Inheritance of pseudo-self compatibility in self-incompatible garden and greenhouse chrysanthemums, Dendranthema grandiflora Tzvelv

Summary Self incompatibility (SI) can be used to alleviate costly hand emasculation and pollination in F₁ hybrid chrysanthemum seed production. SI, however, disrupts the progression of inbreedig (selfing or full-sib mating). Consequently, inbreds are selected for breakdown of the SI system or the presence of pseudo-self compatibility (PSC). PSC inbreds, recombinant inbreds, and noninbred cultivars were selfed and/or intercrossed to determine PSC expression across environments and generate 1–3 inbred generations (I₁–I₃). Percent PSC ranged from 0–68.8% for inbred parents, 0.2–99.7% for recombinant inbreds, and 0.6–25.7% for noninbred cultivars. There was no indication of end-of-season PSC. The majority of parents (78%) were classified as low PSC and this trend continued in the I₁ (70.1%), I₂ (65.6%), and I₃ (83.6%) generations; mid PSC was rarer (11.9–18.8%) and high PSC the least common (4.5–15.6%). PSC distributions were primarily continuous, rather than discrete, indicating quantitative inheritance. In several inbred families, 100% of the I₁ individuals were SI; this was not correlated with parental PSC level. Inbred families derived from selfing low and mid PSC parents were the most likely to reach extinction due to inbreeding depression. High PSC was not highly heritable, since I₁ progeny were predominantly SI or had low to mid PSC levels. Most 153-1 values for PSC:SI segregations (1:1, 1:3, 3:1) were not significant at the 5% level. Realized heritability (H _R ) estimates for PSC ranged from a low of 0.05% to 10.19%, although increased H _R values did not account for inbreeding depression or genetic mechanisms preventing selection for high PSC. The highest individual %PSC increased over the high parent with SI × low PSC or SI × mid PSC parents in all cases; progeny means did so only in SI × low PSC. Low × low crosses were split evenly between an increase and decrease in progeny or highest individual mean. Since all low × low crosses and low selfs produced 43–50% of the progeny with PSC levels higher than the parents, it appears that most low parents possess some unexpressed PSC genes. Mid PSC parents responded similarly to low PSC genotypes. An increase in PSC was found when crossing SI × PSC parents, illustrating a threshold effect. PSC × PSC crosses (high × low, low × low) produced SI, low, and mid PSC I₁ individuals but did not show a heterotic effect, since the PSC parents had already surpassed the PSC threshold. Selfing high PSC parents, however, suggested different genetic control. Progeny and highest individual means behaved the same; 83% decreased and 17% remained the same for PSC levels. The drop in PSC indicated non-additive gene action. Thus, the PSC threshold with additive gene action holds when selection for higher PSC levels is being done from low and mid PSC parents, but once high PSC levels are obtained, non-additive gene action prevails.Abbreviations self pollinated - H _R Realized Heritability - PSC Pseudo-Self Compatibility - SC Self Compatibility - SI Self Incompatibility     

Effects of irradiance and juvenility on the selection of chrysanthemums

Summary Chrysanthemum morifolium seedlings and cuttings from those seedlings were grown to flowering at 17°C, short days (8 h) and two levels of irradiance (10 and 30 W m^–2). The cuttings flowered more readily than the seedlings at both irradiances, Especially at 10 W m^-2 many seedlings remained vegetative. The delay in flowering of the seedlings was accompanied by a higher number of leaves produced before flowering.The number of leaves produced by the seedling in excess of the number of leaves produced by the cutting from that seedling varied considerably between genotypes. The correlations between seedlings and the corresponding cuttings for days to flowering and number of leaves were low but generally significant.It is concluded that cuttings are preferred over seedlings in breeding programmes that aim at developing chrysanthemums for low light winter production.     

Inbreeding depression in garden and glasshouse chrysanthemums: germination and survivorship

Summary Sixty-six chrysanthemum (Dendranthema grandiflora) inbred selections, noninbred cultivars and hybrids, and D. makinoi were the base populations from which up to three generations of inbreds were obtained using multiple plant descent. Each parent possessed pseudo-self compatibility (PSC), which allowed seed set following self pollination. Rapid-generation cycling (laboratory seed ripening to heart stage and subsequent embryo rescue) reduced generation time and minimized confounding maternal with zygotic inbreeding depression during post heart-stage seed development. Selection criteria were male/female fertility and PSC. Two stages of the life cycle were chosen to evaluate inbreeding depression: germination (seed development to germination) and survivorship (fertile individuals at anthesis). PSC was environmentally interactive and genotype dependent, causing variable levels of self seed set between and within populations and generations. By the end of the second generation, families from all noninbred cultivars, D. makinoi, and one F₁ hybrid were eliminated from the experiment due to self incompatibility and/or inbreeding depression. Post-germination inbreeding depression was severe in several advanced inbreds. Inbred progeny of most F₁ or F₂ hybrids expressed less or equal amounts of inbreeding depression compared to advanced inbreds. Linear regression coefficients for either germination or survivorship on percent homozygosity were negative. Correlation coefficients between percent germination and survivorship (as a percent of seed set) were highly significant for Minnesota inbreds (r=0.67, P0.002) and hybrids (r=0.67, P0.006). The correlation coefficient was higher when percent germination and survivorship (as a percent of germinated seedlings) were compared (r=0.95, P0.001).Abbreviations F Fisher's Coefficient of Inbreeding - IBD Inbreeding Depression - PSC Pseudo-self Compatibility - RGC Rapid Generation Cycle - SI Self Incompatibility     

Genetic analysis in Chrysanthemum morifolium. I. Flowering time and flower number at low and optimum temperature

Summary The number of days to flowering and the number of flowers per plant were studied at 12, 13, 15 and 17 C night temperature in 79 F₁ populations from 15 parents. Only a few F₁ plants flowered earlier than their parents; the majority flowered at the same time, later or not at all. The number of flowers of the F₁ plants was higher than that of the parents.Genetic analyses showed a broad sense heritability of 70% for days to flowering and flower number and a highly significant GCA effect for days to flowering.The progenies and parents that flowered early at one temperature were generally early at all temperatures.     

Genetic analysis in Chrysanthemum morifolium. II. Flower doubleness and ray floret corolla splitting

Summary The inheritance of flower doubleness and floret corolla shape was studied in 70 F₁ populations from 16 parents. Singleness is partially dominant over doubleness. The overall segregation for doubleness in crosses of the type double × single was 29% single, 58% semi double and 12% double. The percentages varied with the parents.The anemone flower type is inherited as if it were a single. Crosses between plants with ligulate (split lengthwise) and tubular corollas produced ligulate, tubular and intermediate types in various ratios depending on the parents used. Flower doubleness and corolla shape are not linked.     

A mutant of a mutant of a mutant of a...: Irradiation of progressive radiation-induced mutants in a mutation-breeding programme with Chrysanthemum morifolium Ram.

Summary Radiation-induced sports in Chrysanthemum morifolium Ram. have been reported for several years. It has become an everyday practice to produce flower-colour mutants from outstanding cross-breeding products, even before they are distributed for the commercial production of cut flowers.One of the most successful and recent examples is that of cv. Horim, of which hundreds of mutants were produced by successive use of radiation-induced mutants in the mutation-breeding programme. Over about 4 years a variety of flower-colour mutants was obtained, not only largely including the outstanding characteristics of the original cultivar but sometimes even with an appreciable improvement in quality and yield. It is expected that the latter types, the Miros group, will soon completely supersede the spontaneous or radiation-induced Horim sports and mutants and take over the leading position of the Horim group in the production of all-year-round (AYR) cut flowers.     

Rapid detection of S-glycoproteins of self-incompatible crucifers using Con-A reaction

Summary As S-specific glycoproteins react with concanavalin A (ConA), they could be stained sensitively with ConA-FITC after cellulose acetate isoelectric focusing of stigma proteins in self-incompatible crucifers. By this method the S genotype of an individual plant could be inferred within 4 h using 4 flowers. This simple and rapid method may be applicable for describing S genotypes in practical F₁ breeding.     

Self-incompatibility in Brassica napus: seed set on crossing 19 self-incompatible lines

Summary Seed set after self-pollination confirmed that 19 lines of Brassica napus were self-incompatible. Eight lines, H, J, Q, W, X, K, P and Z, were fully cross-compatible. Line R was cross-compatible with these lines but often had a low seed set as female parent. These results are consistent with the activity of nine distinct S-alleles. Line S was cross-incompatible with K, as was V with P, and F with Z. With each of the lines A, E, B, O, G, L and M at least one reciprocal difference was found so that the number of additional distinct S-alleles could not be inferred, but there must be a minimum of seven.Pollen tube counts of intra- and inter-line pollinations using M, B and X confirmed the homozygosity of these lines with respect to self-incompatibility status and the presence of non-reciprocal compatibility between M and B.The results are interpreted in terms of the activity of both B. oleracea and B. campestris S-loci and the implications for hybrid breeding are discussed.     

Analysis of components controlling early flowering of chrysanthemum at low temperature

Summary Twenty seven clones of chrysanthemum, 19 of which were selected at low temperature, were flowered at 10, 14, 18 and 22°C. The number of days to flower in response to temperature could, for each genotype, be described by a parabola with the parameters: optimum temperature, days to flower at the optimum temperature and curvature which represents the sensitivity to temperature.The low-temperature selections did not have a reduced optimum temperature but reduced sensitivity to temperature accounted for most of the ability to flower at low temperature. The parameters may vary independently. Examples are given to show how these parameters can be combined to construct genotypes with a required temperature response.     

The shriveling of pollinated pistils as an aid to rapid determination of Chrysanthemum pollen viability

Summary Shriveling of pistils observed one day after pollinating the dise florets of Chrysanthemum is normally followed by a good seed set 8 weeks later. When shriveling fails to appear, no seeds will be formed.It is suggested to use the easily visible pistil shriveling as a rapid in vivo test of pollen viability.     

The reaction of Chrysanthemum cultivars to Puccinia horiana and the inheritance of resistance

Summary Four types of reaction of Chrysanthemum morifolium to infection by Puccinia horiana are described. (1) Resistant plants show no macroscopic lesions. Cultivars classified as resistant carry a single dominant gene mostly in a simplex, sometimes in a duplex condition. (2). Incomplete resistance provides good protection but under extreme test conditions some pustules develop slowly. (3) Necrosis inhibits spore formation to a large extent. The progenies of crosses between necrotic and susceptible cultivars are mostly susceptible. (4) Susceptible plants sporulate abundantly.The merits of the three types of resistance are discussed.     

Winter environment increases self seed yield in self-incompatible (SI) Petunia hybrida clones

Summary SI inbreds of P. hybrida and crosses between self-incompatible (SI) petunia plants were brought to flower under winter and summer glasshouse conditions. SI response, as measured by self seed, ranged from zero seed set under both conditions to low or zero seed set during the summer and high seed set during the winter. Some plants produced comparable seed yields during either pollination time. Increased expression of pseudo-self-compatibility (PSC) during the winter months was attributed to a breakdown of the Si system by the low light conditions of Minnesota winters (45°N). Genotypic differences within and between populations in the winter provided differences facilitating selection for increased SI. These differences were masked in some populations by the summer environmental conditions. The use of simulated or natural low light conditions as a technique for selecting for SI should increase selection efficiency.Scientific Journal Series Paper Number 10,499 of the Minnesota Agricultural Experiment Station.     

Induced changes in growth and flowering of chrysanthemum after irradiation and in vitro culture of pedicels and petal epidermis

Summary To increase the genetic variation for yield in Chrysanthemum morifolium cv. White Spider, pedicel segments and petal epidermis were induced in vitro to regenerate adventitious buds either directly from the original tissue or indirectly via callus. Besides, pedicels were irradiated before in vitro culture with an X-ray dose of 8 Gy. All treatments yielded variation in flower morphology. Changes in flower colour were largely restricted to treatments involving irradiation.Treatments yielding many morphological variants also yielded most production variants. Plants regenerated from petals differed from those originating from pedicels. Clones were found that had retained the morphology of White Spider, but outperformed the controls in flowering time and flower number.     

Inheritance of flower color in Dendranthema grandiflora Tzvelev. (Chrysanthemum morifolium Ramat.) using cultivars and inbreds. I. Plastid pigmentation

Summary Progeny from controlled crosses of 28 garden chrysanthemum (Dendranthema grandiflora Tzvelev.) inbreds in various combinations as males with 3 heterozygous cultivars and 10 inbreds as females were evaluated for plastid pigmentation in a series of field and glasshouse environments. Disomic inheritance of a dominant gene, which acts to inhibit pigmentation, best explained the data. However, 1 of 8 families involving a glasshouse cultivar with colorless plastids (Puritan) as female and inbreds with yellow plastids as males deviated significantly from the expected 1:1 ratio. This family consistently exhibited an excess of plants with colorless plastids. Although parent clones and cloned progeny from inbred parents were scored the same in field and glasshouse environments, reevaluation in the field of all 8 Puritan families, first evaluated in a glasshouse environment, revealed phenotypic misclassification apparently due to environmental effects. An excess of plants with colorless plastids appeared in the glasshouse but evaluation of the same genotypes in the field resulted in an excess of plants having yellow plastids. Submerging ray florets in a hot-water bath revealed yellow plastids masked by vacuolar pigments, thus reducing chances of misclassification. The data suggest that evaluation of highly variable families in a single environment can lead to misclassification of plastid pigmentation. However, even with evaluation in 2 or more environments, certain families may exhibit an excess of plants with colorless plastids.     

Radiation-induced low-temperature tolerant solid mutants of Chrysanthemum morifolium Ram.

Summary From irradiated explants of the chrysanthemum cultivar Parliament solid non-chimeric low-temperature tolerant mutants were obtained. Some of the mutants flowered approx. 2 weeks earlier than control plants at a temperature of 12°C D/N and looked otherwise identical to the original genotype. The significance of it is discussed, a.o. in relation to attempts in developing selection methods at cell level for such characteristics.     

Application of RAPD markers in the characterisation of Chrysanthemum varieties and the assessment of somaclonal variation

Characterisation of fifteen commercial varieties of Chrysanthemum was carried out through RAPD analysis. Varieties could be distinguished from each other and the level of similarity between varieties seemed to be not very high. In vitro cultures were establish from four varieties and were subjected to different proliferation conditions. Five individuals from each variety and treatment were analysed using RAPD at the beginning of the treatment and after a month of culture. Variation was detected at both stages of the culture period. The rate of variation found showed differences between varieties, but no significant difference was found between culture conditions. This revised version was published online in August 2006 with corrections to the Cover Date.     

Effects of intergenotypic competition on selection within chrysanthemum progenies

Summary Intergenotypic competition affected the growth and onset of flowering of target plants of four chrysanthemum cultivars planted within plots of the same four cultivars at commercial spacing. The relative competitive abilities of two of these cultivars, Pollyanne and Hurricane, strong and weak respectively, were confirmed when they were further grown in balanced, equal-proportion mixtures. Notwithstanding their different competitive abilities, these two cultivars had similar flower yields in pure stands and, for this character at least, would be judged to have equal commercial worth.In a further series of experiments, target plants of the female parent were grown in pre-determined positions within clonally replicated progeny plots and in equivalent positions in pure stands of the female parent. These two types of plot gave similar proportions of total target plant variance due to position, suggesting that individual plants within a segregating progeny are subjected to similar levels of competition. It is inferred that the level of competition approximates to the overall mean competitive ability of the full-sibs comprising the progeny. Selection procedures are discussed in the light of this conclusion.     

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.     

Sequence of the S 9-RNase cDNA and PCR-RFLP system for discriminating S 1- to S 9-allele in Japanese pear

A new S ₉-allele was discovered in 6 Japanese pear cultivars, ‘Shinkou’, ‘Shinsei’, ‘Niitaka’, ‘Amanogawa’, ‘Nangetsu’ and ‘Nansui’. cDNA encoding S ₉-RNase, a stylar product of S ₉-allele, was cloned from pistils of ‘Shinkou’ and ‘Shinsei’ by 3' and 5' RACE. The S ₉-RNase gene had an open reading frame of 684 nucleotides encoding 228 amino acid residues. S ₉-RNase had a hypervariable (HV) region different from S ₁- to S ₈-RNase and shared higher similarity (95.2%) with apple S ₃-RNase than with 8 Japanese pear S-RNases (from 61.0% to 70.7%). Genomic PCR with primers ‘FTQQYQ’ and ‘anti-(I/T) IWPNV’ provided S ₁- to S ₉-amplicon (product), but could not discriminate the S ₂ from the S ₉ of ca. 1.3 kb. The S ₂ and S ₉ were distinguished by digestion with AflII and BstBI, respectively. The digestion with nine S-allele-specific restriction endonucleases, SfcI, AflII, PpuMI, NdeI,AlwNI, HincII, AccII, NruI and BstBI, distinguished S ₁ to S ₉, establishing that this PCR-RFLP system is useful for S-genotype assignments in Japanese pear harboring S ₁- to S ₉-allele. ‘Shinkou’, ‘Shinsei’, ‘Nangetsu’ and ‘Nansui’ assigned as S ₄ S ₉ were determined to be cross incompatible. This revised version was published online in July 2006 with corrections to the Cover Date.     

Chimerical structure and carotenoid inheritance in Chrysanthemum morifolium (Ramat.)

Summary Progeny analysis showed that yellow-flowered chrysanthemum sports with carotenoid only in L1 are indistinguishable from their white progenitors in breeding behaviour; those with carotenoid in L2 breed quite differently. Chimerical structure has, therefore, to be considered when analysing flower colour inheritance data. Segregations were consistent with the action of the single dominant gene I. Such simply inherited characters may be useful in studies to distinguish between the several alternative patterns of inheritance which can be suggested in the species.