MINISTRY OF AGRICULTURE, FISHERIES AND FOOD CSG 15 Research and Development Final Project Report (Not to be used for LINK projects)

Two hard copies of this form should be returned to: Research Policy and International Division, Final Reports Unit MAFF, Area 6/01 1A Page Street, London SW1P 4PQ An electronic version should be e-mailed to [email protected]

Project title Genetic improvement of resistance to pests and diseases, tree habit and fruit quality in and pear

MAFF project code HH1016STF

Contractor organisation Horticulture Research International and location East Malling West Malling, Kent ME19 6BJ

Total MAFF project costs £ 833,256

Project start date 01/04/97 Project end date 31/03/00

Executive summary (maximum 2 sides A4)

The project has the following objectives:

· to identify in apple and pear sources of resistance to economically important pests and diseases for which no breeding lines have yet been developed · to elucidate in apple inheritance of other agronomically describable trees and fruit characters · to raise selections of apple and pear with desirable combinations of commercial characters · to identify genotypically good parents in apple and pear · to resolve the incompatibility genotype of ‘Meridian’ and of 50 other apple .

Some 3,500 apple selections of normal habit and 300 pear selections were planted out for assessment by the Apple and Pear Breeding Club, along with 700 columnar selections. Some 34,000 seedlings were screened for resistance to scab in the glasshouse, and the survivors planted in an unsprayed seedling nursery, together with 3,500 seedlings combining mildew resistance genes. About 25,000 seedlings not carrying disease resistance were planted in the sprayed nursery. Approximately 45,000 apple flowers and 10,000 pear flowers were hand-pollinated to raise progenies for genetic studies or for screening.

The progeny ‘Elise Rathke’ (weeping) x ‘’ segregated for weeping habit, indicating this character is controlled by a gene. Progenies were raised of two ‘low ethylene’ apple selections. The pears ‘Comice’ (quince compatible) and ‘Williams’ (quince incompatible) were intercrossed.

CSG 15 (Rev. 12/99) 1 Project Genetic improvement of resistance to pests and diseases, MAFF HH1016STF title tree habit and fruit quality in apple and pear project code

Segregation for pink flesh in the progeny ‘Falstaff’ x ‘Pink Pearl’ indicated this novel character is controlled by a dominant gene. However, the progeny ‘Telamon’ x ‘Pink Pearl’ failed to segregate, indicating linkage of the gene for pink flesh with the incompatibility locus, S. This was confirmed later by showing that the incompatibility ribonuclease for S3 cosegregated with pink flesh in the former progeny.

Using AFLPs, bands differentiating DNA bulked variously for apetaly, columnar habit, purple pigmentation, woolly apple aphid resistance and mildew resistance were detected. In the case of mildew resistance, these bands have been tested for cosegregation with phenotype. Four markers were found for Pl-w, one of which has been cloned and sequenced for conversion to a SCAR marker.

A range of pear genotypes was screened in the glasshouse for response to the pear bedstraw aphid, but some inconsistencies with the reported literature were noted. Six apple selections that may have resistance to apple sawfly were moved to an infestation plot. The apple genebank catalogue was updated to incorporate notes explaining the potential use of most of the accessions. An effort was made to obtain donors of all genes in the literature.

By analysing stylar ribonucleases, it was possible to assign incompatibility genotypes to 50 apple cultivars. Analysis of the new ‘Meridian’ revealed only one allele, S5. Use of primers showed that the second allele, S9, was present, even though it was not functional. The genotype of ‘Meridian’ o o can thus be indicated at S5S9 , where S9 represents a stylar mutation. Analysis of a progeny of ‘’ segregating for resistance to woolly apple aphid indicated that the S locus is not linked with the resistance gene, despite previous reports.

Twenty-nine papers describing this work were published or submitted and 30 presentations made to growers or conferences.

The progress in the development of breeding lines and markers opens up opportunities for developing marker-assisted selection for apple, to improve the efficiency of the selection process. A CSG7 has been submitted to MAFF envisaging work on marker conversion, identification of new markers, development of high throughput screening, testing multiplexing and implementation of marker-assisted selection.

Various selections from previous projects that had been passed to the Apple and Pear Breeding Club for evaluation or development performed well. The scab-resistant apple cultivar ‘Saturn’ and the Cox- type ‘Meridian’ were released to growers via the Apple and Pear Research Council. Selections E11- 20 and E88-9 look promising in trial and 20 more apple selections and two pear scion selections were chosen for trialling. The rootstocks AR86-1-20 and AR86-1-25, resistant to woolly apple aphid and crown rot and of similar vigour to MM106, are candidates for release. Five more rootstock selections performed well in trial – AR680-2, AR801-11, AR835-11, AR839-9 and AR852-3.

CSG 15 (1/00) 2 Project Genetic improvement of resistance to pests and diseases, MAFF HH1016STF title tree habit and fruit quality in apple and pear project code

Scientific report (maximum 20 sides A4) BACKGROUND

The UK produces some 300,000 tonnes of apple and pears per annum, worth £100M at wholesale prices. For supermarket sales, fruit must be attractive with good flavour and texture and long shelf- life, and novel features are desirable. In addition, growers need reliable cropping, resistance to pests and diseases, easily managed tree form and, for main crop varieties, good storage performance. Rootstocks are needed to dwarf trees to a manageable size and these rootstocks should be easy to propagate, resistant to soil-borne pests and diseases, well anchored and free from suckers, and should induce precocious and reliable cropping.

HRI-East Malling has identified donors for most, but not all, desirable characters and has developed appropriate breeding lines. In many cases, the genetic control of the characters has been resolved. In some cases, linked molecular markers have been identified, that in future could allow marker- assisted selection. The project aims to develop breeding lines and genetic information, including markers, that can be exploited for the benefit of the industry via the complementary breeding programme funded by the Apple and Pear Breeding Club.

OBJECTIVES

(I) To identify in apple and pear sources of resistance to economically important pests and diseases for which no breeding lines have yet been developed, especially apple sawfly, pear bedstraw aphid and pear psylla.

(II) To elucidate in apple inheritance of other agronomically desirable tree and fruit characters such as dwarfing habit and low ethylene production, including novel ones such as weeping habit and red flesh, by identifying sources of these characters, if necessary, determining the genetic control and, if appropriate, detecting genetic markers.

(III) To raise seedlings of apple and pear with desirable combinations of commercial characters, including sufficient resistance to scab and mildew so that 70% fewer fungicide sprays are needed to apple than for existing cultivars.

(IV) To identify genotypically good parents in apple and pear and to accelerate seedling selection.

(V) To resolve the incompatibility genotype of ‘Meridian’ and of 50 apple cultivars from the National Fruit Collections at Brogdale (year 3 only).

METHODS AND RESULTS

General

Frequent contacts were maintained with the APRC Breeding and Varietal Development Committee and grower organisations such as the East Kent Fruit Society to help ensure the programme responded to the needs of the industry. An open day was held on 1 October 1998 to explain to growers and advisors the process of apple and pear breeding, from the strategic research funded by MAFF through nearer-market work funded by the Apple and Pear Breeding Club.

CSG 15 (1/00) 3 Project Genetic improvement of resistance to pests and diseases, MAFF HH1016S title tree habit and fruit quality in apple and pear project code TF

Advanced selections

Replicated trials of about 30 selections of columnar habit were assessed in all three years. Spring frosts hampered drastically reduced crops in 1997, but in 1998 SA544-28, -39 and -48 (‘Falstaff’ x ‘Obelisk’) were shortlisted. However, texture was disappointing in 1999, perhaps because we misjudged harvest date, and parts of the trial were retained for a further year.

Initial assessment, seedlings and crossing

In the fruit assessment plots, the progeny ‘Falstaff’ x ‘Pink Pearl’ (pink flesh) segregated for pink flesh, potentially a useful novel character, indicating this character is controlled by a dominant gene for which ‘Pink Pearl’ is heterozygous. However, the progeny ‘Telamon’ x ‘Pink Pearl’ failed to segregate, raising the possibility of linkage with the incompatibility locus as described later. Other columnar progenies were evaluated for fruit quality and disease resistance.

On the fruit assessment plots, over 3 years, some 3,500 apple selections of normal habit, worked on M9, were planted as cordons for assessment by the Apple and Pear Breeding Club, along with 325 pear selections on Quince C. About 700 columnar selections, worked on M26, were planted out.

Some 34,000 seedlings were screened over the 3 years for resistance to scab in the glasshouse and resistant survivors were planted out into an unsprayed nursery plot. Around 3,500 seedlings were raised combining different mildew resistant genes for the identification and development of molecular markers. About 25,000 seedlings, not carrying scab or mildew resistance, were raised and planted in the sprayed nursery. The progeny ‘Elise Rathke’ (weeping) and ‘Pink Pearl’ (pink flesh) segregated for weeping habit, suggesting this character is controlled by a single gene, ‘Elise Rathke’ being heterozygous. Progenies were raised of backcrosses of two ‘low ethylene’ selections E19-13 and E55-55 to ‘Gloster 69’. These were planted in the sprayed nursery and will be propagated onto rootstocks to reduce the juvenile phase.

Around 1,300 pear seedlings (including a small rootstock progeny, from a cross between P55-44 and Pyrus betulaefolia) were planted in the unsprayed nursery plots and screened for field resistance and desirable plant habit before being propagated onto Quince C rootstocks.

Many of the crosses made in Year 1 of the project failed, due to severe frosts. However, years 2 and 3 proved more favourable. Approximately 40,000 apple and 10,000 pear flowers were hand- pollinated. Some seeds were produced in year 3 of the cross between ‘Comice’, quince-compatible, x ‘Williams’, quince-incompatible, but sowing was delayed for a year as the progeny size was felt to be too small for the analysis required.

Molecular markers

DNA was extracted from 48 seedlings of a progeny segregating for apetaly and screened for AFLP markers, a more powerful technique than RAPDs, using a bulked segregant approach. Two putative marker bands have been identified, one for apetaly and the other for normal flowers., cosegregation with phenotype has not yet been demonstrated. A complementary approach was tried in which DNA, restricted with a range of enzymes, was probed with the Ap3 and Pi sequences from Arabidopsis, one of which we expect to be homologous to the apetaly gene in apple. However, the probes failed to hybridise well. In 1999, we tired the Ap3 homologue which had been identified in strawberry by David James’s group. This hybridised well with the parental DNA, but seedlings have not yet been checked to see if the RFLPs revealed cosegregate with the apetalous phenotype.

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DNAs were extracted from an apple progeny resulting from the intercross of two scab-resistant selections, ‘Saturn’ and E11-20. These were analysed using SCAR markers linked to the Vf gene for scab resistance to distinguish the homozygous seedlings which should ‘breed true’ for scab resistance, thus avoiding the need for screening in the next generation. Sixteen Homozygous individuals were identified. DNAs were extracted from two apple progenies segregating for two different mildew-resistance genes, Pl-w and Pl-D, and screened for AFLP markers using a bulked segregant approach. Four markers were found for Pl-w, one of which has been cloned and sequenced for conversion to a ‘breeder-friendly’ SCAR marker. A marker has also been found for Pl- D.

The progeny ‘Northern Spy’ x ‘Totem’ was analysed with AFLPs to detect marker bands for columnar habit, purple pigmentation, woolly apple aphid resistance and mildew resistance, using a bulk segregant approach. Candidate markers were detected for all four characters, but have not yet been converted into SCARs and checked for cosegregation with phenotypes.

Donors and genebank

Apples thought to have resistance to sawfly, including A140-10, A810-222 and E249-5, representing three generations of derivatives of M. zumi, as well as M. zumi itself, were maintained as potted trees, and, when they flowered for the first time at the end of year 3, were moved to an unsprayed, infested plot. Two other varieties, ‘Gascoignes’ Scarlet’ and ST14-20, also thought to be resistant, were included.

A range of pear varieties and selections were identified as possible sources of resistance to pear- bedstraw aphid. Actively growing shoots were inoculated in year 3 on potted trees in the glasshouse. The aphids colonised some, but not all the genotypes. However, with respect to the results for ‘Conference’, the results were inconsistent when compared with replicated field data so the trees have been maintained for further testing. It may be that different races of aphid exist.

The apple genebank catalogue was updated to incorporate notes explaining the potential use of most of the accessions. Items of no further value were discarded and an effort was made to obtain donors of all Malus genes described in the literature.

Incompatibility

In a preliminary study, it was found that incompatibility, S, genotypes of apple cultivars could be determined using the stylar ribonuclease array developed at HRI-East Malling. Subsequently, styles were collected from 50 cultivars at the National Fruit Collection, Brogdale, and at HRI-East Malling, of interest to growers or for breeding or genetic studies. The proteins were extracted and separated by isoelectric focusing and the gels stained for ribonuclease activity. Triploids showed two principal bands and triploids three. By comparing with standard cultivars representing the 26 known S alleles, the genotypes were deduced. Three ‘new’ bands were detected and are attributed to S28, S29 and S30. Genotypes are listed below:

Cultivar S alleles Cultivar S alleles Arkansas S2S3S5 Katja S5S24 Barry S5S25 Kent S3S9 Cacanska Pozna S7S28 Kidd’s Orange Red S5S9 Chieftain S7S28 Kinrei S2S28 Calville Blanc d’Hiver S2S3 Kougetsu S3S7 S3S8 Leonie de Sonnaville S5S7

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Cortland S5 S25 S2S5 Cox’s Early Export S1S5 Macoun S3S25 Cox’s Pomona S1S9 Milton S1S25 Dabinette S3S8 Murasaki S7S28 Delicious (Red) S9S28 Obelisk S9S25 Delprim S3S24 Orange de Sonnaville S5S7 Directeur van de Plassche S5S7 Oxford Sunrise S9S30 S1S24 Prince Charles S2S5 Early McIntosh S5S25 Prinses Beatrix S5S7 Elan S2S5 Prinses Irene S5S7 S3S10 Prinses Marijke S5S7 Freyburg S2S5 Redgold S2S28 George Carpenter S1S3 Redsleeves S10S24 Glockenapfel S15S29 Sampion S3 S5 Gloster 69 S28S29 Shamrock S2S25 S2S3 S9S10 Jonamac S7S25 Spencer S2S10 Jonadel S7S28 Stark Blushing Golden S3S7 S5S7S9 Vista Bella S10S24

Styles were likewise analysed of the new cultivar, ‘Meridian’, which comes from the cross ‘Cox’ (S5S9) x ‘Falstaff’ (S2S5). However, only one band, corresponding to S5 was seen. To investigate this anomaly, DNA was extracted and amplified with primers designed from the sequences for the alleles S2, S5 and S9. Amplification was obtained for S5 and S9. Thirty-four young seedlings of the backcross ‘Falstaff’ x ‘Meridian’ were also analysed with these primers. They segregated for the genotypes S2S9 and S5S9. We conclude that a mutation, possibly epigenetic, has occurred whereby the S9 allele in ‘Meridian’, though present, does not code for a functional ribonuclease. The genotype o o of ‘Meridian’ is thus S5S9 , were indicates a stylar mutation. Although more work would be needed to resolve the nature of the suppression of the S9 allele, in the meantime we can conclude that ‘Meridian’ can be pollinated by all other cultivars analysed so far, including ‘Cox’, S5S9.

The failure of ‘Telamon’ x ‘Pink Pearl’ to segregate for pink flesh colour raised the possibility that the two parents share an incompatibility allele and that the pink flesh gene is tightly coupled with one allele. Analysis of stylar ribonucleases showed the two cultivars share S3. And analysis of ‘Falstaff’ x ‘Pink Pearl’, a fully compatible cross, showed that pink flesh indeed cosegregated with S3.

Some early work carried out at HRI-East Malling and the John Innes Institute that suggested woolly apple aphid resistance in ‘Northern Spy’ is tightly linked with the S locus was followed up. The progeny ‘Northern Spy’ x ‘Totem’ was screened for aphid response and for incompatibility alleles using the ribonuclease technique. No evidence of linkage was found.

DISCUSSION AND IMPLICATIONS

The cultivars released and some of the scion and rootstock selections that look promising should, when commercialised, contribute to the profitability of the UK industry by providing genetically improved cultivars meeting modern marketing and growing criteria. Progenies still undergoing evaluation will provide the next wave of advanced selections for assessment in the complementary Apple and Pear Breeding Club project and/or will provide genetic information on the inheritance of particular characters.

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The development of molecular markers for a range of agronomic characters paves the way for marker-assisted selection. This promises to increase the efficiency of the selection process. Success will depend on high throughput systems being developed for DNA extraction and analysis. The markers will need to be PCR-based and amenable to multiplexing, e.g. microsatellites. The work on incompatibility shows how problems of direct importance to the grower, i.e. pollination requirements of new cultivars, can be investigated using appropriate protein and DNA methods more rapidly than the traditional approach of making controlled crosses and monitoring subsequent fruit set.

Rootstock improvement is a particularly slow process. However, the adoption of marker-assisted selection is premature, as few markers for rootstock characteristics have been detected. Nevertheless, there is scope for streamlining the screening process. For example, instead of digging up the roots and measuring the percentage rootbank as a predictor of dwarfing, it may be simpler to bud the seedlings in their second year with a ‘space-saving’ columnar scion and monitor the subsequent growth.

POSSIBLE FUTURE WORK

The work described has developed breeding lines, detected markers, clarified genetics and identified donors. This paves the way for greater emphasis on marker-assisted selection, which promises to increase the efficiency of the selection process.

A CSG7 has been submitted to MAFF to take this work forward. It will concentrate on:

· marker conversion for marker-assisted selection · identification of new markers · development of high throughput technology · multiplex testing and troubleshooting · implementation of marker-assisted selection.

ACTIONS RESULTING (TECHNOLOGY TRANSFER, PUBLICATIONS AND PRESENTATIONS)

Various scion and rootstock selections from previous projects that had been passed to the Apple & Pear Breeding Club for evaluation or development performed well.

The scab-resistant apple ‘Saturn’ (formerly SA15-4) and the Cox-type ‘Meridian’ (formerly A931-15) were released to growers via the Apple and Pear Research Council in 1998 and 1999 respectively. The red-fruited, scab-resistant selection E11-20 is being considered for naming and release. The ‘’-type apple E88-9 was selected at Brogdale for bulking up for replicated trialling on grower farms. Some 20 apple scion selections and two pear scion selections were selected by the Apple and Pear Breeding Club for trialling.

Two rootstock selections, AR86-1-20 and AR86-1-25, are candidates for release. They are resistant to woolly apple aphid and to crown rot and are of similar vigour to MM106. They should be valuable for less intensive planting systems, including orchards. In trials conducted by the Apple and Pear Breeding Club of apple rootstock selections from the MAFF-funded programme, two showed particular promise. AR680-2 and AR801-11 induce tree vigour similar to M9 and M26 respectively, but have much better productivity. Three more recent selections showing early promise are AR839-9 and 852-3, which have vigour similar to M9, and AR835-11, which is between M26 and MM106.

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In all, 29 papers were published in, or prepared for, a wide range of journals, refereed and non- refereed. In addition, 30 presentations were made to visitors or conferences. These are itemised below.

Publications from previous project, and in conjunction with European Mapping Project

WEBSTER, A. D., TOBUTT, K. R., EVANS, K.M., ALSTON, F.H. & JAMES, D.J. (1997). Rootstock breeding and orchard testing at Horticulture Research International, East Malling, Acta Horticulturae, 341: 83-88. MANGANARIS, A.G. & ALSTON, F.H. (1997). Genetics of superoxide dismustase in apple. Theoretical and Applied Genetics, 95:484-489. ROCHE, P.A., BROWN, L.M., KING, G.J., ALSTON, F.H., EVANS, K.M., VRIELINK, R., DUNEMANN, F., MARKUSSEN, T., TARTARINI, S., BROWN, L.M., RYDER, C. & KING, G.J. (1997). RFLP and RAPD markers linked to the rosy leaf curling aphid resistance gene (Sd1) in apple. Theoretical and Applied Genetics 94:528-533. ROCHE, P., van ARKEL, G. & VAN HEUSDEN, A.W. (1997). A sepcific PCR assay for resistance to biotypes 1 and 2 of the rosy leaf curling aphid in apple based on an RFLP marker closely linked to the Sd1 gene. Plant Breeding, 116:567-572. UR-RAHMAN, H., JAMES, D.J., HADONOU, A.M., CALIGAN, P.D.S. (1997). The use of RAPD for verifying the apomictic status of seedlings of Malus species. Theoretical and Applied Genetics, 95:1080-1083. EVANS, K.M. (1997). Breeding with alternative sources of resistance to Podosphaera leucotricha (powdery mildew) in apple; the potential for gene pyramiding. IOBC/WPRC Bulletin, 20(9):101-104. RIDOUT, M.S., (TONG, S.), (VOWDEN, C.J.) & TOBUTT, K.R. (1998). Three-point linkage analysis in crosses of allogamous plant species. Genetical Research 72:111-121. TOBUTT, K.R., BOŠKOVI , R. & RUSSELL, K. (1998). Stylar ribonucleases to predict incompatibility alleles in tree fruit. Report of Horticulture Research International 1996/97, pp 76-77. MALIEPAARD, C., ALSTON, F.H., VAN ARKEL, G., BROWN, L.M., CHEVREAU, E., DUNEMANN, F., EVANS, K.M., GARDINER, S., GUILFORD, P., VAN HEUSDEN, A.W., JANSE, J., LAURENS, F., LYNN, J.R., MANGANARIS, A.G., DEN NIJS, A.P.M., PERIAM, N., RIKKERINK, E., ROCHE, P., RYDER, C., SANSAVINI, S., SCHMIDT, H., TARTARINI, S., VERHAEGH, J.J., VRIELINK-VAN GINKEL, M., KING, G.J. (1998). Aligning male and female linkage maps of apple (Malus pumila Mill.) using multi-allelic markers. Theoretical and Applied Genetics 97:60-73. PETROPOULOU, S.P. & ALSTON, F.H. (1998). Selecting for improved pollination at low temperatures in apple. Journal of Horticultural Science & Biotechnology 73: 507-512. TOBUTT, K.R. & ALSTON, F.H. (Eds) (1999). Proceedings of Eucarpia Fruit Breeding Symposium, Acta Horticulturae, 484. KING, G.J., ALSTON, F.H., BROWN, L.M., CHEVREAU, E., EVANS, K.M., DUNEMANN, F., JANSE, J., LAURENS, F., LYNN, J.R., MALIEPAARD, C., MANGANARIS, A.G., ROCHE, P.A., SCHMIDT, J., TARTARINI, S., VERHAEGH, J., VRIELINK, R. (1998). Multiple field and glasshouse assessments increase the reliability of linkage mapping of the Vf source of scab resistance in apple. Theoretical and Applied Genetics 96:699-708. ROCHE, P., BROWN, L.M., KING, G.J., ALSTON, F.H., EVANS, K.M., MALIEPAARD, C., HEUSDEN, S. VAN, VRIELINK, R., LAURENS, F., DUNEMANN, F., MARKUSSEN, T. & TARTARINI, S. (1999). Identification and development of markers linked to aphid resistance in apple. Acta Horticulturae, 484: 519-522.

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MALIEPAARD, C., ALSTON, F.H., ARKEL, G. VAN, BROWN, L., CHEVREAU, E., DUNEMANN, F., EVANS, K.M., GARDINER, S., GUILFORD, P., HEUSDEN, S. VAN, JANSE, J., LAURENS, F., LYNN, J., MANGANARIS, S., NIJS, T. DEN, PERIAM, N., RIKKERINK, E., ROCHE, P., RYDER, C., SANSAVINI. S., SCHMIDT, H., TARTARINI, S., VERHAEGH, J., VRIELINK, R. & KING, G. (1999). The European Apple Map. Acta Horticulturae, 484: 325-330. EVANS, K.M. (1999). New approaches to apple scion breeding at HRI-East Malling. Acta Horticulturae, 484: 171-174. DUREL, C-E., LESPINASSE, Y., CHEVALIER, M., CHEVREAU, E., NIJS, A.P.M. DEN, DJOUVINOV, V., DUNEMANN, F., EVANS, K.M., FISCHER, C., GESSLER, C., KELLERHALS, M., LAURENS, F., MANGANARIS, A.G., PARISI, L., SANSAVINI, S., SCHMIDT, H., SCHOUTEN, H.J. & SCHREIBER, H. (1999). Genetic dissection of apple resistance regarding pathogen variability: Co-ordination of European research programmes. Acta Horticulturae, 484: 435-441. TOBUTT, K.R. & ALSTON, F.H. (1999). Proceedings of the Eucarpia Symposium on Fruit Breeding and Genetics. Acta Horticulturae 484.

Publications resulting principally from HH1016STF

EVANS, K.M. (1998). Breeding for resistance to diseases. EMRA Members’ Day Report, July 1998. EVANS, K.M. (1998). Breeding for reduced ethylene production. HRI Annual Report, 1996-97, p. 81. BOŠKOVI , R. & TOBUTT, K.R. (1999). Correlation of stylar ribonuclease isoenzymes with incompatibility alleles in apple. Euphytica, 107:29-43. TOBUTT, K.R., BOŠKOVI , R. & ROCHE, P. (2000). Incompatibility and resistance to wooly apple aphid in apple. Plant Breeding 119: 65-69. EVANS K.M. & ALSTON F.H. (2000). Apples - the genetics of fruit quality. In: Encyclopedia of genetics. Ed E.C.R.Reeve. Fitzroy Dearburn. in press KING, G.J., MALIEPAARD, C., LYNN, J.R., ALSTON, F.H., DUREL, C.E., EVANS, K.M., GRIFFON, B., LAURENS, F., MANGANARIS, A.G., SCHREVENS, E. & TARTARINI, S. (2000). Quantitative genetic analysis and comparison of physical and sensory descriptors relating to fruit flesh firmness in apple (Malus pumila Mill.) Theoretical and Applied Genetics (accepted) PHILLIPS, K.L., JAMES, C.M., CLARKE, J.B. & EVANS, K.M. (2000). Identification of molecular markers linked to mildew resistance genes Pl-w and Pl-d in apple. IOBC/WPRS Bulletin. Angers 1999 (in press). ALSTON, F.H., PHILLIPS, K.L. & EVANS, K.M. (2000). A Malus Gene List. Acta Horticulturae (in press). EVANS, K.M., LESPINASSE, Y., PARISI, L., LAURENS, F. & DUREL, C.E. (2000). Durable resistance to scab and mildew in apple - a European project. Pesticide Outlook (in press). ALSTON, F.H. & EVANS, K.M. (2000). Apples – the genetics of resistance to diseases and insect pests. In: Encyclopaedia of Genetics (Ed. E.C.R. Reeve). (Submitted).

Presentations

K.M. Evans. Selecting for fruit quality in new apple varieties. Talk to EMRA members, East Malling, April 1997. K. R. Tobutt. Incompatibility in tree fruits. Talk to East Malling Trustees, East Malling, June 1997. K.R. Tobutt. Use of genetic resources in top fruit breeding. Talk to Nordic Genebank Fruit Group, East Malling, September 1997. K.M. Evans. Apple breeding. Growers walk at Brogdale Fruit Trials, September 1997.

9 Project Genetic improvement of resistance to pests and diseases, MAFF HH1016S title tree habit and fruit quality in apple and pear project code TF

K.R.Tobutt. Top fruit breeding. Talk to students from Zullier, Switzerland, East Malling, September 1997. A.M. Hadonou. Markers for early selection and cultivar identification. Talk to Nordic Genebank Fruit Group, East Malling, September 1997. K.M. Evans. Saturn. Marden Fruit Show Press Conference, October 1997. K.R. Tobutt. Marker-assisted selection and genomics. To HRI Board of Directors, East Malling, April 1998. K.R. Tobutt. Rootstock breeding and opportunities for molecular markers. To EU Concerted Action on Rootstocks, HRI-East Malling, June 1998. K.M. Evans. Breeding for fruit quality. At HRI-East Malling Open Day, June 1998. K.M. Evans. Breeding apples for resistance to diseases. At HRI-East Malling Members’ Day on Reducing Inputs in Fruit Production, July 1998. K.M. Evans & K.R. Tobutt. East Malling’s apple breeding programme. Talk to East Kent Fruit Society, East Malling, August 1998. K.R. Tobutt. Fruit breeding and columnar apples. To KCC visitors, HRI-East Malling, August 1998. K.R. Tobutt. Seedling selection. At the Apple and Pear Breeding Club Day, HRI-East Malling, October 1998. K.M. Evans. Apple and pear breeding. At the Apple and Pear Breeding Club Day, HRI-East Malling, October 1998. C.M. James. Apple breeding at HRI East Malling. At “Apple Day”, Cambridge University Botanic Garden, October 1998. C.M. James. Molecular markers for crop improvement. At HRI East Malling, Open Day, June 1998. C.M. James & A.M. Hadonou. Marker-assisted selection and mapping. For Belgian students, May 1998, HRI-East Malling. C.M. James & A.M. Hadonou. Marker-assisted selection. For Wye College students, May 1998, HRI-East Malling. C.M. James & A. M. Hadonou. Marker-assisted selection in fruit breeding. To Institute of Biologists, April 1998, HRI-East Malling. C.M. James. AFLPs and marker-assisted selection. To Shell International Renewables and Plant Breeding and Biotechnology Groups, June 1998, HRI-East Malling. K.R. Tobutt. Incompatibility genetics in Prunus and Malus. At BBSRC Self-Incompatibiilty meeting, Sutton Bonington, April 1999. K.R. Tobutt. Breeding new varieties of top fruit. To horticultural students from Belgium, East Malling, April 1999. K.R. Tobutt. Top fruit breeding. To George Nobel and MAFF colleagues, East Malling, May 1999. K.R. Tobutt. Plant breeding. To students from Capel Manor, East Malling, June 1999. K.R. Tobutt. Breeding columnar apples. To students from University of Bonn, East Malling, July 1999. K.R. Tobutt. Top fruit breeding. To fruit growers from Aomori, East Malling, August 1999. K.R. Tobutt. Genome analysis in Rosaceae. To BBSRC visitors, Wellesbourne, September 1999. K.R. Tobutt. Incompatibility genetics in Prunus and Malus. Talk to Wye/Imperial/East Malling Plant Science Workshop at Wye, November 1999. K.M. Evans. Breeding for organic apple production. HRI organic workshop, East Malling, February 2000.

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