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a Univ. of Minnesota Southern Research and Outreach Center, 35838 120th Street, Waseca, MN 59093
b Dep. of Agronomy and Plant Genetics, Univ. of Minnesota, St. Paul, MN 55108
c Dep. of Plant Science, South Dakota State Univ., Brookings, SD 57007
d Dep. of Plant Pathology, Univ. of Minnesota, St. Paul, MN 55108
* Corresponding author (chenx099{at}umn.edu)
| ABSTRACT |
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| INTRODUCTION |
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Growing resistant cultivars is a major SCN management strategy. A number of resistant cultivars in Maturity Groups I and II have been developed for the use in the northern soybean-growing areas of the USA. However, additional strategies are necessary to reduce SCN population density and minimize soybean yield loss (S. Chen, unpublished).
Nematicides have been used to control different nematodes including SCN (Schmitt et al., 1983; Noel, 1987; Sasser and Uzzell, 1991; Smith et al., 1991). Nematicides, however, are not acceptable for control of SCN because they are not cost effective and potentially have negative environmental impacts.
Crop rotation is an effective strategy for SCN management, especially in the southern USA. A number of studies have demonstrated that growing nonhost crops reduced SCN population densities and increased soybean yield (Young and Hartwig, 1992; Koenning et al., 1993; Trevathan and Robbins, 1995; Weaver et al., 1995; Young, 1998; Howard et al., 1998). Two or more years of a nonhost crop resulted in low or undetectable SCN densities and acceptable soybean yields (Francl and Dropkin, 1986; Schmitt, 1991). Koenning et al. (1993) demonstrated that 1 yr of a nonhost crop was sufficient to control SCN with no additional benefit beyond 2 yr of growing a nonhost crop. Soybean cyst nematode survival rate is higher in northern region than in southern regions of the USA (Riggs et al., 2001). Consequently, a longer rotation may be needed for effective management of SCN in the northern soybean-growing regions of the USA.
In a previous study, the effect of long-term corn-soybean rotation on SCN population was investigated in Minnesota (Porter et al., 2001). The crop sequences in that study included 5-yr corn rotated with 5-yr SCN-susceptible soybean, corn-soybean rotation, and monoculture of each crop. Two years of corn generally lowered the number of SCN eggs 100 cm-3 soil from thousands by a factor of 10, and 5 yr of corn lowered number of eggs in 100 cm-3 of soil from the thousands by a factor of 100. In that study, however, there was no soybean crop following 2- to 4-yr corn. In 1996, we initiated an experiment at two sites to determine the effectiveness of rotation involving SCN-resistant and SCN-susceptible soybean with 1 to 3 yr of corn for SCN management in southern Minnesota. In this paper, we report results of the first 4-yr rotation cycle.
| MATERIALS AND METHODS |
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The experiment consisted of 18 treatments arranged in a randomized complete block design with four replicates. At both sites, the 18 treatments were (i) continuous SCN-susceptible soybean (S); (ii) continuous SCN-resistant soybean (R); (iii) continuous corn (C); (iv) and (v) resistant soybean rotated annually with corn (RC and CR); (vi) and (vii) susceptible soybean rotated with 1-yr corn (SC and CS); (viiix) susceptible soybean rotated with 2-yr corn (SCC, CSC, CCS); (xixiv) susceptible soybean rotated with 3-yr corn (SCCC, CSCC, CCSC, CCCS); and (xvxviii) a 4-yr rotation of susceptible soybeancornresistant soybeancorn (SCRC, CRCS, RCSC, and CSCR) (Tables 14). The resistant soybean was Freeborn (resistance derived from PI 88788) (Orf and Young, 1997), the susceptible soybean was Sturdy, and corn hybrids were Pioneer 3730 during 1996 through 1998 and Dekalb 493sr in 1999.
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Data Collection
A soil sample composed of 20 cores was taken from the two central rows of each plot near the plant root zone to a 20-cm depth with a 2.5-cm-diam soil probe at sowing and harvest. The soil samples were stored at -20° C before being processed. Each soil sample was thoroughly mixed. Cysts were extracted with a hand-decanting method. A subsample of 100 cm3 of soil was placed in a 1-L beaker containing 500-mL of water, soaked for at least 30 min and stirred with a motorized stirrer at about 3000 rpm for 3 to 5 min to break soil aggregates. The soil suspension was washed into a 2-L bucket. After a few seconds, the soil suspension was poured through an 850-µm-aperture sieve "nested" on a 250-µm-aperture sieve. The bucket was filled with a strong jet of water again and the suspension was poured on the sieves. This procedure was repeated at least three times for each soil sample. Cysts with debris and soil particles on the 250-µm-aperture sieve were collected, and the cysts were separated from the soil particles and debris with centrifugation in 76% (w/v) sucrose solution at 1500 g. Eggs were released from the cysts by breaking the cysts in a 40-mL glass tissue grinder (Fisher Scientific, Pittsburgh, PA). The egg suspension was poured through a 70-µm-aperture sieve nested on a 38-µm-aperture sieve. Eggs were caught on the 38-µm-aperture sieve and collected into a 50-mL tube and stored at 4°C before being counted within 2 wk. If the egg samples could not be counted within 2 wk, they were stored in a freezer until counted.
Yield of corn and soybean was measured from a 6.7-m length of the two central rows with a small plot combine. The soybean yield was standardized at 130 g kg-1 moisture, and corn yield was standardized at 155 g kg-1.
Data Analysis
For plots in which the egg density at sowing was more than 500 eggs 100 cm-3 soil, the SCN population change (Pf/Pi = egg density at harvest/egg density at sowing) was computed. Nematode egg counts were transformed with log10(x + 1) and Pf/Pi with log10(x) to improve homogeneity of variances before analyses of variance (ANOVA). Average Pf/Pi for each crop were back-transformed. Yields were not transformed for the ANOVA. Least significant difference (LSD, P = 0.05) was used to compare means. Regressions of Pf/Pi against Pi with the linear model ln
= ln
+ bln
derived from the equation
= aPib were performed to determine relationships between the population change rate during the growing season and the initial nematode density, and to determine an equilibrium density (Ferris, 1985). Predicated equilibrium (E = carrying capacity) was determined by solving the equation ln
= ln
+ bln
when Pi = Pf then E = Pi = a1/-b. The relationships between soybean yield and Pi were determined by regression with the equation Y = aebPi or ln
= ln
+ bPi (where Y is soybean yield, Pi is egg density at sowing, a is maximum yield, e is base of the natural logarithm, and b is rate parameter) modified from Appel and Lewis (1984) without determining minimum yield.
| RESULTS |
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In 1997, Pf in susceptible soybean (CS or SS) was not significantly different from the sequence of susceptible soybean in 1996 and corn in 1997 (SC), and these rotations supported higher Pf than sequences without susceptible soybean in 1996 and 1997 (CC, RC, CR, and RR) (Table 2).
In 1998, Pf in susceptible soybean (S = CCS, RCS, SCS, and SSS) ranged from 13 259 to 19 823 eggs 100 cm-3 soil, and were higher than any other sequence. There was no significant difference among Pf in susceptible soybean (S) (Table 3). The sequences in which susceptible soybean was grown either in 1997 or 1996 (CSC, SCC, and SCR) often resulted in higher egg density than the sequence without susceptible soybean (CCC, CRC, RCR, and RRR). The sequence in which resistant soybean rotated annually with corn (CRC and RCR) resulted in the lowest egg density, with values less than 300 eggs 100 cm-3 soil (Table 3).
In 1999, the Pf in susceptible soybean (S = SSSS, CSCS, SCCS, CCCS, and CRCS) ranged from 8206 to 20 913 eggs 100 cm-3 soil, and they were not significantly different regardless of the different crops in the previous 3 yr (Table 4). Egg densities after one season of corn following susceptible soybean (SC = SCSC, CCSC, and RCSC) were still high (657511 988 eggs 100 cm-3 soil) and not significantly different from the sequences with susceptible soybean in 1999 (S) (Table 4). Two-year corn (CSCC) resulted in a Pf of 856 eggs 100 cm-3 soil, which was lower than sequences with 1-yr corn (SC) or susceptible soybean in 1999 (S). The SCN population density after 3-yr corn (SCCC) was 1938 eggs 100 cm-3 soil and not significantly different from that after 2-yr corn (CSCC). The lowest Pf was observed in corn monoculture (CCCC) or corn annually rotated with resistant soybean (RCRC and CRCR). The Pf in monoculture of resistant soybean (RRRR) was higher than in resistant soybean annually rotated with corn (CRCR and RCRC).
Lamberton
Overall, the nematode egg density was higher at the Lamberton site than at the Waseca site. In 1996, SCN population density increased from 14 206 to 30 306 eggs 100 cm-3 soil in susceptible soybean but decreased from 18 458 to 11 011 in resistant soybean and from 12 623 to 10 768 eggs 100 cm-3 soil in corn (Table 1).
In 1997, the highest Pf (14 210 eggs 100 cm-3 soil) was observed in the sequence with susceptible soybean in 1996 and corn in 1997 (SC) (Table 2). Susceptible soybean in monoculture (SS) reduced nematode density from 20 780 to 3800 eggs 100 cm-3 soil, whereas susceptible soybean following corn (CS) increased egg density from 6474 to 9031 eggs 100 cm-3 soil. The reduction of egg density in susceptible soybean may have been caused by early-season flooding, which severely damaged soybean growth. The resistant cultivars (RR and CR) also reduced SCN egg density.
Because of the flooding effect in 1997 on the SCN population, interpreting data of the nematode population in the following years was complicated. Nevertheless, treatment effects on the SCN population were obvious in 1998 and 1999. In 1998, Pf in susceptible soybean (S) ranged from 25 068 to 40 142 eggs 100 cm-3 soil and were generally higher than other sequences (Table 3). The Pf in resistant soybean (RCR, SCR, and RRR) ranged from 6573 to 9523 eggs 100 cm-3 soil and did not significantly differ from 1 or 2 yr of corn following susceptible soybean (11 200 for CSC and 15 816 eggs 100 cm-3 soil for SCC). The lowest Pf was observed in monoculture of corn (CCC) and the rotation of corn-resistant soybean-corn (CRC).
In 1999, the Pf in susceptible soybean (S) were 15 250 to 33 350 eggs 100 cm-3 soil, which did not significantly differ from 1-yr corn following susceptible soybean (SC) (Table 4). The Pf in 3-yr corn (SCCC) was 8219 eggs 100 cm-3 soil; it was significantly different from 1-yr corn following susceptible (SC), but not from 2-yr corn (CSCC). The sequence of 2-yr corn (CSCC), however, resulted in final egg density lower than the sequence of 1-yr corn (SC) or susceptible soybean in 1999 (S). As at Waseca, the lowest egg density was observed in corn monoculture (CCCC) or corn annually rotated with resistant soybean (RCRC or CRCR). The Pf in monoculture of resistant soybean (RRRR) was numerically higher than resistant soybean annually rotated with corn (CRCR and RCRC), but the difference was not statistically significant at P = 0.05.
SCN Population Change during a Single Season
The Pf/Pi in corn ranged from 0.23 to 0.86, except the 1998 Lamberton site, in which Pf/Pi was 1.24 (Table 5). The Pf/Pi in resistant soybean was similar to corn, ranging from 0.21 to 0.73 (1.92 in 1988 at Lamberton). The Pf/Pi in susceptible soybean ranged from 1.76 to 9.91 and was higher than in corn or resistant soybean in all years at both sites except 1997 at the Lamberton site where soybean plants were damaged by flooding (Table 5).
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At the Waseca site, resistant soybean produced 386 and 474 kg ha-1 higher yield than susceptible soybean in 1996 and 1997, respectively (Tables 1 and 2). In 1998 resistant soybean in monoculture (RRR) or rotated annually with corn (RCR) produced higher yield than susceptible soybean in monoculture (SSS), or in annual rotation with corn (SCS) (Table 3). Yield of susceptible soybean following 2-yr corn (CCS) was higher than following 1-yr corn (SCS). In 1999, resistant soybean in annual rotation with corn (CRCR) produced the highest yield (3614 kg ha-1), although it was not significantly different from resistant soybean in other sequences (RRRR and CSCR) or from susceptible soybean in rotation with corn and resistant soybean (CRCS) (Table 4). Susceptible soybean in monoculture (SSSS) produced the lowest yield (1549 kg ha-1). No difference in yield of susceptible soybean was observed among rotations with 1-yr corn (CSCS), 2-yr corn (SCCS), and 3-yr corn (CCCS).
At the Lamberton site, resistant soybean produced 503 kg ha-1 higher yield than susceptible soybean in 1996 (Table 1). In 1997, resistant soybean following corn produced the highest yield (1808 kg ha-1), and susceptible soybean in monoculture produced the lowest yield, only 386 kg ha-1 (Table 2). One-year corn increased the soybean yield for both resistant and susceptible cultivars in the following year (CR vs. RR; CS vs. SS). In 1998, susceptible soybean in monoculture (SSS) produced lower yield than any other treatment (Table 3). Yield of resistant soybean in monoculture (RRR) was lower than the yield of resistant soybean in rotation with corn and susceptible soybean (SCR) and yield of susceptible soybean following 2-yr corn (CCS). In 1999, susceptible soybean in monoculture (SSSS) produced only 959 kg ha-1 and was lower than other treatments except soybean rotated with 2-yr corn (SCCS), which produced 1186 kg ha-1 (Table 4). No significant difference in soybean yield was observed among other treatments.
Yield of susceptible soybean was negatively related with the egg density at sowing. Statistical significance (P < 0.05) of the relationship, however, was observed only in 1998 and 1999 at Waseca, and 1996 and 1999 at Lamberton (Table 7). Yield of resistant soybean was also reduced with increasing egg density at sowing in 1998 and 1999 at Waseca, and in 1997 at Lamberton (Table 7).
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At the Lamberton site, corn following soybean (RC and SC) produced 1674 kg ha-1 higher yield than corn following corn (CC) in 1997 (Table 2). No difference in corn yield was observed among the treatments in 1998 (Table 3). In 1999, corn yield following soybean (RC and SC) was 1341 kg ha-1 higher than yield of corn following corn (CC) (Table 4).
| DISCUSSION |
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Egg density decreased with increasing number of corn years within a rotation sequence, and soybean yield was negatively related to egg density. Therefore, we would expect a susceptible soybean to produce higher yield in the sequences following 2 yr (SCCS) or 3 yr (CCCS) than 1 yr (-SCS) of corn. However, soybean yield benefit by growing more years of corn in the preceding year was observed in some, but not all, years (Tables 14), indicating other factors were involved in the rotation effect on soybean yield.
Growing resistant cultivars was the most effective means to lower SCN population density and increase soybean yield. Therefore, an annual rotation of corn with resistant soybean would be a good choice for SCN management before a susceptible soybean cultivar is grown. On the basis of the data of this study and a previous one (Chen et al., 2001a), if egg density is 20 000 eggs 100 cm-3 soil after growing a susceptible soybean, a 5-yr annual rotation of corn with resistant soybean cultivars is needed before the next susceptible soybean could be grown without significant yield loss. If the egg density is 5000 eggs 100 cm-3 soil, a 3-yr rotation of corn-resistant soybean-corn would be adequate to lower SCN density to where a susceptible soybean could be grown with limited or without yield loss. More years of a resistant cultivar may not further reduce the SCN density because the resistant cultivar supported a limited reproduction.
For an effective use of resistant cultivars in rotation, cultivars with different resistant sources or with different resistant genes should be included in the rotation. If a single resistant cultivar is used in the same field over years, genetic compositions of the SCN populations in the field may be changed (Young, 1984). There was a trend indicating that SCN reproduction potential increased in the resistant cultivar Freeborn. Further study, however, is needed to quantify reproduction potential of the SCN populations in soil where resistant soybean has been grown for several years.
Although resistant soybean produced higher yield than susceptible soybean, yield loss of both resistant and susceptible cultivars to SCN was evident in this study (Table 7). The Pearson correlation coefficients for the relationship between soybean yield and SCN was low (Table 7), indicating that other factors were involved in the variation of soybean yield. This phenomenon was also observed in a previous study (Chen et al., 2001b). The damage to soybean by SCN was more severe at Lamberton in 1997 when plots were flooded than other years without severe flooding. In the wet soil, the nematode penetration or infection may have increased root-rot diseases and caused more damage to soybean plants, a phenomenon that has been demonstrated in a previous greenhouse study (Adeniji et al., 1975). Consequently, there was a greater difference in soybean yield between high egg density and lower egg density (CR vs. RR; and SS vs. CS).
This study showed that SCN reproduction during the growing season was density dependent, a phenomenon that has been described previously (Ferris, 1985). Equilibrium of SCN egg density in soil grown with susceptible soybean was generally more than 10 000 eggs 100 cm-3 soil (Table 6). One season of susceptible soybean could increase egg density from hundreds 100 cm-3 soil to near the equilibrium density. Consequently, there was no difference in egg density at the end of season in susceptible soybean following 1 to 3 yr of corn. Crop rotation with corn or resistant soybean to lower SCN density resulted in higher yield of the resistant soybean and susceptible soybean in the following season, but it did not add any benefit to SCN management for future seasons of susceptible soybean. After one season of susceptible soybean, an extensive rotation with nonhost and resistant soybean was again needed before another crop of susceptible soybean can be grown in order to keep SCN population density low.
The equilibrium egg density is dependent on a number of factors, including environmental conditions, soybean growth, biological antagonists, and nematode fecundity. Further study is needed to determine key factors affecting SCN reproduction and survival to find ways to reduce the SCN population equilibrium and increase the efficiency of crop rotation in SCN management. It appeared that SCN egg density in corn-growing seasons declined faster in a previous study (Porter et al., 2001) than in this study. In the previous study, 2 yr of corn generally lowered egg density from thousands to below 200 eggs 100 cm-3 soil. The reason for the difference in SCN survival between the two studies is unclear. Whether fungal parasites of eggs was a major factor in lowering SCN egg density at the two sites in the previous study (Porter et al., 2001) as compared with the two sites in this study could not be determined or eliminated.
| ACKNOWLEDGMENTS |
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| NOTES |
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Received for publication February 8, 2001.
| REFERENCES |
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