2n Eggs Research Articles

Genome sequence analysis provides insights into the mode of 2n egg formation in Solanum malmeanum.

Our genomic investigation confirms the mechanism of 2n eggs formation in S. malmeanum and aid in optimizing the use of wild germplasm. Wild potatoes are a valuable source of agronomic traits. However, substantial reproductive barriers limit gene flow into cultivated species. 2n gametes are instrumental in preventing endosperm abortion caused by genetic imbalances in the endosperm. However, little is known about the molecular mechanisms underlying the formation of 2n gametes. Here, the wild species Solanum malmeanum Bitter (2x, 1EBN, endosperm balance number) was used in inter- and intrapoloid crosses with other Solanum species, with viable seeds being produced only when S. malmeanum was used as the female parent to cross the 2EBN Solanum genus and with the likely involvement of 2n gametes. Subsequently, we substantiated the formation of 2n eggs in S. malmeanum using fluorescence in situ hybridization (FISH) and genomic sequencing technology. Additionally, the transmission rate of maternal heterozygous polymorphism sites was assessed from a genomic perspective to analyze the mode of 2n egg formation in S. malmeanum × S. tuberosum and S. malmeanum × S. chacoense crosses; each cross acquired an average of 31.12% and 22.79% maternal sites, respectively. This confirmed that 2n egg formation in S. malmeanum attributed to second-division restitution (SDR) coupled with the occurrence of exchange events. The high-throughput sequencing technology used in this study has strong advantages over traditional cytological analyses. Furthermore, S. malmeanum, which has a variety of excellent traits not available from present cultivated potato genepool, has received little research attention and has successfully achieved gene flow in cultivated species in the current study. These findings will facilitate the understanding and optimization of wild germplasm utilization in potatoes.

The Hybridization Barrier between Herbaceous Medicago sativa and Woody M. arborea Is Weakened by Reproductive Abnormalities in M. sativa Seed Parents.

Historically, crosses between Medicago sativa (alfalfa) and M. arborea with alfalfa as the seed parent failed, as did crosses using M. arborea as the seed parent. Thus, a reproductive barrier kept the two species isolated until early in this century. The breakthrough came when alfalfa seed parents were identified in Wisconsin USA and Queensland AU that produced partial hybrids (hereafter hybrids). The hybrids were obtained by making large numbers of crosses on selected alfalfa parents. This was the first level of weakening the crossing barrier as reported in Plants in 2013. Further weakening of the barrier is reported herein whereby more hybrids were obtained with fewer crosses. This was accomplished by pedigree selection for new alfalfa seed parents and by using a product of the first hybrids called Alborea. New alfalfa seed parents were crossed with M. arborea, and Alborea parents were backcrossed to M. arborea. Hybrid plants were produced with fewer crosses in both cases. These hybrids, like the first hybrids, have mostly alfalfa traits but also have traits from M. arborea. It was theorized early on that the alfalfa component could be explained by 2n eggs in the alfalfa parents that were fertilized by normal n gametes from M. arborea. Evidence that the Wisconsin alfalfa and Alborea seed parents did in fact produce 2n eggs was reported in Plants in 2022. Moreover, they produced 2n eggs at approximately the same frequency that they produced hybrids. As reported herein, Alborea parents produced the highest frequency of hybrids and thus had the weakest barrier. Importantly, they also have the highest frequency of 2n eggs. It was determined that alfalfa and Alborea parents that produce 2n eggs and hybrids, also produce 2n pollen. In effect, an experiment was undertaken in reverse showing that 2n pollen could be used to screen for plants that produce hybrids. In the thousands of crosses made over the years, fertilization of normal n eggs in alfalfa parents always failed. Normal meiosis appears to be the main barrier to producing interspecific hybrids in our case. Fertilization of abnormal 2n eggs ensures sufficient alfalfa genetic material to continue embryogenesis. Evidently, the meiotic abnormality of 2n eggs is the major factor that weakens the crossing barrier.

Evidence That 2n Eggs Explain Partial Hybrids between Medicago sativa and Medicago arborea.

Selected genotypes of alfalfa (Medicago sativa) produce partial hybrids in sexual crosses with Medicago arborea, as reported in Plants (2013). The hybrids contain mostly alfalfa DNA and traits, but also contain DNA and traits from M. arborea. It was proposed in 2008 that the partial hybrids could be explained by fertilization of 2n eggs in alfalfa by normal pollen from M. arborea, followed by partial loss of M. arborea chromosomes during embryogenesis. In this paper, we confirm the presence of 2n eggs in the first alfalfa parents that produced hybrids. The test for 2n eggs involved pollinating 4x alfalfa with pollen from 8x alfalfa. The production of 8x progeny in the cross proved that selected alfalfa parents produced 2n eggs. Thus, 2n eggs appear to explain how the partial hybrids (hereafter hybrids) contain mostly alfalfa DNA and traits. However, two of the six alfalfa plants that did not hybridize with M. arborea also had 2n eggs. Thus, although 2n eggs explain the alfalfa content of hybrids, 2n eggs are not the only factor involved in weakening the hybridization barrier, and in transferring genes to alfalfa from M. arborea.

The common occurrence of 2n eggs by lily F1 distant hybrids and its significance on lily breeding: a case of analyzing OT hybrids

The common occurrence of 2n eggs by lily F1 distant hybrids and its significance on lily breeding: a case of analyzing OT hybrids

A new way to produce odd‐allotetraploid lily (Lilium) through 2n gametes

AbstractOdd‐allotetraploid lily, such as ‘Honesty’ (LAAA), is a new type of allotetraploids. However, there has been no report on how to breed odd‐allotetraploid lily cultivars so far. According to their genomic composition, such as LAAA, we proposed LA × AAAA and LALA × AAAA would be two ways to breed them since LA lily could produce a small amount of 2n gametes and LALA could produce a large amount of normal 2x gametes. In the present study, an interploid cross, LA × AAAA, was made and its five seedlings were analysed using genomic in situ hybridization. Four of them were odd‐allotetraploid, one was near odd‐allotetraploid. Their intergenomic variations caused by 2n eggs increased the chance of selecting new cultivars. It was concluded that the interploid cross was a promising way to breed odd‐allotetraploid lilies with high genetic variations at chromosome levels.

Frequency of Polyploids of Solanum tuberosum Dihaploids in 2X × 2X Crosses

When breeding diploid potatoes, tetraploid progeny can result from the union of 2n eggs and 2n pollen in 2x-2x crosses. Thirty-three crosses were made to examine tetraploid progeny frequency in 2x-2x crosses. All crosses were between S. tuberosum dihaploids and diploid self-compatible donors, M6 and DRH S6-10-4P17. Using chloroplast counting for ploidy determination, the frequency of tetraploid progeny was as high as 45% in one of the 33 crosses. Based upon single nucleotide polymorphism (SNP) genotyping, the tetraploid progeny were attributed to bilateral sexual polyploidization (BSP), which is caused by the union of 2n egg and 2n pollen. Dihaploids were identified that produce lower frequencies of 2n eggs. The results of this study suggest that S. tuberosum dihaploids with a high frequency of 2n eggs should be avoided in 2x - 2x crosses for diploid breeding programs.

Low temperature increases the frequency of 2n female gametes in the diploid rubber tree (Hevea brasiliensis (Willd. ex A.Juss.) Müll. Arg.)

Triploid breeding via the fusion of 2n gametes and n gametes is an efficient method for breeding new varieties in woody plants. Triploid plants are frequently observed in the seedlings of the diploid rubber clone GT1. However, the factors affecting the occurrence and source of triploid progeny in GT1 are still unknown. In this study, the correlation between temperature and triploid progeny frequency in GT1 in different environments over 6 years was analyzed to evaluate the influence of temperature on the frequency of 2n female gametes. A total of 76 triploid progenies were analyzed using 19 simple sequence repeat markers, with the aim of revealing their genetic origin and the mechanism underlying 2n megagametophyte formation. The results showed that low temperature increased the frequency of triploid progeny in GT1, and the determination of allelic status revealed that all triploids were derived from the 2n female gametes, thereby confirming that low temperature increased the frequency of 2n female gametes. The rate of maternal heterozygosity restitution (HR) of seven loci varied from 2.70 to 67.65% with a mean value of 40.56%, and the rate of maternal HR of 37 triploids varied from 14.29 to 66.67% with a mean value of 41.38%, indicating that the production of 2n eggs in the triploid seedlings of GT1 was by second division restitution. The results of this study are valuable for the polyploid breeding of rubber trees.

Creation of long-day adapted tetraploid potatoes from crosses of Solanum tarijense females with Andigena males

T-type is a common chloroplast DNA type among modern potato varieties (Solanum tuberosum), their progenitor Chilean tuberosum, and a diploid wild species S. tarijense. To recreate Chilean tuberosum, we made 983 pollinations between 10 accessions of T-type chloroplast DNA-holding S. tarijense used as females and 32 Andean tetraploid landraces (Andigena) used as males, from which 14 tetraploid hybrids were obtained. These interspecific hybrids grew vigorously with long stolons, flowered well with high male and female fertilities, and matured 1 month later than modern varieties. Seedlings of selfed and sib-crossed interspecific hybrids were artificially selected for tuber yield under long days. The selected clones were grown in the field, of which two clones produced over 1 kg of edible tubers per plant. These results lend experimental support to the hypothesis that the Chilean tuberosum originated by selection for long-day adaptability from tetraploid hybrids that occurred by fertilization of a 2n egg of S. tarijense and n pollen of Andigena.

Pairing analysis and in situ Hybridisation reveal autopolyploid-like behaviour in Solanum commersonii\xa0\xd7\xa0S. tuberosum (potato) interspecific hybrids

Wild potato relatives are rich sources of desirable traits for introgressive hybridisation into cultivated potato. One of them, Solanum commersonii (2n = 2x = 24, 1EBN, endosperm balance number), is an important species belonging to the potato tertiary genepool. It can be used in potato breeding through bridge crosses and 2n gamete production. Triploid F1 hybrids between S. commersonii (through spontaneous 2n egg formation) and diploid 2EBN Solanum tuberosum Group Phureja were crossed with S. tuberosum Group Tuberosum, resulting in successive backcross 1, 2 and 3 progenies. The main aim of this study was to determine if there are any barriers to homoeologous pairing and recombination in the allotriploid (S. commersonii × potato) hybrids and their backcrosses, and so to predict if S. commersonii chromosomes can be transmitted to the next generation and introgressed into their recipient potato chromosomes. Microscopic observations of spread pollen mother cells suggested no preferential pairing in the triploid hybrids, while chromosome transmission and segregation in further meiotic stages were fairly balanced. Fluorescent in situ hybridisation with BAC probes (BAC FISH) was used to obtain markers to trace the meiotic behaviour of specific chromosome pairs. Moreover, genomic in situ hybridisation (GISH) demonstrated no obvious differences in fluorescence signals between the homoeologues suggesting that repetitive sequences did not diverge much between the parental species. As a consequence, we were not able to trace the course of the S. commersonii chromosomes in the successive introgressive hybridisation backcross generations. Our results strongly point at a high genomic similarity between the homoeologous chromosomes promising high suitability of S. commersonii in introgressive hybridisation breeding of potato.

Sexual Polyploidization in Medicago sativa L.: Impact on the Phenotype, Gene Transcription, and Genome Methylation.

Polyploidization as the consequence of 2n gamete formation is a prominent mechanism in plant evolution. Studying its effects on the genome, and on genome expression, has both basic and applied interest. We crossed two diploid (2n = 2x = 16) Medicago sativa plants, a subsp. falcata seed parent, and a coerulea × falcata pollen parent that form a mixture of n and 2n eggs and pollen, respectively. Such a cross produced full-sib diploid and tetraploid (2n = 4x = 32) hybrids, the latter being the result of bilateral sexual polyploidization (BSP). These unique materials allowed us to investigate the effects of BSP, and to separate the effect of intraspecific hybridization from those of polyploidization by comparing 2x with 4x full sib progeny plants. Simple sequence repeat marker segregation demonstrated tetrasomic inheritance for all chromosomes but one, demonstrating that these neotetraploids are true autotetraploids. BSP brought about increased biomass, earlier flowering, higher seed set and weight, and larger leaves with larger cells. Microarray analyses with M. truncatula gene chips showed that several hundred genes, related to diverse metabolic functions, changed their expression level as a consequence of polyploidization. In addition, cytosine methylation increased in 2x, but not in 4x, hybrids. Our results indicate that sexual polyploidization induces significant transcriptional novelty, possibly mediated in part by DNA methylation, and phenotypic novelty that could underpin improved adaptation and reproductive success of tetraploid M. sativa with respect to its diploid progenitor. These polyploidy-induced changes may have promoted the adoption of tetraploid alfalfa in agriculture.

Evidence of the production of 2n eggs in diploid plants of the autopolyploid complex Turnera sidoides L. (Passifloraceae)

Turnera sidoides is a complex of dystilous perennial rhizomatous herbs with five subspecies in which diploid (2n = 2x = 14) to octoploid (2n = 8x = 56) cytotypes were found. Previous studies have suggested an autopolyploid origin of the complex, and provided evidence for the production of 2n pollen in experimental conditions as well as in natural populations. However, only the production of 2n pollen has been demonstrated so far, while the production of 2n eggs on experimental crosses or in natural populations of T. sidoides remains unknown. In this paper we investigate the production of 2n eggs in diploid individuals to understand how they may have contributed to the origin and establishment of polyploids in diploid populations of T. sidoides. Our progeny test and flow cytometric analysis of seeds collected in natural populations of this species complex, show that the triploid embryos originated from 2n eggs, suggesting that 2n eggs can contribute to the origin of neopolyploids by sexual unilateral polyploidization as well as by sexual bilateral polyploidization. The occurrence of plants that continuously form many 2n eggs and pollen would play a key role in the establishment of neopolyploids in natural diploid populations.

Transcriptomic changes following synthesis of a Populus full-sib diploid and allotriploid population with different heterozygosities driven by three types of 2n female gamete.

Diploid gametes are usually applied to produce triploids of Populus [originating from first-division restitution (FDR), second-division restitution (SDR), and postmeiotic restitution (PMR) 2n eggs]. Three types of 2n gametes transmitted different parental heterozygosities in Populus. Failed spindle formation and no chromosomal separation to opposite poles during meiosis I mean that FDR 2n gametes carry nonsister chromatids that are potentially heterozygous. By contrast, SDR 2n gametes result from failed sister chromatid separation in meiosis II, and therefore, they carry sister chromatid that are potentially homozygous. Completely homozygous 2n gametes can arise from the PMR mechanism. The alteration of gene expression resulting from allopolyploidization is a prominent feature in plants. We compared gene expression in the full-sib progeny of three allotriploid Populus populations (triploid-F, triploid-S, and triploid-P) with that in its parent species, and their full-sib diploid F1 hybrid. Genome-wide expression level dominance was biased toward the maternal in the diploid F1 hybrid and three allotriploid populations, whereas our data indicated important, but different, effects of the transmission of different heterozygosity by 2n female gametes in the expression patterns of allopolyploids. Because of the higher level of heterozygosity, the triploids had higher rates of non-additive and transgressive expression patterns in the triploid-F than in triploid-S and triploid-P. Compared with diploid F1, about 30-fold more genes (251) were differently expressed in the triploid-F than in the triploid-S (9) and triploid-P (8), respectively. These findings indicate that hybridization and polyploidization have immediate and distinct effects on the large-scale patterns of gene expression, and different effects on the transmission of heterozygosity by three 2n female gametes.

Inheritance and variation of Cytosine methylation in three populus allotriploid populations with different heterozygosity.

DNA methylation is an epigenetic mechanism with the potential to regulate gene expression and affect plant phenotypes. Both hybridization and genome doubling may affect the DNA methylation status of newly formed allopolyploid plants. Previous studies demonstrated that changes in cytosine methylation levels and patterns were different among individual hybrid plant, therefore, studies investigating the characteristics of variation in cytosine methylation status must be conducted at the population level to avoid sampling error. In the present study, an F1 hybrid diploid population and three allotriploid populations with different heterozygosity [originating from first-division restitution (FDR), second-division restitution (SDR), and post-meiotic restitution (PMR) 2n eggs of the same female parent] were used to investigate cytosine methylation inheritance and variation relative to their common parents using methylation-sensitive amplification polymorphism (MSAP). The variation in cytosine methylation in individuals in each population exhibited substantial differences, confirming the necessity of population epigenetics. The total methylation levels of the diploid population were significantly higher than in the parents, but those of the three allotriploid populations were significantly lower than in the parents, indicating that both hybridization and polyploidization contributed to cytosine methylation variation. The vast majority of methylated status could be inherited from the parents, and the average percentages of non-additive variation were 6.29, 3.27, 5.49 and 5.07% in the diploid, FDR, SDR and PMR progeny populations, respectively. This study lays a foundation for further research on population epigenetics in allopolyploids.

Analysis of transmission of heterozygosity by 2n gametes in Populus (Salicaceae)

It is well known that polyploidy is a major force in plant evolution and diversification. Evaluating the transmission of heterozygosity by 2n gametes with different formation mechanisms is of crucial importance. In the present study, 120 triploid hybrids [originating from first-division restitution (FDR), second-division restitution (SDR), and postmeiotic restitution (PMR) 2n eggs of the same female parent] and 30 pairs of polymorphic simple sequence repeat (SSR) markers (distributed across all 19 chromosomes of Populus) were used to evaluate the maternal heterozygosity transmitted by each type of 2n eggs. FDR, SDR, and PMR 2n eggs transmitted different parental heterozygosities (0.7480, 0.3958, and 0.3590, respectively) in Populus. In addition, compared with the small coefficient of variation (CV = 13.50 %) of heterozygosity transmitted by the 19 chromosomes of FDR 2n eggs, there was greater variation in heterozygosity transmitted by SDR and PMR 2n eggs (CV = 51.92 % and CV = 63.14 %, respectively). This may have resulted from the locations of the applied SSR markers, in addition to the different formation mechanisms of 2n gametes. Variations in locus-centromere distance resulted in variations in recombination rates, which influence the degree of heterozygosis at the detected locus. In conclusion, our data provides a useful reference point for the suitable applications of 2n gametes in future studies, promoting the further research of polyploidy in plants.

A strategy for characterization of persistent heteroduplex DNA in higher plants.

Heteroduplex DNA (hDNA) generated during homologous recombination (HR) is an important component that shapes genetic diversity in sexually reproducing organisms. However, studies of this process in higher plants are limited. This is because hDNAs are difficult to capture in higher plants as their reproductive developmental model only produces normal gametes and does not preserve the mitotic products of the post-meiotic segregation (PMS) process which is crucial for studying hDNAs. In this study, using the model system for tree and woody perennial plant biology (Populus), we propose a strategy for characterizing hDNAs in higher plants. We captured hDNAs by constructing triploid hybrids originating from a cross between unreduced 2n eggs (containing hDNA information as a result of inhibition chromosome segregation at the PMS stage) with normal male gametes. These triploid hybrids allowed us to detect the frequency and location of persistent hDNAs resulting from HR at the molecular level. We found that the frequency of persistent hDNAs, which ranged from 5.3 to 76.6%, was related to locations of the simple sequence repeat markers at the chromosomes, such as the locus-centromere distance, the surrounding DNA sequence and epigenetic information, and the richness of protein-coding transcripts at these loci. In summary, this study provides a method for characterizing persistent hDNAs in higher plants. When high-throughput sequencing techniques can be incorporated, genome-wide persistent hDNA assays for higher plants can be easily carried out using the strategy presented in this study.

Megasporogenesis, megagametogenesis, and induction of 2n eggs with colchicine in poplar section Aigeiros

Triploid breeding through 2n eggs induction has become one of the most powerful approaches for the improvement of the genus Populus. To enhance the efficiency of triploid production in section Aigeiros, female buds (catkins) of P. × euramericana (Dode) Guinier at different developmental stages were exposed to colchicine for 2n eggs induction. The relationship between flower bud morphological characteristics (or time after pollination) and female meiotic stage (or embryo sac development) was established to guide colchicine treatments. In the resulting progeny, three triploid seedlings were obtained via embryo rescue and identified by flow cytometric analyses. Cytological observation revealed that the leptotene to pachytene stage of meiosis and 12 h after pollination might be the suitable period for 2n eggs induction with colchicine in P. × euramericana. Results and protocols related to 2n eggs induction, effective methodologies for embryo rescue, and ploidy evaluation in this study might be applicable in polyploidy breeding in section Aigeiros.

Assessment of the genetic composition of triploid hybridPopulususing SSR markers with low recombination frequencies

The induction of nonreduced gametes, whether from first-division restitution (FDR), second-division restitution (SDR), or both, is an important approach for polyploidization. However, an accurate method for determining the genetic constitution of polyploid hybrids is not available. In this study, based on both flow cytometric analysis and somatic chromosome counting, 164 triploid hybrids of the female parent Populus pseudo-simonii × P. nigra ‘Zheyin3#’ (2n = 2x = 38, abbreviated ZY3) and male parent P. × beijingensis (2n = 2x = 38, abbreviated BJY) were produced by high-temperature treatment during ZY3 megasporogenesis. Using six simple sequence repeat (SSR) markers with low recombination frequencies to reduce the impact of recombination, we analyzed the allelic configurations of the obtained triploid hybrids. Although the allelic configurations were not always consistent at all six loci, by combining the allelic configurations at the six loci, we inferred that 40 triploids originated from FDR 2n eggs of ZY3, whereas the others originated from SDR 2n eggs. In conclusion, our study provides a novel and effective tool for analyzing 2n gametes and performing early selection to improve triploid poplar breeding programs.

Pollen-mediated gene flow from a commercial potato cultivar to the wild relativeS. chacoenseBitter under experimental field conditions in Argentina

The common potato, Solanum tuberosum ssp. tuberosum (tbr, 2n = 4x = 48; 4EBN), has many closely related wild tuber-bearing species. Around 28 to 35 of them spontaneously grow in Argentina overlapping, in some areas, with the crop and/or experimental transgenic potatoes. Although it is well proven that hybridization barriers in potatoes can be incomplete, information on gene flow between cultivated and wild germplasm is scarce. Thus, a gene flow field experiment with a circular array was set up in Balcarce, Argentina, in 2009, and evaluated over two seasons. The tetraploid tbr cultivar Huinkul MAG and one compatible cloned genotype of the related wild potato S. chacoense Bitter (chc, 2n = 2x = 24; 2EBN), which produced 2n eggs, were used, respectively, as pollen donor and receptor. Berries with hybrid seeds - as revealed by ploidy and RAPD profiles - were obtained in one season, at 30 m from the pollen donor. These results reinforce others previously obtained with the same pollen donor and a male sterile tbr cultivar in a similar array, pointing out to the need of increasing isolation distances in areas of overlap between cultivated and wild potato germplasm to prevent or minimize undesirable pollen-mediated gene flow.

Sexual polyploidization in plants – cytological mechanisms and molecular regulation

In the plant kingdom, events of whole genome duplication or polyploidization are generally believed to occur via alterations of the sexual reproduction process. Thereby, diploid pollen and eggs are formed that contain the somatic number of chromosomes rather than the gametophytic number. By participating in fertilization, these so-called 2n gametes generate polyploid offspring and therefore constitute the basis for the establishment of polyploidy in plants. In addition, diplogamete formation, through meiotic restitution, is an essential component of apomixis and also serves as an important mechanism for the restoration of F1 hybrid fertility. Characterization of the cytological mechanisms and molecular factors underlying 2n gamete formation is therefore not only relevant for basic plant biology and evolution, but may also provide valuable cues for agricultural and biotechnological applications (e.g. reverse breeding, clonal seeds). Recent data have provided novel insights into the process of 2n pollen and egg formation and have revealed multiple means to the same end. Here, we summarize the cytological mechanisms and molecular regulatory networks underlying 2n gamete formation, and outline important mitotic and meiotic processes involved in the ectopic induction of sexual polyploidization.

The effect of 2n gametes on sex ratios in Actinidia

Sex can sometimes lead to complications. In some crops, 2n gametes have been exploited by plant breeders to transfer genetic variation between taxa of different ploidy levels. However, their role and use in dioecious genera have received relatively little attention. In the dioecious genus Actinidia (kiwifruit), seedling populations usually segregate equally for females and males as sex is determined by an XX female/XY male system. While fertilization involving 2n egg cells is not expected to affect the sex ratios of progenies, fertilization involving 2n pollen is likely to produce progenies with excess males. The extent of sex ratio distortion will depend on the relative contributions of first and second division restitution, and the frequency and location of cross-overs in meiosis. In this study, seedlings recovered from crosses between females of hexaploid Actinidia deliciosa and males of two diploid species, Actinidia chinensis and Actinidia eriantha, included a proportion of pentaploid hybrids presumably derived from fertilization involving 2n pollen. Most of these pentaploids were male, and a proportion of them were likely to be carrying two Y chromosomes. If used as parents in further crosses, males with multiple Y chromosomes are likely to cause distorted sex ratios in their immediate progenies. In dioecious genera such as Actinidia, the effects on sex ratios of different mechanisms of ploidy change need to be taken into account when considering the evolution of polyploidy and the design of breeding strategies involving ploidy manipulation.