Plants Of Different Ploidy Levels Research Articles

A comparative study on morphological and fruit quality traits of diploid and polyploid carambola (Averrhoa carambola L.) genotypes

Artificially induced polyploid crops are generally associated with improved agronomic characteristics. Triploids and tetraploids previously created from the diploid carambola (Averrhoa carambola L.) cultivar 'B17′ (2n = 22) were used as experimental materials. The plody level of this new genetic material was checked by the combination of flow cytometry and by chromosome counting. The morphological characteristics and fruit quality of diploid and polyploid carambola were evaluated under field conditions. The results revealed that plants of different ploidy levels demonstrated significantly different phenotypic traits. Compared with their diploid counterparts, polyploids in general had greater vigor and biomass. Triploid and tetraploid plants produced stronger branches and petioles, more leaflets, thicker and larger leaves, larger pollen grains and flowers, bigger and heavier fruits as compared with that of diploid plants. However, a great decrease in the number of flowers was observed in polyploid plants, with 50 % reduction in the triploids as compared with the diploids. Furthermore, the majority of pollen from triploid plants was abnormal with collapsed structure and two different types of pollen grains were observed in the tetraploids using scanning electron microscope. Additionally, a general improvement in the quality of polyploid fruits was found when compared with diploid ones. In particular, the shelf life of polyploid fruits was much longer than their diploid counterparts. A comparative study was conducted among parental diploid carambola plants and their triploid and tetraploid deliverable offsprings based on morphological, physiological and agronomical traits. This study is expected to contribute into the breeding process for the production of new polyploid carambola varieties which will be characterized by special agronomical characteristics, better quality and longer self life period of their fruits.

Morphology mirrors ploidy and reproductive modes in Pilosella officinarum

Pilosella officinarum is represented predominantly by tetraploid (2n = 36), pentaploid (2n = 45) and hexaploid (2n = 54) cytotypes reproducing, to various degrees, both sexually and apomictically. Its current intraspecific taxonomical treatment is based mainly on selected apomictic lineages, the large number of which makes the treatment confusing and not generally applicable. We therefore tested the breeding modes of a representative set of plants from central Europe, cultivated under experimental conditions. Each ploidy level was associated with a different reproductive pattern (4x – sexual, 5x – prevalently apomictic and 6x – sexual and apomictic). Whereas sexual tetraploids occurred in the western part of the study area (the Czech Massif), apomictic pentaploids and hexaploids were scattered in its eastern part (Western Carpathians and Pannonia). Moreover, the hexaploid cytotype formed a distinct exclusively sexual group restricted to steep river canyons of the Czech Massif. Morphometric analyses were performed to determine the set of characters which distinguish major lineages characterized by different ploidy and reproductive modes. Their results confirm the existence of morphological differences between plants of different ploidy levels and, in the case of the hexaploid cytotype, different modes of breeding. Knowledge of ploidy and reproductive modes is therefore essential for elucidating the reticulate infraspecific structure of Pilosella officinarum.

Transgenerational effects of inter-ploidy cross direction on reproduction and F2 seed development of Arabidopsis thaliana F1 hybrid triploids

Key messageReproduction in triploid plants is important for understanding polyploid population dynamics. We show that genetically identical reciprocal F1 hybrid triploids can display transgenerational epigenetic effects on viable F2 seed development.The success or failure of reproductive outcomes from intra-species crosses between plants of different ploidy levels is an important factor in flowering plant evolution and crop breeding. However, the effects of inter-ploidy cross directions on F1 hybrid offspring fitness are poorly understood. In Arabidopsis thaliana, hybridization between diploid and tetraploid plants can produce viable F1 triploid plants. When selfed, such F1 triploid plants act as aneuploid gamete production “machines” where the vast majority of gametes generated are aneuploid which, following sexual reproduction, can generate aneuploid swarms of F2 progeny (Henry et al. 2009). There is potential for some aneuploids to cause gametophyte abortion and/or F2 seed abortion (Henry et al. 2009). In this study, we analyse the reproductive success of 178 self-fertilized inter-accession F1 hybrid triploids and demonstrate that the proportions of aborted or normally developed F2 seeds from the selfed F1 triploids depend upon a combination of natural variation and cross direction, with strong interaction between these factors. Single-seed ploidy analysis indicates that the embryonic DNA content of phenotypically normal F2 seeds is highly variable and that these DNA content distributions are also affected by genotype and cross direction. Notably, genetically identical reciprocal F1 hybrid triploids display grandparent-of-origin effects on F2 seed set, and hence on the ability to tolerate aneuploidy in F2 seed. There are differences between reciprocal F1 hybrid triploids regarding the proportions of normal and aborted F2 seeds generated, and also for the DNA content averages and distributions of the F2 seeds. To identify genetic variation for tolerance of aneuploidy in F2 seeds, we carried out a GWAS which identified two SNPs, termed MOT and POT, which represent candidate loci for genetic control of the proportion of normal F2 seeds obtained from selfed F1 triploids. Parental and grandparental effects on F2 seeds obtained from selfed F1 triploids can have transgenerational consequences for asymmetric gene flow, emergence of novel genotypes in polyploid populations, and for control of F2 seed set in triploid crops.

Postglacial colonization history reflects in the genetic structure of natural populations of Festuca rubra in Europe.

We conducted a large‐scale population genetic survey of genetic diversity of the host grass Festuca rubra s.l., which fitness can be highly dependent on its symbiotic fungus Epichloë festucae, to evaluate genetic variation and population structure across the European range. The 27 studied populations have previously been found to differ in frequencies of occurrence of the symbiotic fungus E. festucae and ploidy levels. As predicted, we found decreased genetic diversity in previously glaciated areas in comparison with nonglaciated regions and discovered three major maternal genetic groups: southern, northeastern, and northwestern Europe. Interestingly, host populations from Greenland were genetically similar to those from the Faroe Islands and Iceland, suggesting gene flow also between those areas. The level of variation among populations within regions is evidently highly dependent on the postglacial colonization history, in particular on the number of independent long‐distance seed colonization events. Yet, also anthropogenic effects may have affected the population structure in F. rubra. We did not observe higher fungal infection rates in grass populations with lower levels of genetic variability. In fact, the fungal infection rates of E. festucae in relation to genetic variability of the host populations varied widely among geographical areas, which indicate differences in population histories due to colonization events and possible costs of systemic fungi in harsh environmental conditions. We found that the plants of different ploidy levels are genetically closely related within geographic areas indicating independent formation of polyploids in different maternal lineages.

Root sprouting in Knautia arvensis (Dipsacaceae): effects of polyploidy, soil origin and nutrient availability

Plants able to resprout from roots have a potential bud bank that gets initiated after injury to overcome meristem limitation after loss of all stem parts and to facilitate regeneration. Knautia arvensis is reportedly able to sprout from its roots on arable land, but information is missing regarding such ability in serpentine populations or how it might differ between diploids and tetraploids. We hypothesized that (1) 'ruderal' non-serpentine populations better tolerate severe disturbance than relic, serpentine ones; (2) tetraploid populations resprout more readily than diploids due to enhanced growth of higher ploidy levels; and (3) plants of different ploidy levels from serpentine soils are, for evolutionary reasons, more similar in their response to disturbances than plants from non-serpentine soils. To test these hypotheses, we conducted a pot experiment. Its results do not support our hypothesis that the ability to sprout from roots is a factor driving the spread of new weedy taxa into central Europe or the hypothesis that it is related to polyploidization in the genus Knautia. Both tetraploids and plants from non-serpentine populations regenerated less vigorously than diploids and plants from serpentine populations. However, the genetically closer populations of serpentine origin were more similar in their response to experimental manipula- tions than their genetically distinct non-serpentine counterparts. The success of non-serpentine taxa in disturbed habitats of central Europe might be related to traits other than the ability to resprout.

Polyploidy enhances the occupation of heterogeneous environments through hydraulic related trade‐offs inAtriplex canescens(Chenopodiaceae)

Plant hydraulic characteristics were studied in diploid, tetraploid and hexaploid cytotypes of Atriplex canescens (Chenopodiaceae) to investigate the potential physiological basis underlying the intraspecific habitat differentiation among plants of different ploidy levels. Populations of A.canescens from different habitats of the Chihuahuan Desert (New Mexico, USA) were analyzed using flow cytometry to determine ploidy levels. Traits related to xylem water transport efficiency and safety against drought-induced hydraulic failure were measured in both stems and leaves. At the stem level, cytotypes of higher ploidy showed consistently lower leaf-specific hydraulic conductivity but greater resistance to drought-induced loss of hydraulic conductivity. At the leaf level, comparisons in hydraulics between cytotypes did not show a consistent pattern, but exhibited high plasticity to proximal environmental conditions related to soil water availability. The results suggest that a trade-off between stem hydraulic efficiency and safety across ploidy levels underlies niche differentiation among different cytotypes of A.canescens. Polyploidization may have been facilitated by environmental heterogeneity related to water availability, and variation in water-related physiology found in the present study suggests an important functional basis for the niche differentiation and coexistence of A.canescens cytotypes in desert environments.

Hybridization between diploidCentaurea pseudophrygiaand tetraploidC. jacea(Asteraceae): the role of mixed pollination, unreduced gametes, and mentor effects

We studied hybridization between the diploid Centaurea pseudophrygia and the tetraploid C. jacea by performing crossing experiments and screening natural populations using flow cytometry. The experiments confirm that the studied species exhibit strong reproductive isolation. Interspecific hybrids were formed at a low frequency, including triploids (originating from reduced gametes) and tetraploids (involving unreduced gametes of the diploids). In contrast, hybrids were almost absent among seeds and adult plants of natural mixed populations and among the offspring from experimental pollinations with a mixture of pollen of both ploidy levels. We found that mixed pollination is an important mechanism for preventing hybridization between plants of different ploidy levels and sustaining the reproduction of the tetraploids. A mentor effect (induced selfing in the presence of pollen of different ploidy levels) was observed in both diploids and tetraploids, reinforcing the reproductive isolation between cytotypes. Higher ploidy levels (pentaploid, hexaploid) involving unreduced gametes of the tetraploid species were identified. Notably, pentaploids were discovered for the first time in Centaurea sect. Jacea. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104, 93–106.

Complex distribution patterns, ecology and coexistence of ploidy levels of Allium oleraceum (Alliaceae) in the Czech Republic.

Despite extensive study of polyploidy, its origin, and ecogeographical differences between polyploids and their diploid progenitors, few studies have addressed ploidy-level structure and patterns of ecogeographical differentiation at various spatial scales using detailed sampling procedures. The pattern of coexistence of polyploids in the geophyte Allium oleraceum at the landscape and locality scale and their ecology were studied. Flow cytometry and root-tip squashes were used to identify the ploidy level of 4347 plants from 325 populations sampled from the Czech Republic using a stratified random sampling procedure. Ecological differentiation among ploidy levels was tested by comparing sets of environmental variables recorded at each locality. Across the entire sampling area, pentaploids (2n = 5x = 40) predominated, while hexaploids (2n = 6x = 48) and tetraploids (2n = 4x = 32) were less frequent. The distribution of tetra- and hexaploids was partially sympatric (in the eastern part) to parapatric (in the western part of the Czech Republic) whereas pentaploids were sympatric with other cytotypes. Plants of different ploidy levels were found to be ecologically differentiated and the ruderal character of cytotypes increased in the direction 4x --> 5x --> 6x with the largest realized niche differences between tetra- and hexaploids. Most populations contained only one ploidy level (77 %), 22 % had two (all possible combinations) and 1 % were composed of three ploidy levels. The majority of 4x + 5x and 5x + 6x mixed populations occurred in sympatry with uniform populations of the participating cytotypes in sites with ecologically heterogeneous or marginal environment, suggesting secondary contact between cytotypes. Some mixed 4x + 6x populations dominated by tetraploids being sympatric and intermixed with uniform 4x populations might represent primary zones of cytotype contact. Almost no mixed accessions were observed on the fine spatial scale in mixed populations. The results provide evidence for adaptive differences among ploidy levels, which may contribute to their complex distribution pattern. The prevalence of asexual reproduction, limited dispersal and equilibrium-disrupting processes may support local coexistence of cytotypes.

In vitro induction of tetraploids in crape myrtle (Lagerstroemia indica L.)

A protocol for in vitro induction of crape myrtle tetraploids using nodes from in vitro-grown shoots (2n = 48) was established. Nodal buds were excised from in vitro-grown shoots, maintained on proliferation medium containing Murashige and Skoog medium supplemented with 4.44 μM 6-benzyladenine , 0.54 μM α-naphthaleneacetic acid, and treated with a range of concentrations of colchicine under three different conditions. Nodal bud explants treated in liquid proliferation medium supplemented with either 15 or 20 mM colchicine for 24 h turned necrotic and died; whereas, those cultured on solid proliferation medium supplemented with either 125 or 250 μM colchicine for 30 days survived, but no tetraploid plants were obtained. However, when explants were cultured in liquid proliferation medium containing 250, 500 or 750 μM colchicine for 10 days, tetraploid plants (2n = 96) were obtained. Incubation of explants in medium containing 750 μM colchicine promoted the highest frequency of survival (40%) of explants and of recovered tetraploids (60%). Morphological and anatomical characteristics of leaves, including leaf index, stomata size and number, stomata index (length/width), and number of chloroplasts in guard cells correlated with ploidy of crape myrtle plants. The number of chloroplasts in guard cells of stomata was a stable and reliable marker in discriminating plants of different ploidy levels. Chromosome counts and flow cytometry confirmed these findings.

Population dynamics of diploid and hexaploid populations of a perennial herb.

Despite the recent enormous increase in the number of studies on polyploid species, no studies to date have explored the population dynamics of these taxa. It is thus not known whether the commonly reported differences in single life-history traits between taxa of different ploidy levels result in differences in population dynamics. This study explores differences in single life-history traits and in the complete life cycle between populations of different ploidy levels and compares these differences with differences observed between different habitat types and years. Diploid and hexaploid populations of a perennial herb, Aster amellus, are used as the study system. Transition matrix models were used to describe the dynamics of the populations, and population growth rates, elasticity values and life-table response experiments were used to compare the dynamics between populations and years. The results indicate that between-year variation in population dynamics is much larger than variation between different ploidy levels and different habitat conditions. Significant differences exist, however, in the structure of the transition matrices, indicating that the dynamics of the different ploidy levels are different. Strong differences in probability of extinction of local populations were also found, with hexaploid populations having higher probability than diploid populations, indicating strong potential differences in persistence of these populations. This is the first study on complete population dynamics of plants of different ploidy levels. This knowledge will help to understand the ability of new ploidy levels to spread into new areas and persist there, and the interactions of different ploidy levels in secondary contact zones. This knowledge will also contribute to understanding of interactions of different ploidy levels with other plant species or other interacting organisms such as pollinators or herbivores.

Possible pathways of the gene flow inTaraxacum sect.Ruderalia

Reproductive behaviour and the pathways of gene flow among ploidy levels were studied experimentally inTaraxacum sect.Ruderalia. Diploid, triploid and tetraploid individuals were sampled from mixed diploid — polyploid natural populations. 136 experimental hybridizations between the plants of different ploidy levels were performed. Seeds resulting from these crosses, those obtained from isolated anthodia as well as from open pollinated anthodia (both from cultivated and wild plants) were subjected to the flow-cytometric seed screening (FCSS) to determine ploidy levels in the progeny and to infer breeding behaviour of maternal plants. Three possible pathways of the gene flow were studied: (A) fertilization of sexuals by pollen of apomicts, (B) BIII hybrid formation, (C) facultative apomixis. Diploid maternal plants when experimentally crossed with triploid pollen donors produced diploids and polyploid progeny, while when pollinated with a mixture of the pollen of diploids and triploids or insect pollinated, no polyploids were discovered. It seems that in the mixture with the pollen of diploids, the pollen of triploids is ineffective. Tetraploids produce hybrids much easier with diploid mothers and their role in wild populations requires further study. Triploid mothers, even those with subregular pollen did not show traces of facultative apomixis. BIII hybrids were present in the progeny of both triploids and tetraploids, in tetraploids in quite high percentages (up to 50% of the progeny in some crosses).