Ph I-induced transfer of leaf and stripe rust-resistance genes from Aegilops triuncialis and Ae. geniculata to bread wheat

Leaf and stripe rusts are severe foliar diseases of bread wheat. Recently, chromosomes 5M^g from the related species Aegilops geniculata that confers resistance to both leaf and stripe rust and 5U^t from Ae. triuncialis conferring resistance to leaf rust have been transferred to bread wheat in the form of disomic DS5M^g(5D) and DS5U^t(5A) chromosome substitution lines. The objective of this study was to shorten the alien segments in these lines using Ph ^I-mediated, induced homoeologous recombination. Putativerecombinants were evaluated for their rust resistance, and by genomic in situ hybridization and microsatellite analyses. One agronomically useful wheat-Ae. geniculata recombinant resistant to leaf and stripe rust was identified that had only a small terminal segment of the 5M^gL arm transferred to the long arm of an unidentified wheat chromosome. This germplasm can be used directly in breeding programs. Only one leaf rust-resistant wheat-Ae. triuncialis recombinant, which consists of most of the complete 5U^t chromosome with a small terminal segment derived from 5AS, was identified. This germplasm will need further chromosome engineering before it can be used in wheat improvement. This revised version was published online in August 2006 with corrections to the Cover Date.     

Utilisation of <Emphasis Type="Italic">Aegilops</Emphasis> (goatgrass) species to widen the genetic diversity of cultivated wheat

Wild Aegilops species related to cultivated wheat (Triticum spp.) possess numerous genes of agronomic interest and can be valuable sources of resistance to diseases, pests and extreme environmental factors. These genes can be incorporated into the wheat genome via intergeneric crossing, following, where necessary, the development of chromosome addition and substitution lines from the resulting hybrids. The transfer of a single segment from an alien chromosome can be achieved by translocations. The Aegilops (goatgrass) species, which are the most closely related to wheat, exhibit great genetic diversity, the exploitation of which has been the subject of experimentation for more than a century. The present paper gives a survey of the results achieved to date in the field of wheat–Aegilops hybridisation and gene transfer. The Aegilops genus consists of 11 diploid, 10 tetraploid and 2 hexaploid species. Of these 23 Aegilops species, most of the diploids (Ae. umbellulata Zhuk., Ae. mutica Boiss., Ae. bicornis (Forssk.) Jaub. & Spach, Ae. searsii Feldman & Kislev ex Hammer, Ae. caudata L., Ae. sharonensis Eig, Ae. speltoides Tausch, Ae. longissima Schweinf. & Muschl.) and several polyploids (Ae. ventricosa Tausch, Ae. peregrina (Hack. In J. Fraser) Marie & Weiller, Ae. geniculata Roth, Ae. kotschyi Boiss., Ae. biuncialis L.) have been used to develop wheat–Aegilops addition lines. Wheat–Aegilops substitution lines were developed using several species, including Ae. umbellulata, Ae. caudata, Ae. tauschii, Ae. speltoides, Ae. sharonensis, Ae. longissima and Ae. geniculata. Translocations carrying genes responsible for useful agronomic traits were developed with Ae. umbellulata, Ae. comosa, Ae. ventricosa, Ae. longissima, Ae. speltoides and Ae. geniculata. A large number of genes were transferred from Aegilops species to cultivated wheat, including those for resistance to leaf rust, stem rust, yellow rust and powdery mildew, and various pests (cereal cyst nematode, root knot nematode, Hessian fly, greenbug). Many molecular markers are linked to these resistance genes. The development of new molecular markers is also underway. There are still many untapped genetic resources in Aegilops species that could be used as resistance sources for plant breeding.     

New durum wheat with Hessian fly resistance from Triticum araraticum and T. carthlicum in Morocco

Hessian fly is an important pest of wheat on the North American continent and the temperate Mediterranean drylands. Yield losses caused by this insect in Morocco are the heaviest in the Mediterranean region and are estimated to be 36% on average. Genetic resistance to Mediterranean Hessian fly biotypes has not been found in durum wheat, although large numbers of durum accessions were screened. Genes for resistance were found in common wheat; some of which are transferable to durum. However, there is a need to broaden the genetic base for resistance in durum wheat. The objective of this work was to introgress resistance from selected Triticum araraticum and T. carthlicum accessions using multiple backcross methodology. The experimental recipient durum wheat included numerous adapted and high‐yielding lines. Testing for Hessian fly resistance under controlled conditions and field yield data showed that this programme yields Hessian fly‐resistant durum lines with good yields and adaptation.     

Screening relatives of wheat for snow mold resistance and freeing tolerance

Four hundred and eighty one accessions of species and subspecies of genera Triticum and Aegilops were evaluated for resistance to speckled snow mold caused by Typhula ishikariensis and for freezing tolerance. All Triticum and seven of Aegilops species were severely affected by snow mold. Among surviving Aegilops spp., only the Ae. cylindrica accessions exhibited resistance similar to that of the most resistant winter wheat cultivar, PI 173438. After repeated screening of accessions of Ae. cylindrica, 12 accessions were identified as having resistance similar to that of PI 173438; eleven of those were considered more freezing tolerant than PI 173438, but less than wheat cv. Valjevskaya, the freezing tolerant check. Accessions of Ae. cylindrica with snow mold resistance exceeding that of PI 173438, and with freezing tolerance, are currently being screened. This revised version was published online in August 2006 with corrections to the Cover Date.     

Wheat pre-breeding using wild progenitors

To facilitate the use of wheat wild relatives in conventional breedingprograms, a wheat pre-breeding activity started at ICARDA in 1994/1995season. Preliminary results of gene introgression from wild diploidprogenitors, Triticum urartu, T. baeoticum, Aegilops speltoides andAe. tauschii and tetraploid T. dicoccoides are described. Crosseswith wild diploid Triticum spp. yielded high variation in plant andspike morphology. Synthetic hexaploids were produced from crosses of alocal durum wheat landrace `Haurani' with two Ae. tauschiiaccessions. Both Ae. tauschii accessions carry hybrid necrosis allelesthat gave necrotic plant phenotypes in crosses with some bread wheats.Backcross progenies with agronomical desirable traits, i.e. high spikeproductivity, short plant stature, earliness, drought tolerance and highproductive tillering, were identified in crosses of durum wheat with wild Triticum spp. and in a cross of one of the hexaploid synthetics with alocally adapted bread wheat cv. `Cham 6'. Resistance to yellow rust wasfound in durum wheat crosses with the three wild Triticum spp. andAe. speltoides and leaf rust resistance was identified in crosses withT. baeoticum and Ae. speltoides. The results show that wheatimmediate progenitors may be a valuable and readily accessible source ofnew genetic diversity for wheat improvement.     

Expression of barley yellow dwarf virus and Russian wheat aphid resistance genes in and fertility of spring wheat × triticale hybrids and backcross lines

Summary The Barley Yellow Dwarf Virus disease (BYDV) and the Russian wheat aphid (RWA) Diuraphis noxia (Mordvilko) have caused significant losses to wheat and barley in several areas of the world. Important sources of resistance to both BYDV and RWA have been found in Triticale. Different generations of interspecific wheat x Triticale crosses were produced and the progenies were screened for BYDV and RWA tolerance. Plants with equal chromosome numbers showed different levels of fertility. A significant correlation was observed between pollen fertility and seed set in primary florets (r=0.57). In generaL, pollen fertility, seed set and the number of euploid plants (2n=42) increased from one generation to the next. The expression of BYDV tolerance varied from population to population. Additive effects were predominant in F₁ and some backcross populations. A dominant effect of rye tolerance genes was also observed in few populations. A monogenic trait or a quantitative (polygenic) character would not agree with the observed segregation patterns. The heritability of this oligogenic tolerance was quite different between populations and in many populations the tolerance genes were only partially expressed. Some transgressive segregation for tolerance and sensitivity was demonstrated. The genes controlling tolerance to RWA in Triticale lines, Muskox 658 and Nord Kivu were not expressed in advanced lines resistant to BYDV. This indicates that tolerance genes for BYDV and RWA in these lines are located on different chromosomes.     

The location and characteristics of novel sources of resistance to Bremia Lactucae Regel (downy mildew) in wild Lactuca L. Species

Summary Twenty-one accessions of 3 wild Lactuca species which could be hybridised with L. sativa, the cultivated lettuce, were inoculated at different stages of plant development with 3 multivirulent isolates of Bremia lactucae. Nineteen sources of resistance to B. lactucae, not attributable to the previously recognised resistance factors 1–11 were identified. Two lines of L. serriola showed similar resistance patterns as lines carrying R11. The resistance of some accession was incomplete particularly at the seedling stage and this phenomenon may be race specific.Tests on segregating F₂ populations of crosses between 2 different L. serriola accessions and L. sativa cultivars showed that the resistance in one line (LSE/18) appears to be inherited as a single dominant gene, which is sometimes incomplete in expression and allelic to either Dm6 or R7. The segregation patterns for resistance in PI 281876 did not give readily interpretable ratios.To assess the frequency of occurrence in B. lactucae populations of virulence factors to overcome this novel resistance, 11 of the novel sources of resistance were inoculated with numerous collections of the pathogen from the UK, Czechoslovakia and elsewhere and found to show a high level of resistance.     

Association of KASP markers with Hessian fly resistance in wheat of diverse origin

Hessian fly (Mayetiola destructor (Say)) is a major pest in wheat producing areas of United States, Canada, Europe, and North Africa but has not been found in Australia. Host plant resistance is thought to involve a similar strategy used against plant pathogen, and in this study, we sought to investigate whether disease-resistance gene markers can also be useful in selecting against Hessian fly attack. The genome-wide association study involved 251 wheat genotypes of diverse origin and 72 SNPs, selected on the basis of significant similarity of their associated contig sequences to putative disease resistance genes. A novel statistical approach for genome-wide scan was applied, which utilised genotype data scored as Null alleles in the mixed model, instead of deleting or treating them as missing alleles. The analyses identified four markers with significant associations at the 5% level, after applying the false discovery rate. These were located on chromosomes 4A, 4B, 5A and 7D, with the 5A locus mapping to the cluster of major genes that confer resistance to multiple Hessian fly biotypes. Amongst the diverse wheat accessions analysed, most of the susceptible phenotypes carried the A–G–C–C haplotype at the BS00064369, BS00007416, BS00077047 and HfrDrd_nt_143 loci, respectively. When heterozygotes were excluded, all the Australian wheat accessions carried this allelic combination. The combination of alleles conferring resistance depended on the origin of the wheat accessions, with ICARDA accessions carrying a preponderance of the C–C–CG–T. Of the 11 USA accessions used for this study, only Lola carried a favourable combination of alleles for resistance at these loci.     

Intra species variation in Atylosia scarabaeoides (L.) Benth., a wild relative of pigeonpea (Cajanus cajan (L.) Millsp.)

Summary Atylosia scarabaeoides (L.) Benth., a wild relative of pigeonpea, possesses several useful genes which can be utilized for pigeonpea improvement. In the present study, 33 accessions of A. scarabaeoides were evaluated at ICRISAT Center during the 1987 rainy season for variation in some useful traits to identify parents for inter-generic hybridization. A large variation was observed for leaf components, seed size, pod length, seeds/pod, days to flowering, seed protein, sulphur amino acids, resistance to cyst nematode, phytophthora blight, sterility mosaic, fusarium wilt, pod borer, pod fly, and pod wasp. Only four accessions were found to have more than 28% protein content. Methionine and cystine contents were marginally higher than in pigeonpea but the variation was not large enough to utilize them in the breeding program. In A. scarabaeoides. accessions resistant to fusarium wilt, phytophthora blight, sterility mosaic, and cyst nematode were detected. Compared to pigeonpea, the A. scarabaeoides accessions were less susceptible to lepidopteran borer and were immune to pod fly damage. Accessions ICPW 89 and ICPW 111 in short- (100–120 days), and ICPW 94 and ICPW 118 in medium-duration (140–180 days) were identified as potential parents for use in inter-generic hybridization.ICRISAT Journal Article No. 967     

Adaptation of wheat rusts to the wheat cultivars in former Czechoslovakia

Summary In former Czechoslovakia virulence of rusts attacking wheat was studied since the sixties. Since the same time genes for resistance in the registered cultivars were identified. The role of Berberis and Thalictrum as alternate hosts for stem rust and leaf rust, respectively, was investigated as well. Determined changes of virulence in rust populations could only partially be ascribed to changes of resistance genes in the grown cultivars. Unnecessary genes for virulence had no negative effect on the fitness of the pathogen. All tested samples of aeciospores from barberries attacked rye, not wheat. None of Thalictrum species occurring in the Czech and Slovak Republics was found to host wheat leaf rust. However, the sexual stage of wheat stem rust and wheat leaf rust could be induced on Berberis vulgaris and Thalictrum speciosissimum, respectively. General epidemiological conclusions are drawn from the results and experience of the last 35 years.     

Population genetic structure of the clonal plant Zingiber zerumbet (L.) Smith (Zingiberaceae), a wild relative of cultivated ginger, and its response to Pythium aphanidermatum

Zingiber zerumbet (L) Smith, a wild clonal species related to the cultivated ginger (Zingiber officinale Roscoe), is a potential resistance donor for soft rot disease in ginger caused by Pythium aphanidermatum (Edson) Fitzp. In this study we evaluated the genetic diversity and P. aphanidermatum resistance of 74 Z. zerumbet accessions belonging to 15 populations from eight districts in Kerala state, India. The disease index (DI) of the accessions varied from 0% to 72.24% and the accessions could be separated into six frequency classes according to their DI values. More than 65% of the accessions had a DI < 20%. Eight accessions were found to be immune to the infection. The relative frequency of resistant accessions was higher in the central and northern regions of Kerala. Amplified fragment length polymorphism (AFLP) analysis of Z. zerumbet accessions using five primer combinations yielded 215 bands in total, of which 175 (81.4%) were polymorphic. Nei’s genetic diversity (h) of 0.2738 and Shannon information index (I) of 0.4012 revealed a high genetic diversity in Z. zerumbet unexpected for a clonal species. In the UPGMA dendrogram, accessions were clustered mostly according to their geographical origin and no clear correspondence was observed between the clustering pattern of accessions and their responses to Pythium aphanidermatum. The study revealed high genetic diversity and variability for pathogen resistance among Z. zerumbet accessions and confirmed the value of Z. zerumbet as a potential donor for soft rot resistance for the genetic improvement of ginger.     

Molecular mapping determines that Hessian fly resistance gene H9 is located on chromosome 1A of wheat

Hessian fly [Mayetiola destructor (Say)] is one of the major insect pests of wheat (Triticum aestivum L.) worldwide. Hessian fly resistance gene H9 was previously reported to condition resistance to Hessian fly biotype L that is prevalent in many wheat‐growing areas of eastern USA and an RAPD marker, OPO05₁₀₀₀, linked to H9 in wheat was developed using wheat near‐isogenic lines (NILs). However, marker‐assisted selection (MAS) with RAPD markers is not always feasible. One of the objectives in this study was to convert an RAPD marker linked to the gene H9 into a sequence characterized amplified region (SCAR) marker to facilitate MAS and to map H9 in the wheat genome. The RAPD fragment from OPO05₁₀₀₀ was cloned, sequenced, and converted into a SCAR marker SOPO05₉₀₉, whose linkage relationship with H9 was subsequently confirmed in two F₂ populations segregating for H9. Linkage analysis identified one sequence tagged site (STS) marker, STS‐Pm3, and the eight microsatellite markers Xbarc263, Xcfa2153, Xpsp2999, Xgwm136, Xgdm33, Xcnl76, Xcnl117 and Xwmc24 near the H9 locus on the distal region of the short arm of chromosome 1A, contrary to the previously reported location of H9 on chromosome 5A. Locus Xbarc263 was 1.2 cM distal to H9, which itself was 1.7 cM proximal to loci Xcfa2153, Xpsp2999 and Xgwm136. The loci Xgwm136, Xcfa2153 and SOPO05₉₀₉ were shown to be specific to H9 and not diagnostic to several other Hessian fly resistance genes, and therefore should be useful for pyramiding H9 with other Hessian fly resistance genes in a single genotype.     

Potential new genes for resistance to <Emphasis Type="Italic">Mycosphaerella graminicola</Emphasis> identified in <Emphasis Type="Italic">Triticum aestivum</Emphasis> × <Emphasis Type="Italic">Lophopyrum elongatum</Emphasis> disomic substitution lines

Lophopyrum species carry many desirable agronomic traits, including disease resistance, which can be transferred to wheat by interspecific hybridization. To identify potentially new genes for disease and insect resistance carried by individual Lophopyrum chromosomes, 19 of 21 possible wheat cultivar Chinese Spring × Lophopyrum elongatum disomic substitution lines were tested for resistance to barley yellow dwarf virus (BYDV), cereal yellow dwarf virus (CYDV), the Hessian fly Mayetiola destructor, and the fungal pathogens Blumeria graminis and Mycosphaerella graminicola (asexual stage: Septoria tritici). Low resistance to BYDV occurred in some of the disomic substitution lines, but viral titers were significantly higher than those of two Lophopyrum species tested. This suggested that genes on more than one Lophopyrum chromosome are required for complete resistance to this virus. A potentially new gene for resistance to CYDV was detected on wheatgrass chromosome 3E. All of the substitution lines were susceptible to Mayetiola destructor and one strain of B. graminis. Disomic substitution lines containing wheatgrass chromosomes 1E and 6E were significantly more resistant to M. graminicola compared to Chinese Spring. Although neither chromosome by itself conferred resistance as high as that in the wheatgrass parent, they do appear to contain potentially new genes for resistance against this pathogen that could be useful for future plant-improvement programs.     

The exploration of genetic resources of Portuguese cabbage and kale for resistance to several Brassica diseases

Summary Twenty three accessions of nine Portuguese cabbage and kale land races from different geographic origins were tested at the seedling stage for resistance to several important brassica diseases. Resistance to downy mildew (Peronospora parasitica), expressed as necrosis of the cotyledon mesophyll, was found in all the accessions. Type A resistance to cabbage yellows (Fusarium oxysporum f. sp. conglutinans race 1) was present in most of the landraces. Resistance to clubroot (Plasmodiophora brassicae race 6) was found in one accession of the Portuguese tree kale. High resistance to blackleg (Leptosphaeria maculans) and white rust (Albuco candida) was not detected, although several accessions showed 20 to 30% of plants with intermediate expression of resistance. All Portuguese cole accessions were susceptible to blackrot (Xanthomonas campestris pv. campestris).     

Tan-Spot Resistance Screening of Aegilops Species

Two hundred and twelve accessions of 8 diploid and 10 polyploid species of Aegilops were evaluated for resistance to tan-spot disease of wheat, caused by Pyrenophora tritici-repentis (Died.) Drechs., under greenhouse conditions. One or more accessions of Ae. bicornis, Ae. biuncialis, Ae. Crassa, Ae. columnaris, Ae. cylindrica, Ae. speltoides, Ae. squarrosa. Ae. triaristata. Ae. triuncialis, and Ae. Ovata showed resistance following a 24-hour post-inoculation wet period. With a 36-hour wet period, diploids became only slightly or moderately susceptible and resistant polyploids became susceptible. A 48-hour wet period resulted in still greater susceptibility of both diploid and polyploid species.     

Identification of Resistance Genes Against Powdery Mildew in Four Accessions of Hordeum Vulgare SSP. Spontaneum

Summary Four newly detected accessions of wild barley (Hordeum vulgare ssp. spontaneum) resistant to powdery mildew caused by Blumeria graminis f. sp. hordei were studied with the aim of finding the number of genes/loci conferring the resistance of individual accessions, the type of inheritance of the genes and their relationships to the Mla locus. F₂ populations after crosses between the winter variety ‘Tiffany’ and four wild barley accessions and use of microsatellite DNA markers were focused on the identification of individual resistance genes/loci by means of their chromosomal locations. In PI466495, one locus conferring powdery mildew resistance was identified in highly significant linkage with the marker Bmac0213. This location is consistent with the known locus Mla on chromosome 1HS. In the other three accessions the resistance was determined by two independent loci. In PI466197, PI466297 and PI466461, one locus was identified on chromosome 1HS and three new loci were revealed on chromosomes 2HS (highly significant linkage with Bmac0134), 7HS (highly significant linkage with Bmag0021) and 7HL (significant linkage with EBmac0755). Our prospective aim is identification of further linked DNA markers and the exact location of the resistance genes on the barley chromosomes.     

Sources of resistance to diseases of sugar beet in related Beta germplasm: II. Soil-borne diseases

Between 580 and 700 accessions of related cultivated and wild species of the genus Beta were assessed for resistance to four soil-borne diseases of sugar beet: two seedling damping-off diseases caused by the fungi Aphanomyces cochlioides and Pythium ultimum and two diseases of more mature plants, Rhizoctonia root and crown rot, caused by the fungus R. solani, and Rhizomania, caused by Beet necrotic yellow vein virus (BNYVV), a furovirus transmitted by the plasmodiophorid Polymyxa betae. Analysis of resistance data (assessed on an international standardised 1–9 scale of Resistance Scores) indicated that the highest levels of resistance ({RS} 2) to A. cochlioides and P. ultimum were to be found amongst accessions of the more distantly related sections Corollinae (93% of accessions tested) and Procumbentes (10%), respectively; although useful levels could also be found in the more closely related, and sexually compatible, section Beta (1–6%). Resistance to Rhizoctonia was also found in section Beta (5–7%), depending on whether field or glasshouse tests were used, but there was little evidence of generally high levels of resistance to Rhizomania among accessions of this section. None of the accessions of sections Corollinae and Procumbentes exhibited any notable resistance to Rhizoctonia. However, all sections Procumbentes and some sections Corollinae (4%) accessions were highly resistant to Rhizomania. Individuals with high levels of resistance to Rhizomania were identified from within some section Beta and Corollinae accessions, in which there was evidence of segregation.     

Inheritance of seed resistance to bruchids in cultivated mungbean (Vigna radiata, L. Wilczek)

Bruchid beetles or seed weevils are the most devastating stored pests of grain legumes causing considerable loss to mungbean (Vigna radiata (L.) Wilczek). Breeding for bruchid resistance is a major goal in mungbean improvement. Few sources of resistance in cultivated genepool were identified and characterized, however, there has been no study on the genetic control of the resistance. In this study, we investigated the inheritance of seed resistance to Callosobruchus chinensis (L.) and C. maculatus (F.) in two landrace mungbean accessions, V2709BG and V2802BG. The F₁, F₂ and BC generations were developed from crosses between the resistant and susceptible accessions and evaluated for resistance to the insects. It was found that resistance to bruchids in seeds is controlled by maternal plant genotype. All F₁ plants derived from both direct and reciprocal crosses exhibited resistance to the bruchids. Segregation pattern of reaction to the beetles in the F₂ and backcross populations showed that the resistance is controlled by a major gene, with resistance is dominant at varying degrees of expressivity. Although the presence of modifiers was also observed. The gene is likely the same locus in both V2709BG and V2802BG. The resistant gene is considered very useful in breeding for seed resistance to bruchids in mungbean.     

Resistance in spelt wheat to yellow rust

Summary Seven spelt wheat accessions of different origin were hybridized with the susceptible bread wheat cultivar Taichung 29 in order to study the genetics of their resistance to yellow rust (Puccinia striiformis Westend. f. sp. tritici). One Iranian and five European accessions were found to carry Yr5 of Triticum aestivum ssp. spelta var. album, whereas a factor for resistance in the Iranian accession 415 was confirmed to be genetically distinct from Yr5. The alleles for resistance in each of the accessions studied showed a monogenic dominant mode of inheritance. Twenty-eight spelt wheat accessions, including those studied for their resistance to yellow rust, were subjected to polyacrylamide-gel-electrophoresis to study variation for gliadin storage protein patterns. Thirteen distinct patterns were revealed, implying the presence of duplicates within the studied spelt wheat collection.     

Evaluation of French Brassica oleracea landraces for resistance to Plasmodiophora brassicae

A total of 240 kale, 38 cabbage and 126 winter cauliflower French landraces from the B. oleracea genepool of INRA were assessed for resistance to clubroot caused by Plasmodiophora brassicaeWoron. Two French isolates of the pathogen (K and SJ) were used in the experiments under controlled conditions. The reaction of the 126 cauliflower accessions to naturally occurring clubroot was also evaluated in field trials. Kales exhibited considerable variation for expression of disease resistance and high levels of resistance were found in several accessions. In this group, single resistant plants were observed in most of the morphological types and from quite different geographical origins. Cabbage accessions were moderately to highly susceptible to both isolates. All cauliflower populations proved to be highly susceptible to K isolate and moderately susceptible to SJ isolate. In field trials, cauliflowers were also severely infected. Two lines selected from a resistant kale population were highly resistant against a large range of pathotypes of the pathogen. These lines presented a sufficient level of resistance to be directly useful in the breeding program in order to develop cauliflower and broccoli hybrids resistant to clubroot. This revised version was published online in July 2006 with corrections to the Cover Date.