Colchicine induced manifestation of abnormal male meiosis and 2n pollen in Trachyspermum ammi (L.) Sprague (Apiaceae)

Unreduced gametes are the key source for the natural polyploidization in plants, but process of its formation is very low in nature. Meiotic mutants are second source for the formation of 2n pollen. In this cytological investigation, the meiotic aberrations and its impact on post-meiotic products were analysed in autotetraploid Trachyspermum ammi (L.) Sprague (4n=36). The seedlings of T. ammi (L.) Sprague were treated with 3 different concentrations of colchicine (0.2, 0.4 and 0.5%, w/v) for 3 different durations. Six polyploid plants were induced which was confirmed on the basis of cytological analysis. Colchicine, an anti-microtubular drug induced different meiotic and post-meiotic abnormalities such as chromosomal bridges, lagging chromosomes, scattering, precocious, fragments, dyads, triads, and polyads. The formation of several abnormal sporads clearly signifies the meiotic restitution. The tendency of univalents to scattered in the cytoplasm at metaphase was identified as a peculiar aberration asynapsis. Pollen variability and fusion of pollen walls was reported and pollen fertility was calculated. The morphological analysis of the pollen allowed us to confirm the occurrence of 2n pollen.

Meiotic Aberrations Leading To Aneuploidy In Cauliflower (Brassica Oleracea L. var. Botrytis)

Aneuploid cauliflower plants (Brassica oleracea L. var. botrytis) display abnormal curd phenotypes causing serious commercial problems in offspring populations. Despite extensive breeding efforts, selection of genotypes producing euploid gametes remains unsuccessful due to unknown genetic and environmental factors. To reveal the origin of aneuploid gametes, we analyzed chromosome pairing, chiasma formation and chromosome segregation in pollen mother cells of selected cauliflower genotypes. To this end we compared different genotypes exhibiting Low with < 5%, Moderate with 5-10% and High with > 10% aberrant offspring. Microscopic observations revealed regular chromosome pairing at pachytene. However, cells at diakinesis and metaphase I showed variable numbers of univalents, suggesting that chiasma formation during meiotic prophase is incomplete or disrupted and results in a partial desynaptic phenotype. Cells at anaphase I – telophase II exhibited various degrees of unbalanced chromosome numbers explaining the aneuploid offspring. Immunofluorescence probed with an MLH1 antibody demonstrated fluorescent foci in all genotypes, but their lower numbers do not correspond to the putative sites of chiasmata. Interchromosomal connections between chromosomes and bivalents are common at diakinesis and metaphase I, and they contain centromeric and 45S rDNA tandem repeats, but such threads seemed not to affect proper disjoin of the half bivalents at anaphase I. Moreover, male meiosis in the arabidopsis APETALA1/ CAULIFLOWER double mutant with the typical cauliflower phenotype did show interchromosomal connections, but there were no indications for partial desynapsis. We now hypothesize that the occurrence of desynapsis in cauliflower is a developmental out-of-phase phenomenon partially or completely controlled by genes involved in flower and curd development.

Sex chromosomes in meiotic, hemiclonal, clonal and polyploid hybrid vertebrates: along the ‘extended speciation continuum'

We review knowledge about the roles of sex chromosomes in vertebrate hybridization and speciation, exploring a gradient of divergences with increasing reproductive isolation (speciation continuum). Under early divergence, well-differentiated sex chromosomes in meiotic hybrids may cause Haldane-effects and introgress less easily than autosomes. Undifferentiated sex chromosomes are more susceptible to introgression and form multiple (or new) sex chromosome systems with hardly predictable dominance hierarchies. Under increased divergence, most vertebrates reach complete intrinsic reproductive isolation. Slightly earlier, some hybrids (linked in ‘the extended speciation continuum') exhibit aberrant gametogenesis, leading towards female clonality. This facilitates the evolution of various allodiploid and allopolyploid clonal (‘asexual’) hybrid vertebrates, where ‘asexuality' might be a form of intrinsic reproductive isolation. A comprehensive list of ‘asexual' hybrid vertebrates shows that they all evolved from parents with divergences that were greater than at the intraspecific level (K2P-distances of greater than 5–22% based on mtDNA). These ‘asexual' taxa inherited genetic sex determination by mostly undifferentiated sex chromosomes. Among the few known sex-determining systems in hybrid ‘asexuals', female heterogamety (ZW) occurred about twice as often as male heterogamety (XY). We hypothesize that pre-/meiotic aberrations in all-female ZW-hybrids present Haldane-effects promoting their evolution. Understanding the preconditions to produce various clonal or meiotic allopolyploids appears crucial for insights into the evolution of sex, ‘asexuality' and polyploidy. This article is part of the theme issue ‘Challenging the paradigm in sex chromosome evolution: empirical and theoretical insights with a focus on vertebrates (Part II)’.