Influence of three resistance sources in winter wheat derived from TAM 107 on yield response to Russian wheat aphid

A study to determine yield response to the Russian wheat aphid, Diuraphis noxia (Mordvilko), was conducted during the 1997-1998 and 1998-1999 growing seasons at three eastern Colorado locations, Akron, Fort Collins, and Lamar, with three wheat lines containing either Russian wheat aphid-resistant Dn4 gene, Dn6 gene, or resistance derived from PI 222668, and TAM 107 as the susceptible control. Russian wheat aphids per tiller were greater on TAM 107 than the resistant wheat lines at the 10x infestation level at Fort Collins and Akron in 1999. Yield, seed weight, and number of seeds per spike for each wheat line were somewhat affected by Russian wheat aphid per tiller mainly at Fort Collins. The infested resistant wheat lines harbored fewer Russian wheat aphids and yielded more than the infested susceptible wheat lines. Wheat lines containing the Dn4, Dn6, and PI 222668 genes contain different levels of antibiosis or antixenosis and tolerance. Although differences existed among sites and resistance, there is a benefit to planting resistant wheat when there is a potential for Russian wheat aphid infestations.     

Response of resistant and susceptible barley to infestations of five Diuraphis noxia (Homoptera: Aphididae) biotypes

Since 2003, four new biotypes of the Russian wheat aphid, Diuraphis noxia (Kurdjumov) (Homoptera: Aphididae), RWA2-RWA5, have been discovered that have the ability to damage most of the wheat germplasm resistant to the original Russian wheat aphid population (RWA1). Barley germplasm lines with resistance to RWA1 have not yet been evaluated against the newest biotypes. Our study compared how biotypes RWA1-RWA5 affected the growth and leaf damage of RWA1-resistant germplasm (STARS 9301B, STARS 9577B), moderately resistant germplasm (MR-015), and susceptible varieties (Schuyler, Harrington, and Morex) under greenhouse conditions. Russian wheat aphid population levels also were determined 14 d after plant infestation. STARS 9301B exhibited strong resistance by showing only small differences in leaf damage and growth parameters from the feeding by the biotypes. STARS 9577B showed greater differences in damage by the Russian wheat aphid biotypes than STARS 9301B, yet, the ratings were still within the resistant category (e.g., chlorosis rating 2.3-4.9). Leaf chlorosis ratings for MR-015 ranged from 5.0 to 6.9 and fell within the moderately resistant to susceptible categories for all the biotypes. The greatest difference in leaf chlorosis occurred in Morex where RWA2 showed less virulence than the other biotypes. Feeding by the Russian wheat aphid biotypes produced only small differences in leaf rolling and plant growth within plant entries. Population levels of the Russian wheat aphid biotypes did not differ within barley entries (n = 610-971) at the termination of the study (14 d). From our research, we conclude that the new Russian wheat aphid biotypes pose no serious threat to the key sources of resistance in barley (STARS 9301B and 9577B).     

Categories of resistance at different growth stages in halt,a winter wheat resistant to the Russian wheat aphid (Homoptera: Aphididae)

This study was designed to categorize the resistance to the Russian wheat aphid, Diuraphis noxia (Mordvilko), resistant hard red winter wheat, Halt, as compared with susceptible wheat, TAM 107, at four different growth stages. Antixenosis was expressed in Halt at growth stage Zadoks 30. Antibiosis in Halt affected fecundity, number of aphids produced per reproductive day, maximum number of nymphs produced in one day, and intrinsic rate of increase. Fecundity was lower on Halt than TAM 107, and more nymphs were produced on both varieties at growth stage 20 than 10 and 40. Fewer nymphs were produced per reproductive day and on maximum production days by aphids reared on Halt than by those reared on TAM 107. The intrinsic rate of increase of Russian wheat aphids reared on Halt was lower than aphids reared on TAM 107. Differences in plant height and plant dry weight did not occur. Chlorosis ratings showed greater damage at the earlier stages in Halt and TAM 107 and significantly more damage in TAM 107 than Halt at growth stages 10, 20, and 30. Leaf rolling occurred on infested plants of TAM 107 at growth stages 10, 20, and 30, but not growth stage 40. Halt plants did not exhibit leaf rolling. The presence of a significant level of tolerance could make Halt compatible with other integrated pest management programs. However, care should be taken with cultivars containing evidence of antixenosis or antibiosis that could cause selective pressure on the Russian wheat aphid, potentially causing biotypes to be produced.     

Reproductive rates of Russian wheat aphid (Hemiptera: Aphididae) biotypes 1 and 2 on a susceptible and a resistant wheat at three temperature regimes

The reproductive rates of Russian wheat aphid, Diuraphis noxia (Kurdjumov) (Hemiptera: Aphididae), Biotype 1 (RWA 1) and Biotype 2 (RWA 2) were compared in the laboratory at three temperature regimes on a Russian wheat aphid resistant cultivar ('Prairie Red') and a susceptible cultivar ('TAM 107'). The objective of this study was to expose RWA 1 and RWA 2 to three temperature regimes and two levels of resistance to find whether there were reproductive differences that may occur within each biotype as well as differences in reproduction between biotypes. In addition, temperature effects of the Dn4 gene on biotype reproduction were noted. Differences in reproductive rates between the two biotypes seem to be driven by temperature. For both biotypes, longevity and reproductive rate parameters, except for intrinsic rate of increase, were lower at the 24-29 degree C temperature regime than the 13-18 degree C and 18-24 degree C temperature regimes. The intrinsic rate of increase was higher for both biotypes at the 18-24 degree C and 24-29 degree C temperature regimes than at the 13-18 degree C temperature regime. Reproductive rates between biotypes were similar at the two higher temperature regimes, but the fecundity for RWA 1 was less than RWA 2 at the 13-18 degree C temperature. The change in fecundity rates between RWA 1 and RWA 2 at lower temperatures could have ecological and geographical implications for RWA 2.     

世界麦双尾蚜研究进展：防治方法和策略*

The Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko), was a worldwide cereal pest. The control measures to this pest were reviewed, emphasizing on natural enemies and plant resistance. First, spring wheat with earlier planting dates had higher yield and could resist RWA infestation to a more extent, while winter wheat with later planting dates could escape infestation of Russian wheat aphid with very few exceptions. So, manipulation of wheat planting dates was suggested in worldwide scale for the aphid control. Second, the natural enemies were considered as the most important factor to reduced the pest status. Introduced and native natural enemies were evaluated for their potential as biological agents in South Africa, United States, and Australia. In South Africa, an introduced parasitoid and a predator were selected for releasing. In the United States, the project on exploring and releasing the natural enemies was unprecedented in biological control history. The endeavor in USA has been proved primarily successful today and will be afterward. The RWA control in Chile was considered most successful, partly because of their introduction of natural enemies before the aphid arrival. The native enemies together with other factors in central Asia and Europe apparently suppressed the aphids to a low level. The screen for resistant wheat was another important research project in fighting with RWA. In South Africa and USA, resistant wheat and barley were bred, and some of them had been put in commercial use for RWA control. The overwhelming mechanisms in resistant wheat varieties were antibiosis, tolerance or their combination. Though chemical insecticide spraying was proved as an effective method for aphid control, more and more research has switched from this method to non chemical control measures as required by IPM. Future research should put more emphasis on augmentation of the natural enemies, revealing the relationship between RWA and agricultural ecosystem and integration of all effective measures.     

麦双尾蚜发生数量与小麦播种期的关系

新疆伊犁和塔城冬、春麦田的麦双尾蚜Diuraphis noxia (Mordvilko) 有虫株率和百株蚜量,与小麦播种时间密切相关。在小麦正常播种期内,冬小麦晚播可以显著减少麦双尾蚜数量,每晚播种10天,第二年麦双尾蚜有虫株率可以下降40%～70%;春小麦每晚播种10天,麦双尾蚜有虫株率增加30%～88%。     

Effect of greenbugs (Homoptera: Aphididae) on yield loss of winter wheat

The effect of greenbug, Schizaphis graminum (Rondani), feeding on the yield of four winter wheat cultivars commonly grown in Oklahoma was studied. Cultivars tested were 'Karl', a recent derivative 'Karl-92', and '2163', all greenbug-susceptible cultivars; and 'TAM-110', a cultivar with resistance to biotype E greenbugs. The objectives were to determine the effect of different greenbug densities during fall and spring on yield of winter wheat, and to develop mathematical models to quantify the effect of greenbugs on yield loss. The intensity of greenbug infestations achieved in plots by artificial infestation varied among years and growing seasons within a year, but was generally sufficient to cause a reduction in yield. Among yield components, the number of heads per square meter and the number of seeds per head were frequently negatively correlated with the accumulated number of greenbug-days per tiller. Seed weight was rarely affected by greenbug infestation. A regression model estimated yield loss for greenbug-susceptible cultivars at 0.51 kg/ha loss of yield per greenbug-day in years with near normal precipitation, and a loss of 1.17 kg/ha under severe drought conditions. The susceptible winter wheat cultivars exhibited similar yield loss in relation to the intensity of greenbug infestation, as indicated by a common slope parameter in the regression model. Results suggest that the model is robust for predicting yield loss for susceptible cultivars.     

麦类品种对麦双尾蚜自然感虫性的初步观察

在新疆伊犁地区1993年和1994年分别观察了195个和36个麦类品种对麦双尾蚜Diuraphis noxia(Mordvilko)的自然感虫性。36个品种两年的结果进行比较表明,感虫率级别相同的品种有18个,占50%;其中“一粒小麦”稳定地表现为不感虫性;感虫率稳定地表现在20%以下的品种有13个,即“86-2-4-2-3-3”、“京772”、“M85 189”、“T579”、“永良13”、“T494”、“浮纳尔”、 “Mg4521”、 “广引74”、“Mg5824”、“Mg8586”、“T1008”和“短芒黑边红”; 感虫率稳定在20%～40%的品种2个,即“额敏黑芒”和“白长穗”; 感虫率稳定在40%～60%的品种2个,即“黑芒红”和“勾毛白”。其余品种,有14个两年的感虫率相差一个级别（20%）;“黑芒长穗”、“高原602”、“伊宁黄库尔班”和“82-10-42-1-1”等4个品种感虫率相差二个或三个级别。另外,1996年塔城引种试验的2个品种“春麦雄性不育系 901”和“啤酒大麦(石引1号)”均遭受麦双尾蚜的严重为害。     

麦双尾蚜及其天敌在不同海拔高度的分布

在新疆塔城,麦双尾蚜Diuraphis noxia (Mordvilko) 在春麦田最集中分布于海拔600～800 m 高度,在冬麦田最集中分布于700～800 m 高度;冬麦田和同海拔高度的春麦田麦双尾蚜数量密切相关(r=0.91, P<0.01)。麦双尾蚜寄生性天敌蚜茧蜂类和蚜小蜂类,在春麦田最集中分布的高度为500～600 m,随着时间的延后,分布的高度范围逐渐扩展;冬麦田最集中分布的高度为600～800 m,略高于春小麦田。捕食麦双尾蚜的斑腹蝇幼虫在春麦田前期最集中分布的高度为500 m处,后期分布高度上升;冬麦田较集中分布的高度为600～800 m,也随着时间的延后也上升。     

新疆塔城地区麦双尾蚜的发生与防治

在新疆塔城地区,麦双尾蚜Diuraphis noxia (Mordvilko) 分布在塔城市、额敏县、裕民县、托里县和布克赛尔县,夏季气候干热的乌苏县和沙湾县未发现麦双尾蚜。根据两年田间试验确定冬、春小麦的防治指标（被害株率）分别为2.8%～11.9%和1.0%～6.2%,麦双尾蚜发生较重的年份和小麦生长早期防治指标较严。在塔城地区,大部分年份麦双尾蚜受自然控制因素制约而为害较轻。     

Differential induction of apoplastic peroxidase and chitinase activities in susceptible and resistant wheat cultivars by Russian wheat aphid infestation

The intercellular peroxidase and chitinase activities of three wheat cultivars [Triticum aestivum L. cvs `Tugela DN', `Molopo DN' (Gariep) and `Betta DN'] containing the Dn-1 gene for resistance to the Russian wheat aphid (RWA) Diuraphis noxia (Mordvilko) and the corresponding near-isogenic susceptible cultivars (`Tugela', `Molopo' and `Betta') were studied under conditions of infestation and non-infestation. The aim was to gain information on the mechanism of resistance. The resistance response was induced by RWA infestation. Infestation rapidly induced the activities of both enzymes selectively in resistant wheat to levels of magnitudes higher than those in susceptible wheat. The genetic background in which the Dn-1 resistance gene is bred played a role and the level of activity corresponded to the level of resistance. Immunologic studies confirmed that the induction of enzyme activities was due to the induction of higher protein levels. These results indicate that peroxidase and chitinase may have a role in insect resistance. Received: 20 June 1997 / Revision received: 9 April 1998 / Accepted: 5 June 1998     

Distribution and diversity of russian wheat aphid (Hemiptera: Aphididae) biotypes in North America

Wheat, Triticum aestivum L., with Russian wheat aphid, Diuraphis noxia (Kurdjumov) (Hemiptera: Aphididae) resistance based on the Dn4 gene has been important in managing Russian wheat aphid since 1994. Recently, five biotypes (RWA1-RWA5) of this aphid have been described based on their ability to differentially damage RWA resistance genes in wheat. RWA2, RWA4, and RWA5 are of great concern because they can kill wheat with Dn4 resistance. In 2005, 365 Russian wheat aphid clone colonies were made from collections taken from 98 fields of wheat or barley, Hordeum vulgare L., in Oklahoma, Texas, New Mexico, Colorado, Kansas, Nebraska, and Wyoming to determine their biotypic status. The biotype of each clone was determined through its ability to differentially damage two resistant and two susceptible wheat entries in two phases of screening. The first phase determined the damage responses of Russian wheat aphid wheat entries with resistance genes Dn4, Dn7, and susceptible 'Custer' to infestations by each clone to identify RWA1 to RWA4. The second phase used the responses of Custer and 'Yuma' wheat to identify RWA1 and RWA5. Only two biotypes, RWA1 and RWA2, were identified in this study. The biotype composition across all collection sites was 27.2% RWA1 and 72.8% RWA2. RWA biotype frequency by state indicated that RWA2 was the predominant biotype and composed 73-95% of the biotype complex in Texas, Oklahoma, Colorado, and Wyoming. Our study indicated that RWA2 is widely distributed and that it has rapidly dominated the biotype complex in wheat and barley within its primary range from Texas to Wyoming. Wheat with the Dn4 resistance gene will have little value in managing RWA in the United States, based on the predominance of RWA2.     

Antioxidative enzymes and the Russian wheat aphid (Diuraphis noxia) resistance response in wheat (Triticum aestivum)

A crucial function of antioxidative enzymes is to remove excess reactive oxygen species (ROS), which can be toxic to plant cells. The effect of Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko), infestation on the activities of antioxidative enzymes was investigated in the resistant (cv. Tugela DN) and the near-isogenic susceptible (cv. Tugela) wheat (Triticum aestivum L.). RWA infestation significantly induced the activity of superoxide dismutase, glutathione reductase and ascorbate peroxidase to higher levels in the resistant than in susceptible plants. These findings suggest the involvement of antioxidative enzymes in the RWA-wheat resistance response, which was accompanied by an early oxidative burst. The results are consistent with the role of ROS in the resistance response and the control of their levels to minimise toxic effects.     

Yield response and categories of resistance to Russian wheat aphid in four Dn4 hard red winter wheat cultivars

A field experiment was conducted to determine whether resistance to Russian wheat aphid, Diuraphis noxia (Mordvilko), conferred by the Dn4 gene is affected by genetic background. This was done by comparing the yield responses to Russian wheat aphid-resistant wheat containing Dn4, derived through the backcross method, to those of the corresponding recurrent parents. Infested resistant cultivars had fewer Russian wheat aphids per tiller than infested susceptible cultivars at the Lamar and Fort Collins, CO sites but not at the Akron, CO site. At the Lamar site, resistant cultivars yielded more than the susceptible cultivars. 'Prairie Red' and 'Yumar' were more resistant than 'Prowers', especially at the higher infestation level. Resistance in these cultivars was categorized in a laboratory experiment to confirm this differential expression of resistance. Resistance in Prairie Red, 'Halt', 'Prowers 99', and Yumar was categorized at three plant growth stages. Antibiosis was expressed as reductions in maximum number of nymphs produced per 24 h and intrinsic rate of increase. The maximum number of nymphs produced per 24 h was reduced in Halt and 'Lamar'. Averaged over cultivars, the intrinsic rate of increase was less at jointing than at the seedling or tillering growth stages. Tolerance was expressed in the resistant cultivars as reduced chlorosis and leaf rolling. Growth reductions in infested Prowers 99 plants was less than the other cultivars. This study confirms that some cultivars containing Dn4 may express antibiosis and tolerance, whereas others may not show the same categories. Thus, expression is affected by genetic background.     

The reactive oxygen species are involved in resistance responses of wheat to the Russian wheat aphid

The effect of Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko), infestation on the hydrogen peroxide (H(2)O(2)) content and NADPH oxidase (EC 1.6.3.1) activity was studied in the resistant (cv. Tugela DN) and near-isogenic susceptible (cv. Tugela) wheat (Triticum aestivum L.). The objective of this study was to investigate the involvement of the reactive oxygen species (ROS) during the resistance responses against the RWA. Infestation significantly induced an early accumulation of the H(2)O(2) and increase of NADPH oxidase activity to higher levels in the resistant than susceptible plants. Results of inhibitory studies using diphenylene iodonium (DPI), a suicide inhibitor of NADPH oxidase, strongly suggested a possible signalling role for H(2)O(2) during RWA resistance response by activation of downstream defence enzymes [intercellular peroxidase (EC 1.11.1.7) and beta-1,3-glucanase (EC 3.2.1.39)].     

Influence of the Russian wheat aphid <Emphasis Type="Italic">Diuraphis noxia</Emphasis> (Kurdjumov) (Homoptera,Aphididae) on productive qualities of spring bread wheat and barley grown from the seeds from aphid-infested spikes

The aftereffects of the Russian wheat aphid (RWA) Diuraphis noxia on sowing and productive qualities of barley and spring bread wheat grain were assessed. Seeds of 4 cultivars of barley (Volgar, Povolzhsky 65, Kazak, and Povolzhsky 16) and 4 cultivars of spring wheat (Kinelskaya 59, Kinelskaya Otrada, Kinelskaya Niva, and Kinelskaya 2010) from spikes infested and uninfested with RWA in 2007 and in 2014 were sown in the subsequent years, using 0.5 m² experimental plots in four replications, at a seeding rate of 300 grains/m². The least significant difference (LSD_0.5) was used to compare the mean ± standard deviation (SD) values. The field germination rate of seeds from spring wheat spikes damaged by RWA was reduced by 15%. Of the components of grain yield, barley and spring wheat grown from seeds from the infested spikes showed a 23-31% smaller number of productive tillers before harvesting, a 16% smaller number of grains per spike, a 13-16% lower grain weight per spike, and a total yield loss of 33-42%. In hulless bread wheat RWA fed on the developing kernels inflicting greater damage, whereas the hulled barley seeds were practically not damaged. The mean yield loss of the barley and spring wheat spikes infested with RWA was 24-32% and 50-66%, respectively. Due to the greater tillering capacity and formation of secondary productive tillers in barley, about 52% of the productive barley tillers and 37-39% of spring wheat ones were infested with RWA, which resulted in a comparable yield loss (20-25% in barley and 19-23% in spring wheat). Resistance to RWA was higher in spring wheat and barley cultivars with a shorter vegetation period, looser spikes, and thinner culm walls. The length of productive tillers damaged by RWA was reduced by 21-28%, which determined a lower incidence of leaf diseases.     

Physiological and biochemical responses of resistant and susceptible wheat to injury by Russian wheat aphid

We examined the physiological and biochemical responses of resistant ('Halt' and 'Prairie Red') and susceptible ('TAM 107') wheat, Triticum aestivum L., to injury by the Russian wheat aphid, Diuraphis noxia (Mordvilko). Photosynthetic capacity was evaluated by measuring assimilation/internal CO2 (A/Ci) curves, chlorophyll fluorescence, chlorophyll, and nonstructural carbohydrate content. Total protein and peroxidase specific activity also were determined. No significant differences were detected in chlorophyll concentration between aphid-infested and control TAM 107 plants. The aphid-infested resistant cultivars had similar or significantly higher chlorophyll concentrations compared with their respective control plants. Measurements over time showed that infested Halt plants had delays in photosynthetic senescence, Prairie Red plants had photosynthetic rate changes that were similar to control plants, and TAM 107 plants displayed accelerated photosynthetic senescence patterns. The photochemical and nonphotochemical quenching coefficients were significantly higher in infested Halt plants compared with their respective control plants on day 3. Infested TAM 107 plants had significantly higher photochemical quenching compared with control plants at all times evaluated, and they had significantly higher nonphotochemical quenching on day 3. Throughout the experiment, infested Prairie Red plants exhibited photochemical and nonphotochemical quenching coefficient values that were not significantly different from control plants. Total protein content was not significantly different between aphid-infested and control plants for all cultivars. Differences between physiological responses of infested susceptible and resistant cultivars, particularly temporal changes in photosynthetic activity, imply that resistant Halt and Prairie Red wheat tolerate some impacts of aphid injury on photosynthetic integrity.     

Biotypic diversity in Colorado Russian wheat aphid (Hemiptera: Aphididae) populations

The biotypic diversity of the Russian wheat aphid, Diuraphis noxia (Kurdjumov) (Hemiptera: Aphididae), was assessed in five isolates collected in Colorado. Three isolates, RWA 1, RWA 2, and an isolate from Montezuma County, CO, designated RWA 6, were originally collected from cultivated wheat, Triticum aestivum L., and obtained from established colonies at Colorado State University. The fourth isolate, designated RWA 7, was collected from Canada wildrye, Elymus canadensis L., in Baca County, CO. The fifth isolate, designated RWA 8, was collected from crested wheatgrass, Agropyron cristatum (L.) Gaertn., in Montezuma County, CO. The four isolates were characterized in a standard seedling assay, by using 24 plant differentials, 22 wheat lines and two barley, Hordeum vulgare L., lines. RWA 1 was the least virulent of the isolates, killing only the four susceptible entries. RWA 8 also killed only the four susceptible entries, but it expressed intermediate virulence on seven wheat lines. RWA 6, killing nine entries, and RWA 7, killing 11 entries, both expressed an intermediate level of virulence overall, but differed in their level of virulence to 'CO03797' (Dn1), 'Yumar' (Dn4), and 'CO960293-2'. RWA 2 was the most virulent isolate, killing 14 entries, including Dn4- and Dny-containing wheat. Four wheat lines, '94M370' (Dn7), 'STARS 02RWA2414-11', CO03797, and 'CI2401', were resistant to the five isolates. The results of this screening confirm the presence of five unique Russian wheat aphid biotypes in Colorado.