05 probability level using the Fisher protected LSD. Orthogonal contrasts compare Bt and non-Bt hybrids.
| Results and discussion |
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, average of 20 leaves per plant), which agrees with the results of Bennett et al. (1989), Muchow and Carberry (1989), and Abrecht and Carberry (1993). However, in 1998, drought-stressed plants had 20 leaves, but the nonstressed ones had only 19 leaves
. The Bt and non-Bt plants did not differ in total leaf number (
in 1997 and
in 1998), and the water deficit x hybrid interaction was not significant for this plant characteristic (
in 1997 and
in 1998).
Water deficit during the first ECB generation delayed tasseling by 3 d in 1998
, but did not have a significant effect in 1997
(Table 2)
. There was no effect of water deficit during the second ECB generation since this treatment was imposed after the end of the vegetative period. The Bt and non-Bt plants did not have different tasseling or silking dates in either of the years. Silking occurred the same day or 1 d after tasseling for both years and for all treatments. The duration of the period from emergence to tasseling was 59 d (683 degree-days) in 1997 and 64 d (758 degree-days) in 1998. A difference of 5 d in the prediction of the date of tassel emergence is significant, particularly near tasseling when up to 8% of grain yield can be lost per day by soil water deficit (Shaw, 1988).
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. In 1998, however, the effect lasted longer, with plants stressed during the first ECB generation remaining significantly shorter than the nonstressed plants from 12 to 22 July (
, respectively) (Fig. 4)
. Gavloski et al. (1992) also observed a decrease in plant height following the withdrawal of water from one or more sections of the root system. The reduction of plant height by drought stress may affect the level of infestation by ECB in natural conditions, since ECB adults are attracted to the taller plants in an area during the first generation egg-laying period (Mason et al., 1996). We did not observe significant differences between the Bt and non-Bt hybrids for crop height.
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in 1997 and 0.01 in 1998) and biomass yield (
in 1997 and 1998) (Table 3)
, and also the number of kernels per ear (
in 1997, and 0.0001 in 1998). The effect on the 100-seed weight was significant in 1998 only (
in 1997 and 0.002 in 1998). The lowest grain yields were observed with treatments WD1 and WD5 in 1997 and with treatments WD2 and WD5 in 1998. Treatments WD2 and WD5 were imposed immediately after tasseling and silking, phases known to be the most sensitive to water deficit regarding grain yield (Robins and Domingo, 1953; Denmead and Shaw, 1960; Claassen and Shaw, 1970b; Harder et al., 1982; Bennett et al., 1989; Grant et al., 1989; Sinclair et al., 1990).
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and for grain in 1998
. Between varieties over both years, the CIBA varieties expressed the smallest grain or biomass yield loss when considering Bt and non-Bt designation. The biomass or grain dry matter losses ranged from 10 to 18% for NK varieties and 0 to 5% for CIBA varieties. For grain yield, this response seemed related to 100-seed weight because seed numbers were not significantly different (Table 3). The CIBA 100-seed weights were similar for Bt designations, whereas, 100-seed weights of NK non-Bt varieties averaged 9% lower than Bt counterparts. This response may be related to ECB tunneling damage, which will be discussed later. Harvest index (grain yield/aboveground plant dry weight) was affected by water deficit only when it was imposed during anthesis (Table 3). Its value was 0.59 in 1997 and 0.53 in 1998 for nonstressed plants. However, with plants stressed at or after tasseling, it was as low as 0.31 in 1997 and 0.28 in 1998. The Bt and non-Bt plants did not differ in harvest index. Infestation with first generation ECB larvae did not result in any significant difference in stalk tunneling between water deficit treatments or hybrids (P > 0.05) (Table 4 and Fig. 5) . There also was no effect of water deficit on the second generation ECB tunneling, and the interaction with hybrid was not significant in 1997 nor 1998. In both 1997 and 1998, second generation ECB tunneling was greater than first generation tunneling (4.74 and 0.49 cm in 1997, 2.17 and 0.16 cm in 1998). This is probably due to the presence of DIMBOA (2,4-dimethyl-7-methoxy-1,4-benzoxazine-3-one), a naturally occurring chemical produced by most commercial maize hybrids that allows plants to combat first generation ECB attacks (Pedigo, 1996).
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in 1997 and
in 1998), but significant from second generation larvae (
in 1997 and
in 1998), with Bt plants affected the least (Fig. 5). Tunneling by second generation larvae averaged 2.01 and 7.47 cm in 1997 and 1.33 and 3.01 cm in 1998 for Bt and non-Bt plants, respectively. Indeed, Bode and Calvin (1990) indicated that yield loss per ECB larva was about 6 to 7% during the first generation and about 2 to 3% during the second generation. | Conclusion |
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Infestation with first generation ECB larvae did not result in any significant difference in stalk tunneling among water deficit treatments or hybrids (Bt or non-Bt). Hybrids did differ in the amount of second ECB generation tunneling, with Bt plants affected the least. Bt plants had greater grain and biomass yields than their non-Bt counterparts, but these differences were significant only for biomass in 1997 and for grain in 1998. The largest differences in grain yield Bt or non-Bt response were associated with the NK variety. The cause for this response is unknown, but could relate to observed ECB tunneling damage. The NK varieties expressed no tunneling damage for Bt plants, whereas CIBA Bt plants did. Thus, the relative tunneling damage difference between Bt and non-Bt plants was probably greater for the NK variety than the CIBA variety. If the tunneling damage affected transport of dry matter to the filling kernel, it is noteworthy that CIBA 100-seed dry weights were quite similar between Bt and non-Bt plants; this was not true for the NK Bt and non-Bt plants.SAS Institute 1985
| ACKNOWLEDGMENTS |
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| NOTES |
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Received for publication June 2, 1999.
| REFERENCES |
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