Patterns of resistance to three cereal aphids among wheats in the genus Triticum (Poaceae)

Forty-one accessions of wild and cultivated wheats belonging to 19 Triticum species were tested in the field for resistance to three species of aphids, Rhopalosiphum padi Linnaeus, Sitobion avenae Fabricius and Schizaphis graminum Rondani. Antibiotic resistance was estimated by the increase in biomass of aphids over 21 days on adult plants. Overall resistance was estimated by the plant biomass lost due to aphid infestation. All three species of aphids survived and reproduced on all wheats, and reduced spike biomass compared to uninfested controls. The level of antibiosis varied among wheat species and among accessions, with accessions from three, five and one species showing antibiosis to R. padi, S. avenae and S. graminum, respectively. Overall resistance to the three aphid species was observed in five to seven accessions per aphid species. Resistance was usually specific to one aphid species. The frequency of accessions with antibiosis or overall resistance was associated with the ploidy level of the plant species. Except for overall resistance to R. padi, resistance was highest for diploid species and lowest for hexaploid species. No consistent relationship between resistance and level of domestication was detected. Accessions of the wild wheats, Triticum boeoticum Bois, Triticum tauschii (Coss.) Schmal. and Triticum araraticum Jakubz. exhibited high levels of resistance to aphids, as did Triticum monococcum L. which is derived from T. boeoticum. Nevertheless, individual susceptible or resistant accessions occurred at all levels within the evolutionary tree of wheat.     

Tolerance of antibiotic and susceptible cereal seedlings to the aphids Metopolophium dirhodum and Rhopalosiphum padi

Some cereal seedlings exhibit antibiotic and antixenotic resistance to the aphids Metopolophium dirhodum (Walker) and Rhopalosiphum padi (L.), because the seedlings contain hydroxamic acids or gramine. The association between tolerance to aphids and aphid antibiosis was investigated for three cereals, Dollarbird wheat Vulcan wheat and Yagan barley. The dry biomass gained by the aphids and the simultaneous reduction in the biomass of the plants (biomass conversion ratio) quantified tolerance. Biomass production and the density dependence of biomass production by the aphids quantified antibiosis more effectively than fecundity. Vulcan wheat, which has more hydroxamic acid than Dollarbird wheat showed the highest level of antibiosis, and the barley was not antibiotic for either aphid. The biomass conversion ratio was a constant; the biomass of an infested plant was reduced by 3 mg for each mg of aphid biomass gained, regardless of aphid species, plant cultivar, or aphid density. The three plants showed no differential tolerance to the aphids, and therefore tolerance is not associated with antibiosis in this case.     

Laboratory assessments of resistance to the aphids Sitobion avenae and Metopolophium dirhodum in three Triticum species and two modern wheat cultivars

Three ancient varieties of wheat and two spring-sown modern varieties were screened in the laboratory to assess their resistance to the cereal aphids Sitobion avenae and Metopolophium dirhodum. Resistance was measured in terms of non-preference and antibiosis on plants at three growth stages. The ancient variety Einkorn was the most resistant in terms of both non-preference and antibiosis to both species of aphids at all growth stages examined. The ancient variety Emmer and the modern Sicco exhibited some resistant properties, whereas the ancient Spelt and modern Timmo were relatively susceptible to aphid attack.     

Sources of variation in the interaction between three cereal aphids (Hemiptera: Aphididae) and wheat (Poaceae)

The relative contributions of host plant, herbivore species and clone to variation in the interaction between cereal aphids and wheat were investigated using five clones each of three species, Rhopalosiphum padi (Linnaeus), Sitobion avenae (Fabricius) and Schizaphis graminum (Rondani), on seedlings of two cultivars of Triticum aestivum L. and one cultivar of Triticum durum Desf. More individuals and biomass of R. padi than of the other two species were produced on seedlings. The three wheat cultivars lost similar amounts of biomass as a result of infestation by aphids, with the amount lost depending on aphid species: S. avenae caused the lowest loss in biomass. Variation in aphid biomass production was due mostly to differences among aphid species (70%), less to the interaction between wheat type and aphid species (7%), and least to aphid clone (1%). The specific impact of the aphids on the plants ranged from 1.7 to 3.7 mg of plant biomass lost per mg of aphid biomass gained, being lowest for R. padi and highest for S. graminum. Variation in plant biomass lost to herbivory was due mostly to unknown sources of error (95%), probably phenotypic differences among individual seedlings, with 3% due to aphid species and none attributable to aphid clone. For these aphid-plant interactions, differences among aphid clones within species contributed little to variation in aphid and plant productivity; therefore, a small sample of clones was adequate for quantifying the interactions. Furthermore, one clone of each species maintained in the laboratory for about 200 parthenogenetic generations was indistinguishable from clones collected recently from the field.     

Resistance to Rhopalosiphum padi (Homoptera: Aphididae) in three triticale accessions

Experiments were conducted to identify and characterize host plant resistance to bird cherry-oat aphid, Rhopalosiphum padi (L.), in various wheat and wheat-grass hybrids. Initial tests screened for resistance to R. padi among 12 grass accessions (eight wheat [Triticum aestivum L.], three triticale [XTriticosecale Wittmack], and 1XElytricum [Elytrigia elongata [Host] Nevski x Triticum aestivum hybrid]). R. padi had less population growth on triticale accessions '8TA5L' (PI 611760) and 'Stniism 3' (PI 386156) than on other accessions, but nymphiposition by R. padi did not differ among the 12 accessions. Follow-up experiments were conducted to characterize antibiosis, antixenosis, and tolerance to R. padi in three wheat and three triticale accessions. In antibiosis experiments, Stniism 3 and triticale 'H7089-52' (PI 611811) prolonged time to reproduction by R. padi compared with that on wheat accessions 'Arapahoe' (PI 518591), 'KS92WGRC24' (PI 574479), and 'MV4' (PI 435095), whereas time to reproduction on 8TA5L was intermediate and did not differ from that on the other five accessions. Also, R. padi produced fewest progeny on Stniism 3, and fewer progeny on 8TA5L than on H7089-52, Arapahoe, KS92WGRC24, and MV4. Stniism 3 showed antixenosis, because fewer winged R. padi selected Stniism 3 than Arapahoe, H7089-52, or MV4 in choice tests. In tolerance experiments, a 300 aphid-day infestation of R. padi limited shoot length of Arapahoe and KS92WGRC24 plants. Shoot lengths did not differ between infested and noninfested seedlings of MV4, 8TA5L, H7089-52, and Stniism 3, indicating tolerance to R. padi in these accessions. Triticale accessions 8TA5L, H7089-52, and Stniism 3 and MV4 wheat may be meaningful sources of R. padi resistance for small-grain breeding programs, and Stniism 3 may be particularly valuable, given reports of its additional resistance to the Russian wheat aphid, Diuraphis noxia (Mordvilko).     

Interactions of Russian wheat aphid, a hymenopterous parasitoid and resistant and susceptible slender wheatgrasses

Interactions among three trophic levels of resistant and susceptible slenderwheat grasses, Elymus trachycaulum (Link) Goule ex Shinners ex. H.F. Lewis, Russian wheat aphid, Diuraphis noxia (Mordvilko), and a hymenopterous parasitoid were studied in the laboratory and greenhouse. These relationships were compared with a commercial susceptible wheat Triticum aestivum L. variety. Aphids reared on the resistant entries showed significantly lower weights and numbers. Significant reduction of parasitoid mummy weight and adult size was positively correlated with the effects on the aphids. Resistant entries also induced a longer prereproductive period for both the aphids and parasitoids. Numbers of aphids and aphid damage were significantly modified by the addition of parasitoids. Parasitism was higher on plants that did not have leaf rolling. These findings may indicate that antibiosis resistance studied here is not the most desirable because it decreases natural enemy vitality.     

Expression of high zinc efficiency of Aegilops tauschii and Triticum monococcum in synthetic hexaploid wheats

The effect of varied zinc (Zn) supply on shoot and root dry matter production, severity of Zn deficiency symptoms and Zn tissue concentrations was studied in two Triticum turgidum (BBAA) genotypes and three synthetic hexaploid wheat genotypes by growing plants in a Zn-deficient calcareous soil under greenhouse conditions with (+Zn=5 mg kg^-1 soil) and without (−Zn) Zn supply. Two synthetic wheats (BBAADD) were derived from two different Aegilops tauschii (DD) accessions using same Triticum turgidum (BBAA), while one synthetic wheat (BBAAAA) was derived from Triticum turgidum (BBAA) and Triticum monococcum (AA). Visible symptoms of Zn deficiency, such as occurrence of necrotic patches on leaves and reduction in shoot elongation developed more rapidly and severely in tetraploid wheats than in synthetic hexaploid wheats. Correspondingly, decreases in shoot and root dry matter production due to Zn deficiency were higher in tetraploid wheats than in synthetic hexaploid wheats. Transfer of the DD genome from Aegilops tauschii or the AA genome from Triticum monococcum to tetraploid wheat greatly improved root and particularly shoot growth under Zn-deficient, but not under Zn-sufficient conditions. Better growth and lesser Zn deficiency symptoms in synthetic hexaploid wheats than in tetraploid wheats were not accompanied by increases in Zn concentration per unit dry weight, but related more to the total amount of Zn per shoot, especially in the case of synthetic wheats derived from Aegilops tauschii. This result indicates higher Zn uptake capacity of synthetic wheats. The results demonstrated that the genes for high Zn efficiency from Aegilops tauschii (DD) and Triticum monococcum (AA) are expressed in the synthetic hexaploid wheats. These wheat relatives can be used as valuable sources of genes for improvement of Zn efficiency in wheat. This revised version was published online in June 2006 with corrections to the Cover Date.     

Phenotypic mechanisms of host resistance against greenbug (Homoptera: Aphididae) revealed by near isogenic lines of wheat

Interactions between biotype E greenbug, Schizaphis graminum (Rondani), and wheat, Triticum aestivum L., were investigated using resistant and susceptible near isogenic lines of the greenbug resistance gene Gb3. In an antixenosis test, the greenbugs preferred susceptible plants to resistant ones when free choice of hosts was allowed. Aphid feeding resulted in quick and severe damage to susceptible plants, which seemed to follow a general pattern spatially and was affected by the position where the greenbugs were initially placed. Symptom of damage in resistant plants resembled senescence. Within-plant distribution of aphids after infestation was clearly different between the two genotypes. Significantly more greenbugs fed on the first (oldest) leaf than on the stem in resistant plants, but this preference was reversed in the susceptible one. After reaching its peak, aphid population on the susceptible plants dropped quickly. All susceptible plants were dead in 10-14 d after infestation due to greenbug feeding. Aphid population dynamics on resistant plants exhibited a multipeak curve. After the first peak, the greenbug population declined slowly. More than 70% of resistant plants were killed 47 d after infestation. Performance of both biotype E and I greenbugs on several Gb3-related wheat germplasm lines were also examined. It seems that the preference-on-stem that was characteristic of biotype E greenbugs on the susceptible plants was aphid biotype- and host genotype-dependent. Results from this study suggested that antixenosis, antibiosis, and tolerance in the resistant plants of wheat might all contribute to resistance against greenbug feeding.     

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.     

Genetic control of the wheat Triticum monococcum L. resistance to powdery mildew

Using hybrid analysis and test-clone method, 102 accessions of Triticum monococcum L. from the collection of the Vavilov All-Russia Institute of Plant Industry have been studied. This species of wheat has been found to by considerably polymorphic with respect to the resistance to the fungus Erysiphe graminis DC. f. sp. tritici Marchal. causing powdery mildew. The resistance of most accessions to the fungus population and clones is determined by dominant genes. In rare cases, the resistance was determined by recessive genes or one, two, or three oligogenes. A group of einkorn wheat accessions has been found in which the resistance to powdery mildew was determined by the same dominant factor or different but closely linked ones. Recessive resistance genes of T. monococcum differ from the recessive gene pm5 determining the resistance of T. aestivum plants. The genome of T. monococcum contains various genes of resistance to powdery mildew and is a potential source of effective genes to be used when selecting cultivated species of wheat for immunity.     

Quantification of genetic relationships among A genomes of wheats.

The genetic relationships of A genomes of Triticum urartu (Au) and Triticum monococcum (Am) in polyploid wheats are explored and quantified by AFLP fingerprinting. Forty-one accessions of A-genome diploid wheats, 3 of AG-genome wheats, 19 of AB-genome wheats, 15 of ABD-genome wheats, and 1 of the D-genome donor Ae. tauschii have been analysed. Based on 7 AFLP primer combinations, 423 bands were identified as potentially A genome specific. The bands were reduced to 239 by eliminating those present in autoradiograms of Ae. tauschii, bands interpreted as common to all wheat genomes. Neighbour-joining analysis separates T. urartu from T. monococcum. Triticum urartu has the closest relationship to polyploid wheats. Triticum turgidum subsp. dicoccum and T. turgidum subsp. durum lines are included in tightly linked clusters. The hexaploid spelts occupy positions in the phylogenetic tree intermediate between bread wheats and T. turgidum. The AG-genome accessions cluster in a position quite distant from both diploid and other polyploid wheats. The estimates of similarity between A genomes of diploid and polyploid wheats indicate that, compared with Am, Au has around 20% higher similarity to the genomes of polyploid wheats. Triticum timo pheevii AG genome is molecularly equidistant from those of Au and Am wheats.     

我国麦蚜抗药性及研究现状

麦蚜是危害我国小麦Triticum aestivum L.生产的主要害虫,具有分布广、数量大、繁殖力强以及远距离迁飞等特点,不仅直接吸食小麦汁液,还传播多种植物病毒,每年造成小麦减产10%~30%。目前对麦蚜的防控主要以化学防治为主,但由于化学杀虫剂长期或不合理的使用,多地麦蚜对常用杀虫剂产生了不同程度的抗性。本文从麦蚜抗药性测定方法、抗性水平及交互抗性、代谢和靶标抗性机制、以及麦蚜抗药性综合治理等方面进行了综述,以期为麦蚜的防治及杀虫剂的持续合理使用提供理论参考和依据。     

Categories and inheritance of resistance to Russian wheat aphid (Homoptera: Aphididae) biotype 2 in a selection from wheat cereal introduction 2401

The Russian wheat aphid, Diuraphis noxia (Kurdjumov) (Homoptera: Aphididae), is one of the most devastating insect pests of wheat (Triticum spp.) and barley (Hordeum spp.) in the world. Yield losses and control costs are valued at several hundred million dollars each year. The use of D. noxia-resistant cultivars is an ecologically, economically, and biologically sound method of managing this pest. Several D. noxia resistance (Dn) genes from wheat have been used to develop cultivars resistant to D. noxia. However, a new U.S. D. noxia biotype (biotype 2) in Colorado is virulent to all known Dn genes except the Dn7 gene from rye (Secale spp.). Hence, there is an immediate need to identify and characterize unique sources of D. noxia resistance to biotypes. In this article, we report resistance to D. noxia biotype 2, identified in a selection from wheat cereal introduction (CItr) 2401, that is controlled by two dominant genes. CItr2401 has a strong antibiosis effect that is exhibited as a reduced intrinsic rate of increase of D. noxia biotype 2. CItr2401 plants also exhibit tolerance to leaf rolling and chlorosis. No antixenosis was detected in CItr2401.     

Development and reproductive potential of the cereal aphid Sitobion avenae on resistant wheat lines (Triticum monococcum)

The effect of wheat resistance in lines of Triticum monococcum L., on the reproductive performances of the cereal aphid (Sitobion avenae F.) was investigated. Aphids were reared from birth to adult moult either on resistant or susceptible wheat lines, and transferred as apterae to both host genotypes. The influence of these transfers on the subsequent adult weight, gonad status and reproductive performances was evaluated. Aphids transferred from resistant to susceptible plants proved able to compensate for their poor nymphal growth, mainly through additional embryo growth and an increase in the number of matured embryos within the first 10 days of their adult life. Most aphids transferred from susceptible to resistant plants died within the first week following the transfer. Their most advanced embryos matured and were born, but subsequent embryo growth was quickly reduced. The reproductive strategies adopted by S. avenae when facing plant resistance, and the hypothesis of a resistant mechanism based on a poor nutritional state of the resistant plants are discussed.     

Winter wheat (Triticum aestivum) response to Barley yellow dwarf virus at various nitrogen application rates in the presence and absence of its aphid vector,Rhopalosiphum padi

Barley yellow dwarf (BYD) is one of the most common diseases of cereal crops, caused by the phloem‐limited, cereal aphid‐borne Barley yellow dwarf virus (BYDV) (Luteoviridae). Delayed planting and controlling aphid vector numbers with insecticides have been the primary approaches to manage BYD. There is limited research on nitrogen (N) application effects on plant growth, N status, and water use in the BYDV pathosystem in the absence of aphid control. Such information will be essential in developing a post‐infection management plan for BYDV‐infected cereals. Through a greenhouse study, we assessed whether manipulation of N supply to BYDV‐infected winter wheat, Triticum aestivum L. (Poaceae), in the presence or absence of the aphid vector Rhopalosiphum padi L. (Hemiptera: Aphididae), could improve N and/or water uptake, and subsequently promote plant growth. Similar responses of shoot biomass and of water and N use efficiencies to various N application rates were observed in both BYDV‐infected and non‐infected plants, suggesting that winter wheat plants with only BYDV infection may be capable of outgrowing infection by the virus. Plants, which simultaneously hosted aphids and BYDV, suffered more severe symptoms and possessed higher virus loads than those infected with BYDV only. Moreover, in plants hosting both BYDV and aphids, aphid pressure was positively associated with N concentration within plant tissue, suggesting that N application and N concentration within foliar tissue may alter BYDV replication indirectly through their influence on aphid reproduction. Even though shoot biomass, tissue N concentration, and water use efficiency increased in response to increased N application, decision‐making on N fertilization to plants hosting both BYDV and aphids should take into consideration the potential of aphid outbreak and/or the possibility of reduced plant resilience to environmental stresses due to decreased root growth.     

Characterization of greenbug (Homoptera: Aphididae) resistance in synthetic hexaploid wheats

Twelve greenbug (Schizaphis graminum (Rondani)) biotype E-resistant synthetic hexaploid wheats synthesized by crossing Triticum dicoccum Schrank. and Aegilops tauschii (Coss.) Schmal. were evaluated for the three known insect resistance categories, including antibiosis, anti-xenosis, and tolerance. Different methods were evaluated for calculating antibiosis and tolerance. Calculating intrinsic rate of population increase and measuring leaf chlorophyll content with a SPAD chlorophyll meter proved to be time- and labor-efficient for antibiosis and tolerance determination, respectively. The resistance in all synthetic hexaploids proved to be the result of a combination of antibiosis, antixenosis, and tolerance, which makes them valuable sources of greenbug resistance. To assist plant breeders in selecting the best germplasm for greenbug resistance, a plant resistance index was created that revealed differences among the synthetic hexaploid wheats.     

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.     

The potential of hydroxamic acids in tetraploid and hexaploid wheat varieties as resistance factors against the bird‐cherry oat aphid,Rhopalosiphum padi

The potential for exploiting natural wheat resistance to control the cereal aphid Rhopalosiphum padi, the most important aphid pest of small grain cereals in the UK, was investigated as an alternative approach to the use of insecticides. The investigation focussed on a group of secondary metabolites, the hydroxamic acids or benzoxazinones, present naturally as glucosides, but which hydrolyse on tissue damage to give biologically active aglycones, e.g. 2,4‐dihydroxy‐7‐methoxy‐1,4‐benzoxazin‐3‐one (DIMBOA) which are associated with natural plant defence. These can be important for resistance against insects, fungi, bacteria and nematodes for a range of cultivated monocotyledonous plants and could ultimately be combined with other defence mechanisms to provide a general approach to cereal aphid control. Levels of hydroxamic acids, particularly DIMBOA‐glucoside, were determined in hexaploid (Triticum aestivum) and tetraploid (Triticum durum) wheat varieties and differences were found between species and varieties. The effect of feeding by R. padi on the level of hydroxamic acids in the leaf tissue was also investigated. Thus, after 24 h of aphid feeding, as an apparently localised hydrolytic defence reaction in the leaf, levels of DIMBOA‐glucoside decreased noticeably. When aphids were fed on sucrose solution containing low doses of DIMBOA there was a significant mortality compared to the sucrose control. However, the levels of and variation in hydroxamic acids in the wheat varieties investigated were insufficient for significant differences in aphid behaviour and development.     

Effect of wheat resistance, the parasitoid Aphidius rhopalosiphi, and the entomopathogenic fungus Pandora neoaphidis, on population dynamics of the cereal aphid Sitobion avenae

The influence of wheat (Triticum aestivumL.) resistance, the parasitoid Aphidius rhopalosiphiDe Stephani-Perez (Hymenoptera: Braconidae) and the entomopathogenic fungus Pandora neoaphidis(Remaudière et Hennebert) Humber (Zygomycetes: Entomophthorales) on the density and population growth rate of the cereal aphid Sitobion avenae(F.) (Hemiptera: Aphididae) was studied under laboratory conditions. Partial wheat resistance was based on hydroxamic acids, a family of secondary metabolites characteristic of several cultivated cereals. The partial resistance of wheat cultivar Naofén, the action of the parasitoid and the joint action of the parasitoid and fungus, reduced aphid density. The lowest aphid densities were obtained with the combination of the parasitoid and the fungus, but wheat resistance under these circumstances did not improve aphid control. Significant reductions of population growth rate (PGR) of aphids were obtained with the joint action of wheat resistance and natural enemies. In particular, the combined effects of parasitoids and fungi showed significantly lower PGR than the control without natural enemies in both wheat cultivars. Our results support the hypothesis that wheat resistance and the utilization of biological control agents could be complementary strategies in an integrated pest management program against cereal aphids.