Pollen and pollination experiments. I. the contribution of stray pollen to the seed set of depetalled,hand-pollinated flowers of apple

Summary Using the finger nails to remove most of the petals is a fast method of preparing apple flowers for hand pollination in the balloon stage. Such flowers still appeared quite attractive to insects; left to open pollination, the seed set per flower averaged two thirds of that of similarly treated, but hand-pollinated flowers. With the aid of marker pollen of scab or mildew resistant donors and the subsequent screening of the seedling progenies for resistance, it was shown that the stray pollen was responsible on average for one of every three seeds formed by unbagged depetalled and hand-pollinated flowers.     

The interaction between compatible and self-incompatible pollen of apple and pear as influcenced by their ratio in the pollen cloud

Summary The production of self-seed was investigated in apple and pear with the aid of viable and compatible mentor/ pioneer pollen in relation to the proportion of self-imcompatible pollen present in the pollen cloud.Treatments consisted of mixtures of compatible and self-incompatible pollen at ratios of 1:1, 1:2 and 1:9 applied once or twice or followed by self-pollination. Selfing only, whether once or twice, produced virtually no fruits or seeds, while mixed pollinations did. Generally, fruit set tended to decrease and the self-seed to increase with increasing amounts of self pollen in the pollinations.The pioncer-pollen method (compatible in advance of self pollen) appeared more effective than the mentor pollen method (compatible and self pollen mixed). The use of viable instead of dead mentor/pioneer pollen causes competition for the ovules with the self pollen, but has the advantage that, in addition to some self-seed, other seeds are formed which, e.g. in apple, are necessary to keep the fruits on the tree until harvest.Fruit set was moderately reduced at ratios of compatible pollen to self pollen not exceeding 1:9 with pear and 1:5 with apple which constitute maximal ratios in practice as regards pollinator trees: trees of the main cultivar. In fruit orchards and probably in nature with other species, the interaction between compatible and self pollen may result in more self-seed, and so suggests that inbreeding played a greater part in the evolution of self-incompatible species, than the meager results of articial self-pollination imply.     

The performance of double or mixed pollinations with compatible and self-incompatible or incongruous pollen of pear and apple

Summary Six pear and five apple trials were carried out to ascertain the outcome of combinations of compatible pollen (C) with self (S) or incongruous pollen (I) as to the pollination index (PI=seeds/pollinated flower). The PI of the mixture C+I (1:5) was consistently depressed as compared to that of the control C. The results of the double pollinations S/C and I/C were affected by the temperature at pollination; their PI's at <15°C were twice as high as those at >15°C, being well above and below the PI of C in the former and latter case respectively. The opposite was true for the C/S combination, the PI of which increased with the pollination temperature; the PI of C/I did not differ much from the PI of C, irrespective of temperature. The conclusion was reached that the interaction previously and presently found between compatible and self-incompatible pollen also exists to a fair extent between compatible and incongruous pollen. However, in pear neither the mentor nor the pioneer pollen technique proved to aid its hybridization with apple, the formation of self seed was not observed either. In apple the production of apple × pear hybrids was likewise doubtful, but the double pollinations S/C and C/S formed 4–10% self seed.     

Embryo development and fruit-set in pear induced by untreated and irradiated pollen

Summary Flowers of the pear cultivars Conference, Doyenné du Comice and Gieser Wildeman were pollinated with untreated and 50-krad-irradiated pollen. Fruitlets were sectioned at regular intervals up to 6 weeks after pollination. Tubes of the irradiated pollen grew slower, but within two weeks nearly as many embryos were initiated as in the case of untreated pollen. Thereafter, the proportion of degenerated embryos increased, although the endosperm sometimes persisted. Six weeks after pollination only about 21% of the seeds induced by the irradiated pollen still contained normal embryos. At or before that time in-vitro culture of the immature seeds will be necessary because few embryos survive until harvest. The irradiated pollen stimulated a rather large parthenocarpic fruit-set, presumably associated with endosperm rather than embryo development during the first 4–6 weeks after pollination. The possibilities of haploid production are discussed.     

Dwarfing and lethal genes in apple progenies

Summary Three dwarf types are described, early dwarf, determined by two recessive genes d ₁ and d ₃, in the double homozygous state, and an additional recessive gene d ₄ and crinkle dwarf and sturdy dwarf each also determined by a recessive gene.The gene l, for pale green lethal was found to be closely linked to the gene V _f , for scab resistance.There was evidence of a regrowth promoting gene, G, in some seedlings homozygous for d ₁ and d ₃.     

Restrictions of the efficiency of haploid plant production in apple cultivar Idared,through parthenogenesisin situ

Summary For haploid plant production in Idared apple, parthenogenesisin situ was induced by irradiated pollen. The most important restrictions were seed set, embryo germination and green plant recovery from germinated embryos. The efficiency of green plant production was strongly influenced by irradiation dose, picking time and quality of the irradiated pollen. 250 Gy pollen was more efficient for green plant production than 500 Gy pollen. Using 250 Gy pollen with reduced germination capacity resulted in a five-fold lower green plant yield. Delaying picking of the fruits (140 instead of 70 days after pollination) had a significant positive effect. Up to 20 plants per 1000 pollinated flowers, showing no expression of a marker gene present in the pollen parent, were obtained in some treatments. Induction efficiency was high when seed weight was high. This offers the possibility of selecting seeds with the best chances for haploid plant production. Heavy seeds generally contained visual normal developed embryos, which germinated better than those from lighter seeds and gave normal plant development. Delayed picking increased the proportion of large seeds, suggesting that pollination with irradiated pollen retards embryo development. From treatments with irradiated pollen of poor quality more small seeds were harvested.     

Optimal sampling strategies for evaluating fruit softening after harvest in apple breeding

Environmental variance components associated with year, tree, and harvest date were estimated for fruit softening after harvest in apple (Malus × domestica Borkh.) to determine their relative importance and design optimum sampling strategies to discriminate genotypes in apple breeding. Fruit were stored after harvest under 20± 2 ^∘C and 80± 5%RH. Softening was evaluated by adapting the change in firmness during storage to a linear regression and defining the regression coefficient as the softening rate. Environmental variances associated with genotype × year interaction, among trees, year × tree interaction, and among harvest dates were all very small, namely, 2.7, 0.1, 5.2, and 5.7%, respectively, to the total variance obtained from the analysis of variance for the softening rate. The variance associated with genotype, at 57.3%, was very large. On the basis of the number of fruit necessary for firmness measurements, two times harvest is an efficient strategy to determine a genotype mean for the softening.     

Genetics and histology of powdery mildew resistance in apple

Summary Eight scab-resistant cultivars and selections along with eight commercial apple cultivars were evaluated for powdery mildew resistance in greenhouse and nursery tests. Dayton, Liberty, Delicious and Tolman Sweet were rated moderately resistant to infection in both greenhouse and nursery tests. Segregation of seedlings among 14 progenies for mildew reaction indicated that mildew resistance is polygenically controlled in this material with additive gene effects. Recovery of mildew resistant seedlings from crosses involving a scab-resistant parent(s) suggested that this material can be useful in developing scab- and mildew-resistant apple cultivars. Histological investigations were conducted to describe mildew symptoms of infected leaves.     

Pollen and pollination experiments. II. The influence of the first pollination on the effectiveness of the second one in apple

Summary Pollinating apple cultivars twice with compatible pollen at an interval of one or two days produced about twice as many seeds per pollinated flower as a single pollination. With the aid of scab- or mildew resistant marker pollen, it could be shown that the second pollen formed on average twice as many seeds as the first. The first pollen appears to pave the way-partly at its own cost-for the second and was therefore called pioneer pollen.     

Screening for resistance to Nectria galligena Bres. in cut shoots of apple

Summary To find a fast and reliable test to assess resistance to Nectria galligena in apple, different methods of inoculation were compared using macroconidia of N. galligena and one-year-old cut shoots from mature trees of Cox's Orange Pippin, IVT 69078-19, James Grieve and Jonathan.With the best inoculation method 11 genotypes were screened for resistance. Elstar, Golden Delicious, Jonathan and Lombart's Calville were highly resistant and the level of resistance of Ingrid marie, Gloster, Melrose, IVT 69078-19, Cox's Orange Pippin, James Grieve and Idared decreased in this order.The best inoculation method proved to be simple, giving results within four to nine weeks after inoculation.     

Evidence of gene introgression in apple using RAPD markers

Summary A genomic remnant of Malus floribunda clone 821 introgressed into the cultivated apple M. x domestica Borkh. was identified using randomly amplified polymorphic DNA (RAPD) markers obtained by the polymerase chain reaction (PCR). Using a set of 59 oligonucleotide decamer primers, polymorphic DNA markers were identified among three pooled DNA samples. Based on the presence or absence of bands among bulked apple scab-resistant selections and cultivars, bulked scab-susceptible cultivars, and a M. floribunda clone 821 sample, one primer, A 15, identified amplified fragments in the scab-resistant bulked sample that was also unique to the M. floribunda clone 821. The unique band from M. floribunda clone 821 was amplified in four out of 17 scab-resistant selections/cultivars. This RAPD, designated OA15₉₀₀, identifies an introgressed fragment that has as yet no known function.     

Quantitative genetics in an apple breeding strategy

Summary Open-pollinated seed of 500 genotypes of commercial cultivars, breeding selections and wild species of apples (Malus × domestica) was collected from overseas mixed-clone repositories. This is to generate the first generation of the new Apple Breeding Population.Subsequent generations will be intercrossed in a pair-crossing design. Recurrent selection for general combining ability (GCA) is the breeding method used for genetic improvement of the Breeding Population. Every parent in the control-pollinated breeding population is tested for GCA using a polycross mating design.The best individuals of the Breeding Population, intensively selected between families using the polycross GCA estimates, and also within families are intermated to produce a Cultivar Production Population. Trees selected from this have then to be evaluated in grafted clonal tests before commercial release.Each generation, a new Breeding Population of 500 genotypes is selected from the pair-cross families, based on parental GCA values (from polycross tests), full-sib family means and individual performance and then intercrossed. Greatest emphasis will be on within-family selection not between-family, so as to avoid reducing the effective population size.     

Inheritance and selection of some fruit characters of apple. II. The relation between leaf and fruit pH as a basis for preselection

Summary The leaf pH of two year old apple seedlings was found to be sufficiently related to the fruit pH, measured when the seedlings were six to seven years old, to serve as a basis for pre-selection. Discarding all seedlings with an above average leaf pH (40% of the total) eliminated the majority (74%) of seedling with low-acid cum sweet fruits. The method was not effective for reducing the proportion of seedlings with high-acid cum sour fruits.     

The effects of apple rootstock genotypes on mineral content of scion leaves

Summary The relative effects of year to year variation and rootstock genotypes on the major content of leaves of four apple scion cultivars were studied. For nitrogen and phosphorus the main source of variation was due to years. For potassium, magnesium and calcium there were significant genetic effects. The results are discussed in relation to breeding for improved efficiency in mineral uptake in apple rootstocks. Particular reference is made to uptake of minerals associated with physiological disorders of stored apples.     

Induced mutants of Cox's Orange Pippin apple with increased self-compatibility. 1. Production and selection

Summary More than 7000 MV₂ trees derived from 225 MV₀ grafts of Cox's Orange Pippin apple that had been treated with approximately 7.5 kR gamma-rays were grown as a solid block with no pollinators in an isolated orchard. The orchard was sited in South Wales well away from any external source of apple pollen and due to this lack of pollination the great majority of the trees produced very few fruit. However, a few individual trees produced acceptable quantities of fruit under these conditions. Twelve trees maintained this performance for seven years and these trees have been selected for trial to test for mutants with increased self-compatibility.     

Pollen and pollination experiments. VI. Heat resistance of pollen

Summary Pollen of dry apple, pear, lily and rose pollen was heated up to 48 h at a range of temperatures. About half or more than half of the pollen grains survived 48 h at 40 C, 24 h at 50 C, 8 16 h at 60 C. 4 8 h at 70 C, more than one hour at 80 C. and between 10 and 20 min at 90 C. Presumably, pollen able to withstand low humidity is also heat resistant, a property which may be usable to make pollen virus free through heat treatment and perhaps to overcome incompatibility.     

Microsatellite (SSR) variation in a collection of Malus (apple) species and hybrids

A collection of 142 accessions of 23 Malus species, derived hybrids and cultivar accessions from the USDA-ARS Plant Genetic Resources Unit's core collection, which represents an extensive range of Malus species, was screened with a set of previously described SSR (simple sequence repeat) markers. The markers were used to determine genetic identities, estimate genetic diversity, identify genetic relationships among the accessions, and determine the utility of SSR primers developed from Malus ×domestica for making genetic assessments across the whole Malus genus. All eight primer pairs amplified multiple fragments when used in polymerase chain reactions with DNA from these accessions. High levels of variation were detected with a mean of 26.4 alleles per locus and a mean direct count heterozygosity across all eight loci equal to 0.623. The eight primer pairs used in this study unambiguously differentiated all but five pairs of accessions in this collection of 142 accessions of 23 Malus species, derived hybrids and cultivars. These SSR data were not useful in identifying genetic relationships among this diverse collection of accessions, with the majority of the accessions not clustering in ways concordant with taxonomic information and/or geographic origin. The resulting phenogram resolved only two meaningful clusters, for the taxonomically isolated Section Chloromeles and for M. fusca accessions, reflecting genetic relationships arising from geographic origin. The detection of identical accessions in the collection, which were previously considered to be unique, highlights the critical need to further bolster collections of certain Malus species. This revised version was published online in July 2006 with corrections to the Cover Date.     

A comparison of apple and pear seedlings with reference to the juvenile period I. Seedling growth and yield

Summary Several thousand apple and pear seedlings of many progenies were studied in connection with their juvenile period (J.P.). The initially significant inverse relation between the vigour (stem diameter) and the J.P. of the seedlings became insignificant as the trees grew older, due to a retardation of growth occurring when the seedlings become generative. This relation can be used effectively in pre-selecting for vigour in the nursery. Cumulative yields were higher when the J.P. was shorter, but there was no evident link between the J.P. and annual yield in full bearing, that is to say, precocity and preductivity are not directly connected. Better growing conditions and pre-selection have shortened the mean progeny J.P. of either crop by three years since the 1950's. Generally, the pear seedlings appear to grow faster, to become generative slower and to yield lower than comparable apple seedlings. The mean J.P. of apple and pear progenies averaged 4.2 and 6.0 years respectively. The difference between apple and pear may be attributable to a greater selection pressure on both precocity and productivity with apple than with pear.This possibly also led to a smaller tree in the productive phase on the assumption that tree size at first flowering-small at a short, large at a long J.P.-expresses reproductive efficiency and as such is reflected in the size of the adult tree.Also presented at the Juvenility Symposium held at West-Berlin, November 1976.     

The stability of induced compact mutant clones of Bramley's Seedling apple

Summary Twelve selected, compact, clones of Bramley's Seedling induced by gamma radiation treatment were checked for stability. Representative trees were used as vegetative parents for large scale multiplication, and further buds were treated with gamma radiation to disrupt the structure of their meristems. The results indicate that seven of the clones are as stable as the original cultivar, and therefore probably homohistont, containing only cells with compact mutant genotype. The other five clones proved to be unstable and gave rise to a large proportion of apparently normal trees. It is hypothesized that in these clones the L₁ (epidermis) consists of normal unchanged tissue, while the bulk of the plant tissue layers are of mutant cells, i.e. that they are periclinal chimaeras with the genotypes of the different cell layers coded for different growth forms.     

Induced mutants of Cox's Orange Pippin apple with apparent increased self-compatibility. 2. Laboratory and orchard assessment and testing

Summary Fruit set on clones of Cox's Orange Pippin apple which had been produced by gamma-irradiation, and found in a previous trial to crop when isolated from the pollen of other cultivars, was compared after open or hand-pollination. Some clones set more fruit than the unirradiated control trees when open pollinated or when hand-pollinated with pollen from the same tree or control Cox trees. Pollen from some mutant clones also improved set on standard Cox (EMLA). Estimates of the numbers of pollen tubes reaching the base of the style indicated that the increased set was due to enhanced tube growth.