The relationship between geographic distribution and ploidy level of taro,Colocasia esculenta

Summary The distribution of diploid and triploid domesticated and wild taro was investigated. It was found that going from south to north the frequency of diploid accession decreased, while that of triploids increased.     

Cytogenetic studies and the occurrence of triploidy in the wild potato species Solanum commersonii Dun.

Summary The South American wild potato species Solanum commersonii Dun. exists as two subspecies, subsp. commersonii and subsp. malmeanum (Bitter) Hawkes and Hjerting, and as both diploid and triploid cytotypes. The two subspecies were completely cross fertile but there were reciprocal differences in the number of seeds set per berry and the mean seed weight. Crosses between diploids and colchicine-induced autotetraploids yielded only a few seeds per berry, but 16 of the 18 germinated seeds were triploid or nearly so. Such a high proportion of triploids is unusual for such crosses in potatoes. Some triploids has a high percentage of stainable pollen. The origin of naturally occurring triploids is discussed in the context of 2n gametes, a triploid block, and endosperm balance numbers.     

Bilateral polyploidization inDactylis glomerata L. subsp.lusitanica: occurrence,morphological and genetic characteristics of first polyploids

Summary InDactylis glomerata L. subsp.lusitanica, triploid and tetraploid plants were obtained by bilateral sexual polyploidization in crosses between diploid parents known to produce 2n gametes. The polyploid and diploid progeny were compared for allozyme diversity (allele number and heterozygosity), phenological (pollen fertility, inflorescence emergence date), cellular (stomatic cell size) and morphological characters (vegetative biomass, seed weight, total seed number per plant, panicle number and seed number per panicle length) in four contrasting growing conditions. Stomatal cell size and the total number of alleles were significantly lower in diploids than in polyploids which sometimes recovered more than two distinct alleles at a locus via 2n gametes possessing heterozygous loci. Pollen fertility was similar in diploids and tetraploids, but significantly lower in the triploids. Seed weight was significantly higher in polyploids, although the average number of seeds per panicle length and the total number of seeds was higher in the diploid sibs. No phenological differences occurred among ploidy levels whereas such differences were observed inDactylis in all the natural areas of sympatry between diploids and their related polyploids. Our results provide evidence for an instantaneous effect of polyploidization on certain traits and suggest that natural selection would act subsequently on several other traits to allow the establishment of new polyploids.     

Crossability relationships of andean potato varieties of three ploidy levels

Summary The formation of triploids in 4x×2x crosses was studied in primitive potato varieties from the Andes of Peru and Bolivia. Triploid frequencies were as high as 80–100% in some crosses, although the actual number of seeds was low. The triploid block varied according to the tetraploid parent, and was never entirely absent. The likelihood of triploid formation was studied in 4x×2x crosses where pollen from both tetraploids and diploids of known genotype was used. Even though the diploids were chosen for their non-production of 2n gametes, as detected in pollen grain size, 66% of the hybrids were tetraploid. The amount and probability of gene-flow between the tetraploid and diploid gene-pools, either directly through 2n gametes or through triploid bridges was examined. Most of the 20 cultivars of S. x chaucha produced seed in crosses with tetraploids and diploids, although the fertility of the latter crosses was four times higher. The female fertility of some of the triploids thus allows them to exchange genes with the tetraploids and diploids, although gene-flow occurs more rapidly through direct transfer from diploids to tetraploids via 2n gametes.     

Aneuploids of perennial ryegrass (Lolium perenne)

Summary Aneuploid plants of perennial ryegrass (Lolium perenne L.) with 2n=15 to 30 chromosomes were obtained by crossing a near-triploid (2n=3x+1=22) with a diploid or on open-pollination with diploids and tetraploids. Aneuploids occurred with a frequency of 83% in near triploid × diploid progeny and 92% on open-pollination with diploid and tetraploid plants. Aneuploid plants with 15 to 18 chromosomes resembled diploids in morphology and those with 19 to 30 chromosomes were akin to tetraploids. Meiotic studies suggested that most aneuploid plants resulted from transmission of aneuploid egg cells (n=8 to 23). Aneuploid plants with 2n=27 to 30 chromosomes in the progeny of 22×14 cross originated from unreduced egg cells. Plants with 19 to 21 chromosomes were recovered only by immature seed culture. Aneuploid plants with 26 to 30 chromosomes and triploids (2n=21) had higher pollen fertility and bigger seeds than plants with 15 to 22 chromosomes.     

Male fertility and cytology of triploid hybrids between tetraploidSolanum commersonii (2n=4x=48, 2EBN) and Phureja-Tuberosum haploid hybrids (2n=2x=24, 2EBN)

Summary Solanum commersonii Dun. is a diploid (2n=2x=24, 1EBN) wild species of potential value for potato breeding. It is a reproductively isolated species and cannot be crossed with Tuberosum haploids (2n=2x=24, 2EBN) or other diploid 2EBNSolanum species. In order to overcome the EBN barriers, triploid hybrids were produced between Phureja-Tuberosum haploid hybrids, which form 2n pollen grains by parallel spindles, and tetraploidS. commersonii. Microsporogenesis analysis of the triploids indicated a trend towards low values of chromosome distribution at Anaphase I; lagging chromosomes were often observed as well. Despite these abnormalities, the percentage of stainable pollen was very high, ranging from 5.0% to 74.3%. A high variation in pollen grain diameter was also evident. Parallel and tripolar orientation of spindles at Metaphase II of microsporogenesis was a common feature of all the triploids analyzed, but dyads and triads were observed at a very low frequency. Therefore, also the frequency of 2n pollen was very low; the different size of stainable pollen appears to represent the ploidy levels which are possible according to the distribution of chromosomes in Anaphase I. The results obtained also suggest thatS. commersonii could have minor genes acting at the end of meiosis in such a way that, despite the presence of parallel/tripolar spindles, dyads/triads are not formed.Contribution no. 124 from the Research Center for Vegetable Breeding.     

Intergeneric Hybrids between Aegilops squarrosa and Secale cereale and their Meiotic Chromosome Behaviour

Three hybrid plants with different combinations of D and R genomes were produced from crosses between Aegilops squarrosa (2×: and 4×) and Secale cereale (2× and 4×) using embryo culture. Production frequencies of mature hybrids, having the genomes DR, DDR and DDRR, were 2.0%, 5.2% and 2.2 %, respectively, of florets pollinated. Amphidiploids were obtained directly from the cross between tetraploid parents. The majority of their morphological features were intermediate between those of the parents but, as with rye, the rachis was tough. Diploid and triploid hybrids were completely seed-sterile, whereas the amphidiploid had an average self-fertility of 4.5 % with a range of 0 to 45 %. The somatic chromosome numbers of diploids, triploids and amphidiploids were 2n = 14, 21 and 28, respectively. At meiotic metaphase I, the mean chromosome associations were 0.01 III + 0.26 II + 13.4 I in diploids, 6.96 II + 7.1 I in triploids, and 10.5 II + 7.9 I in amphidiploids. In diploids, pairing between D and R chromosomes was observed in the form of end-to-end types without chiasmata. Homologous pairing between D chromosomes was predominant in plants having two sets of D genomes. Amphidiploids showed a diploid-like chromosome behaviour. Homoeologous pairing was not observed in either triploids or amphidiploids.     

Modes of inheritance of two apomixis components,diplospory and parthenogenesis,in Chinese chive (Allium ramosum) revealed by analysis of the segregating population generated by back-crossing between amphimictic and apomictic diploids

To investigate the mode of inheritance of apomixis in Chinese chive, the degrees of diplospory and parthenogenesis were evaluated in F₁ and BC₁ progenies derived from crosses between amphimictic and apomictic diploids (2n = 16, 2x). The F₁ population was generated by crossing three amphimictic diploids 94Mo13, 94Mo49 and 94Mo50 with an apomictic diploid KaD2 and comprised 110 diploids and 773 triploids. All the diploid F₁ plants examined were completely or highly eusporous and completely syngamic. All the triploid F₁ plants examined were highly diplosporous and highly parthenogenetic. KaD2 could not transmit its high level of apomixis via monoploid pollen grains. The BC₁ population, generated by crossing 94Mo49 with apomictic triploids found in the F₁ offspring, exhibited heteroploidy; it comprised haploid, diploid, triploid, tetraploid and various aneuploid individuals. In this generation, clear segregation was observed between diplospory and parthenogenesis. Analysis of the BC₁ population suggests that diplospory and parthenogenesis are each controlled by single dominant genes, D and P, respectively. However, all the BC₁ plants characterized as parthenogenetic were diplosporous. The absence of phenotypically eusporous parthenogenetic plants can be explained by assuming that the presence of diplospory gene is a prerequisite for the parthenogenesis gene expression in Chinese chive.     

Note on the probable cause of the occurrence of tetraploid plants in commercial triploid varieties of sugar beet

In crossing diploid male-sterile beets (18 chromosomes) with a tetraploid pollinator (36 chromosomes) it was established that of 532 individuals from this cross 8.4% were diploid, 83.9% triploid and 7.7% tetraploid.The most reasonable explanation was that the diploid male-sterile plants had given a number of unreduced egg cells. This could be put to a test by crossing diploid male-sterile plants with normal diploids. The average number of triploids found by chromosome counts was 2.4%; in a normal diploid no triploids were found.The type of cytoplasm associated with the male-sterility may lead to abnormal egg cells.Trirave (Triplex) with 84% of triploids, represents an enormous advance over normal polyploid types, which very commonly have no more than 40 per cent triploids and very rarely have more than 50 per cent.Bush Johnsons Limited and D.J. van der Have n.v.Paper read on the XXIInd Winter Meeting of the Institut International de Recherches Betteravières on February 24th, 1959.     

Polyploidy in the glasshouse cucumber (Cucumis sativus L.)

Summary An autotetraploid form of the parthenocarpic variety Butchers Disease Resisting was produced by colchicine treatment. Immersing the seeds in water before treatment in 0.2% colchicine for 24 hours gave the highest percentage of vigorous tetraploids. The yield of fruit from tetraploids was better than the diploids at first but it declined later in the cropping period owing to slow sideshoot production.Self-fertility of autotetraploids was lower than diploids. When crossed with diploids, triploid plants were obtained. These were self-sterile but they could be crossed with diploids to give trisomic plants. Aneuploids with one additional chromosome were found amongst tetraploids and triploids.     

Breeding of polyploid spinach

2x, 3x and 4x forms of spinach were compared in 1959, 1960 and 1961.At the same stage of development the tetraploids outyielded the diploids in total green matter and leaf blades in 1959 and 1961. In 1960 there was no significant difference. This year the tetraploids had a very small increase in green matter in the period between the two harvest dates. Yield of dry matter was determined in 1960 and 1961, and in both years the tetraploids outyielded the diploids. The rate of development decreased with increasing degree of ploidy. Compared at the same age, when the diploids were about to start bolting they were superior to the polyploids with regard to total yield, yield of leaf blades and yield of dry matter.The onset of flowering was considerably delayed in the polyploids. Generally the triploids were intermediate between the diploids and the tetraploids. The triploids and tetraploids had a larger proportion of monoecious plants, and a smaller proportion of males than the diploids.Report No. 27     

Nematode resistance in bananas: screening results on some wild and cultivated accessions of Musa spp.

Bananas cultivated for export all belong to Cavendish cultivars and are all recognized as very susceptible to nematodes, particularly to the burrowing nematode Radopholus similis and the lesion nematode Pratylenchus coffeae. Even if there have been many changes in the management of banana nematodes in large commercial banana plantations, chemical control still remains most often the last resort method to manage the nematodes, although the number of registered products is definitely declining. Therefore, nematode control though genetic improvement is gaining new interest worldwide. In this study, 55 banana accessions mostly diploids from the Musa acuminata genome group (AA) but including some triploid accessions (AAA), some diploids of the Musa balbisiana genome group (BB) and some interspecific hybrids (AAB, AB) were evaluated for resistance to four nematode species R. similis, P. coffeae, Meloidogyne incognita and M. arenaria. These experiments were conducted in a growth chamber under controlled conditions. All banana accessions were susceptible to nematode species, although many different levels of susceptibility were detected. This study confirmed the good resistance status to R. similis of some cultivars from the Pisang jari buaya and Pisang batuau subgroups and the partial resistance of 17 diploid accessions significantly different from the susceptible reference cv. Grande Naine. This study also showed that 12 diploid accessions exhibited a partial resistance to P. coffeae, including some usual or potential genitors belonging to the wild diploids subspecies burmannica (cvs. Long Tavoy 1 and 2) and burmannicoides (cv. Calcutta 4). No source of resistance to Meloidogyne spp. was found. These screening results, combining for the first time four nematode species, are discussed within the scope of banana breeding in order to produce parental diploid lines with single or combined nematode resistances and further develop triploids that can substitute existing susceptible commercial cultivars.     

Polyploidy and sexual polyploidization in the genus Vaccinium

Genus Vaccinium,consisting of blueberries, cranberries, lingonberries and many related wild species, includes diploid, tetraploid, and hexaploid species. Most evidence indicates that the tetraploid species are autotetraploids, with non-preferential bivalent chromosome pairing. Although homoploid interspecific crosses usually produce numerous fertile hybrids if the parents are from the same section of the genus, inter sectional crosses at the diploid level normally produce no seedlings, weak seedlings, or seedlings that are very low in fertility. There is a strong but not complete triploid block within Vaccinium. Even with insections, tetraploid × diploid (and the reciprocal) crosses normally give only a few tetraploid and a few triploid hybrids. Hexaploid × diploid crosses within sections are very hard to make, and the few hybrids that have been obtained are pentaploid. The frequency of 2n gametes varies,both among genotypes within species and among species. Vaccinium pollen is normally shed in tetrads, and the frequency of large pollen grains shed in dyads can be used to estimate 2n gamete frequency. Cultivated blueberries occur at both the tetraploid and the hexaploid levels, and there are important genetic resources in the diploids. Unreduced gamete production has permitted transfer of genetic material from the diploid to the tetraploid level and from the diploids and tetraploids to the hexaploid level via triploid hybrids. Intersectional crosses can occasionally produce tetraploid Vaccinium hybrids that appear to behave as amphidiploids and have medium to high fertility. CommercialVaccinium cultivars are normally propagated by cuttings. Intersectional hybridization, chromosome doubling, and asexual propagation could permit the production of novel hybrid combinations with value as ornamentals or in fruit production. This revised version was published online in July 2006 with corrections to the Cover Date.     

Geographical differentiation of Asian taro, Colacasia esculenta (L.) Schott, detected by RAPD and isozyme analyses

Geographical differentiation and phylogenetic relationships of Asian taros, Colocasia esculenta (L.) Schott, and related species were analyzed by random amplified polymorphic DNA (RAPD) and isozymes of 13 enzyme systems with special interest in the accessions from the Yunnan area of China, which supposedly has served the secondary center of taro diversification and dispersal into the temperate Far East Asia. The RAPD analysis was found to be better suited for detecting genetic differences within taros and among its related species. However, both RAPD and isozyme analyses estimated quite similar genetic relationships within taros and between related species. Genetic differentiation was evident in the taro accessions of Nepal, Yunnan and other Asian areas; but, phylogenetic relationships between the differentiated taro groups were not clearly determined. When taro cultivation was introduced to a new area, only a small fraction of genetic variability in heterogeneous taro populations was transferred possibly causing random differentiation among locally adapted taro populations. Some of the Yunnan and Japanese accessions were found to be direct descendants of the common triploid population. Further analysis on taros from the eastern China and Korea is necessary to clarify the Yunnan's role in taro diversification and dispersal. The significant local differentiation in Asian taros was clearly demonstrated by RAPD and isozyme analyses in this study, and the results of this study will serve as a base to establish evolutionary and genetic relationships among Asian taros. This revised version was published online in August 2006 with corrections to the Cover Date.     

Polyploidy in tulips (Tulipa L.). The occurrence of diploid gametes

Summary From data available in the literature and supplemented by information from private tulip breeders, it appeared that triploid and tetraploid varieties raised in the past, resulted from crosses between diploid tulips and between diploids and tetraploids. This can only be explained by the occurrence of diploid gametes in diploid (2n=24) cultivars. This phenomenon which may affect both female and male gametes, appears to be specific for certain varieties. Consequences for practical breeding are discussed.     

Genotypic variability in vegetable amaranth (Amaranthus tricolor L for foliage yield and its contributing traits over successive cuttings and years

Summary Twenty nine strains of vegetable amaranth (Amaranthus tricolor were grown for two successive seasons to study different selection parameters for foliage yield and its nine contributing morphological and quality traits. The strains AV-38 (5.06 kg/plot) and AV-31 (5.04 kg/plot) recorded highest foliage yield, followed by AV-30 (4.78 kg/plot) and AV-23 (4.70 kg/plot). The protein and carotenoid content averaged 1.24 ± 0.03 mg/100 mg and 0.83 ± 0.02 mg/g respectively. The leaves of A. tricolor also have considerable quantities of ascorbic acid (112.33 ± 5.00 mg/100 g) and fibre (8.39 ± 0.10%). The mean of individual cuttings for plant height, leaf size, stem diameter, foliage yield, protein, ascorbic acid and fibre content increased with successive cuttings till third cutting and thereafter showed a decline. Genotypic coefficient of variation (GCV) values ranged from 6.80 to 28.25%. However, the fibre content, branches/plant, leaves/plant, plant height and stem diameter showed lowest values of GCV. The values of heritability estimates were high for all the traits in all the cuttings as well as on pooled basis and ranged from 0.89 for branches/plant to 0.98 for foliage yield. Highest expected genetic advance was noticed for ascorbic acid (57.48%), followed by foliage yield (48.30%) and leaf size (29.51%).     

Production of intraspecific aneuploids in the genus Solanum. 1. Doubling the chromosome number of seven diploid species

Summary The variation which persisted in interspecific aneuploids even after one or two generations of backcrossing prevented the morphological distinction of the aneuploids from their diploid sibs. An attempt was made to unmask the effects of aneuploidy by producing aneuploids within the more uniform genetic background of a single species.The procedure to produce trisomics and aneuploids of diploid species was to vegetatively double clones or seed of the species with colchicine, cross the induced tetraploids to diploids to produce triploids, and then cross the triploids to the original diploids. An attempt was made to use the same diploid clone and/or introduction of a species in all successive stages of the procedure in order to obtain as much uniformity as possible. This was largely unsuccessful because self-incompatibility and inbreeding reduced seed set. Seventy-three different tetraploids were obtained from seven species. Tuber treatments were noticeably unsuccessful in producing doubled plants. The two vegetative treatments were most successful and produced 41 different tetraploid clones while 30 doubled plants were obtained from four seed treatments.Morphologically the tetraploids resembled the diploid and there was no noticeable increase in size of leaflets and floral parts in the tetraploid. The average number of chloroplasts in ten pairs of guard cells was a distinctive feature which could efficiently be used in distinguishing the induced tetraploid plants from diploids. The tetraploids were generally less fertile than their diploid counterparts and the majority of them exhibited self-compatibility in comparison to self incompatibility of the diploids.Cooperative investigations of the Agricultural Research Service, U.S. Department of Agriculture and the Wisconsin Agricultural Experiment Station. Supported in part by grants from the Research Committee of the Graduate School, University of Wisconsin.     

埃塞俄比亚芥与白菜型油菜间六倍体杂种的获得及其生物学特性研究

通过杂交和染色体加倍人工合成了埃塞俄比亚芥×白菜型油菜间的三倍体和六倍体杂种，并对其进行了细胞学鉴定、气孔观察以及一些生理生化特性的分析。在染色体记数、原位杂交和分子标记印证的基础上，比较了两种不同染色体加倍方法的效率；生理生化分析显示，六倍体杂种植株叶片的游离脯氨酸含量、质膜相对透性、Vc含量和叶绿素含量与三倍体杂种植株相比都有明显的差别。     

The enigma of triploid potatoes

Triploids are comparatively difficult to produce in tuberous Solanums as is the case in many other genera. Amongst the wild species triploids are infrequent whereas in the cultivars the opposite is true. It is argued that their survival and increase under cultivation means that they possess certain selective advantages as cultivars, advantages often concomitant with triploidy per se. Triploids make superior cultivars in a number of crops: by analogy therefore, it is suggested that the relative merits of triploid cultivated potatoes warrants investigation.     

Identification of triploid Citrus by isozyme analysis

Summary Seedlessness is a desirable horticultural attribute in Citrus and is positively associated with triploidy. The conventional cytological method for triploid identification is a laborious technique involving the preparation of root tips for chromosomal analysis. Digital densitometry of isozymes, however, offers the possibility of distinguishing triploid Citrus from large populations of seedlings both quickly and cheaply. Where there are no gene dosage regulation effects, greater band density should be evident in the allozyme contributed by the diploid gamete for a heterozygous locus. The isozymes of 4 enzymes; malate dehydrogenase, 6-phosphogluconate dehydrogenase, shikimate dehydrogenase, and phosphoglucose isomerase, were investigated with polyacrylamide gel electrophoresis. Band densities of these isozymes for triploid Citrus, their diploid siblings and diploid progenitors were measured using a digital densitometer. Of the 4 enzymes investigated only allozymes for shikimate dehydrogenase exhibited consistent differences over a wide range of Citrus cultivars. Greater band density was evident in the allozyme contributed by the diploid gamete. The band density ratio between allozymes for triploid Citrus was close to 0.5, while for diploid Citrus band density ratios were close to 1.0. This effect is due to the extra protein coded by the additional gene dose and was not observed in diploids. Shikimate dehydrogenase proved to be an accurate molecular marker for distinguishing between diploid and triploid Citrus for heterozygous progeny.Abbreviations PAGE Polyacrylamide gel electrophoresis - MDH malate dehydrogenase - 6PGD 6-phosphogluconate dehydrogenase - SkDH shikimate dehydrogenase - PGI phosphoglucose isomerase