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11.0: Prelude to Meiosis and Sexual Reproduction - Biology

11.0: Prelude to Meiosis and Sexual Reproduction - Biology


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The ability to reproduce in kind is a basic characteristic of all living things. In kind means that the offspring of any organism closely resemble their parent or parents. Hippopotamuses give birth to hippopotamus calves, Joshua trees produce seeds from which Joshua tree seedlings emerge, and adult flamingos lay eggs that hatch into flamingo chicks. In kind does not generally mean exactly the same. Whereas many unicellular organisms and a few multicellular organisms can produce genetically identical clones of themselves through cell division, many single-celled organisms and most multicellular organisms reproduce regularly using another method. Sexual reproduction is the production by parents of two haploid cells and the fusion of two haploid cells to form a single, unique diploid cell. In most plants and animals, through tens of rounds of mitotic cell division, this diploid cell will develop into an adult organism. Haploid cells that are part of the sexual reproductive cycle are produced by a type of cell division called meiosis. Sexual reproduction, specifically meiosis and fertilization, introduces variation into offspring that may account for the evolutionary success of sexual reproduction. The vast majority of eukaryotic organisms, both multicellular and unicellular, can or must employ some form of meiosis and fertilization to reproduce.


Biology

Publication date 2013 Usage Attribution 4.0 International Topics Biology -- Textbooks, Biology Publisher Houston, Texas : OpenStax College, Rice University Collection opensource Language English

1 online resource (1517 pages) :

Biology is grounded in an evolutionary basis and includes exciting features that highlight careers in the biological sciences and everyday applications of the concepts at hand. To meet the needs of today's instructors and students, some topics have been condensed and combined for example, phylogenetic trees are presented in the various ways they are currently being developed by scholars, so instructors can adapt their teaching to the approach that works best in their classroom. The book also includes an innovative art program that incorporates critical thinking and clicker questions to help students understand-and apply-key concepts

"Senior contributors, Yael Avissar, cell biology Jung Choi, genetics Jean DeSaix, evolution Vladimir Jurukovski, animal physiology Robert Wise, plant biology Connie Rye, general content lead"--Page 10

Resource simultaneously available in high- and low-resolution PDF files, direct-access HTML version, EPUB format, and iBooks format

The study of life -- The chemical foundation of life -- Biological macromolecules -- Cell structure -- Structure and function of plasma membranes -- Metabolism -- Cellular respiration -- Photosynthesis -- Cell communication -- Cell reproduction -- Meiosis and sexual reproduction -- Mendel's experiments and heredity -- Modern understandings of inheritance -- DNA structure and function -- Genes and proteins -- Gene expression -- Biotechnology and genomics -- Evolution and the origin of species -- The evolution of populations -- Phylogenies and the history of life -- Viruses -- Prokaryotes : bacteria and archaea -- Protists -- Fungi -- Seedless plants -- Seed plants -- Introduction to animal diversity -- Invertebrates -- Vertebrates -- Plant form and physiology -- Soil and plant nutrition -- Plant reproduction -- The animal body : basic form and function -- Animal nutrition and the digestive system -- The nervous system -- Sensory systems -- The endocrine system -- The musculoskeletal system -- The respiratory system -- The circulatory system -- Osmotic regulation and excretion -- The immune system -- Animal reproduction and development -- Ecology and the biosphere -- Population and community ecology -- Ecosystems -- Conservation biology and biodiversity

Low-resolution PDF file (revision BM-1-001-DW, viewed November 19, 2014) title from title page of low-resolution PDF file


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Final product will be 4 haploid (half the number of chromosomes) cells 2 sister chromatids separate haploid diploid meiosis i (reduction division) meiosis ii (equational homologs separate result:

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Source: image3.slideserve.com

Algebra 2 worksheets with answers. Final product will be 4 haploid (half the number of chromosomes) cells 2 sister chromatids separate haploid diploid meiosis i (reduction division) meiosis ii (equational homologs separate result: By the way, about chapter 11.4 meiosis worksheet answers, we have collected particular related photos to complete your references. Some cells go through a programmed death. Verbs of perception + present participle.

Match the following occurances with their appropriate phase in meiosis. Even something as easy as guessing the beginning letter of long words can assist your child improve his phonics abilities. Present simple and present continuous past continuous worksheet 10 : Verbs of perception + present participle. It is held with both hands and the thumbs are used to handle the direction and action buttons.

This worksheet is intended to reinforce concepts related to meiosis and sexual reproduction. Infection (11) instrumentation (34) laboratory tests (8) microbiology pictures (9) microscopy (22) molecular biology (59) mycology (28) parasitology (7) pharmacology (10) product review (1) protocols (12) report and. In interphase, the dna is copied, the cell grows and the. Unspecialized cells are either embryonic or adult. Mendel and meiosis worksheet answers 440 x 320 440 x 320 from meiosis worksheet answer key source.

The process of meiosis happens in the male and female reproductive organs. Section 11 4 meiosis worksheet answers. Students worked on the section 1 reinforcement worksheet. Answer key to the meiosis worksheet. Go back and take a closer look at the information.

Name answer key date class chapter 9 worksheet section 1: Even something as easy as guessing the beginning letter of long words can assist your child improve his phonics abilities. Section 11 4 meiosis worksheet answers. Present perfect l) build up questions and give answers: Hi, i think those are still just the math calc section questions since there are 38 questions and not 20.

Source: ecdn.teacherspayteachers.com

I s there an answer sheet? In interphase, the dna is copied, the cell grows and the. By the way, about chapter 11.4 meiosis worksheet answers, we have collected particular related photos to complete your references. The process of meiosis happens in the male and female reproductive organs. 3 describing functions and features.

The answers to today's worksheet are posted on the attachment page for unit 3, chapter 10 as mitosis.

Source: www.williamwithin.com

Present perfect tense worksheet 11 :

Even something as easy as guessing the beginning letter of long words can assist your child improve his phonics abilities.

Infection (11) instrumentation (34) laboratory tests (8) microbiology pictures (9) microscopy (22) molecular biology (59) mycology (28) parasitology (7) pharmacology (10) product review (1) protocols (12) report and.

The process of meiosis happens in the male and female reproductive organs.

Source: image.slidesharecdn.com

3 describing functions and features.

This worksheet is intended to reinforce concepts related to meiosis and sexual reproduction.

16) psat 11/2/16 and backup link and answers.

Then, as they read the section, have students take notes that will help answer the questions they've read.

18.01.2019 · ahead of preaching about biology section 11 4 meiosis worksheet answer key, you need to know that instruction is actually all of our crucial for a much better down the road.

It is held with both hands and the thumbs are used to handle the direction and action buttons.

I s there an answer sheet?

They go through metaphase, divide, and do the process.

Source: cdn.slidesharecdn.com

2 a) stages of meiosis goal:

Martin wants to a sell a flat.

2 a) stages of meiosis goal:

Some cells go through a programmed death.

Unspecialized cells are either embryonic or adult.

Source: image.slidesharecdn.com

16) psat 11/2/16 and backup link and answers.

Source: image3.slideserve.com

Section 11 4 meiosis worksheet answers.

The process of meiosis happens in the male and female reproductive organs.

Source: cdn.slidesharecdn.com

On each of the images, label the phase of meiosis and cytokinesis.

16) psat 11/2/16 and backup link and answers.

It is held with both hands and the thumbs are used to handle the direction and action buttons.

Infection (11) instrumentation (34) laboratory tests (8) microbiology pictures (9) microscopy (22) molecular biology (59) mycology (28) parasitology (7) pharmacology (10) product review (1) protocols (12) report and.


Mitosis and Meiosis Comparison - PowerPoint PPT Presentation

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Meiosis Review - PowerPoint PPT Presentation

PowerShow.com is a leading presentation/slideshow sharing website. Whether your application is business, how-to, education, medicine, school, church, sales, marketing, online training or just for fun, PowerShow.com is a great resource. And, best of all, most of its cool features are free and easy to use.

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For a small fee you can get the industry's best online privacy or publicly promote your presentations and slide shows with top rankings. But aside from that it's free. We'll even convert your presentations and slide shows into the universal Flash format with all their original multimedia glory, including animation, 2D and 3D transition effects, embedded music or other audio, or even video embedded in slides. All for free. Most of the presentations and slideshows on PowerShow.com are free to view, many are even free to download. (You can choose whether to allow people to download your original PowerPoint presentations and photo slideshows for a fee or free or not at all.) Check out PowerShow.com today - for FREE. There is truly something for everyone!

presentations for free. Or use it to find and download high-quality how-to PowerPoint ppt presentations with illustrated or animated slides that will teach you how to do something new, also for free. Or use it to upload your own PowerPoint slides so you can share them with your teachers, class, students, bosses, employees, customers, potential investors or the world. Or use it to create really cool photo slideshows - with 2D and 3D transitions, animation, and your choice of music - that you can share with your Facebook friends or Google+ circles. That's all free as well!


Concluding remarks

Aphids are remarkable for their ability to express alternate discrete phenotypes given the same genome. In order for sexual organisms to evolve facultative asexuality, we assume at least three changes to occur: prevention of recombination, lack of homologue association and prevention of genome reduction ( d'Erfurth et al., 2009 ). Evidence described here suggests the occurrence of these changes in pea aphids. Reduced recombination machinery activity may allow asexual aphids to prevent formation of crossovers necessary for the meiosis I division (D. Srinivasan and D. Stern, unpublished data). The absence of an aphid Rec8 homologue may limit homologue association in asexual aphids. Duplications and differential expression of cell cycle regulatory genes (Geminin, Cdt1, Cdk1, Wee1, Cdc25, Polo kinases, Aurora kinases, Cks proteins) may help alter the meiotic cell cycle program of asexual oocytes. Downstream targets of the expanded regulatory genes in the pea aphid should also be examined to determine if whole pathways have expanded or if plasticity is generally associated with changes in genes at the top of pathways. Further examination of expression patterns and functions in facultative asexuality will be important in understanding how aphids can exhibit plasticity in meiosis and sexual reproduction.


Results

Plant size and sexual system were significantly correlated, irrespective of the type of coding used (continuous or discrete) or whether the association was corrected for phylogenetic relatedness (Table 3). Species with separate sexes were significantly more likely to have larger gametophores than those with combined sexes (Table 3). In addition, maximum likelihood analysis of gametophore size, coded as a discrete character, showed evidence for the conditional evolution of separate sexes. Transitions from combined to separate sexes were significantly more likely in lineages that had large gametophores (q42 > q31, LR = 5.56, Table 4). The relationship between sexual system and size differed between moss clades, consistent with our hypotheses that the association would be influenced by growth form. Mosses in subclass Dicranidae, which have sexual structures at the tips of the (generally) upright growing shoot, showed a strong correlation between sexual system and size, whereas the effect was marginal or nonsignificant for subclass Bryidae (which include both acrocarpous and pleurocarpous mosses), and pleurocarpous-only Hypnanae (Table 3).

Trait Change to state 1 conditional upon breeding system state Change to state 0 conditional upon breeding system state Change to combined sexes depends on state of other trait Change to separate sexes depends on state of other trait Combined sexes evolves before change to state 1 Separate sexes evolves before change to state 0
Test q12=q34 q21=q43 q13=q24 q31=q42 q12=q13 q42=q43
Gametophore size (0 small, 1 large) 2.06 1.96 0.23 5.56* trend: separate sexes gained in lineages with large gametophores 0.611 2.48
Spore size (0 small, 1 large) 0.25 0.67 0.25 9.56* trend: separate sexes gained in lineages with small spores 2.06 2.06
Chromosome number (0 ≤ 12, 1 >12) 2.32 1.20 3.81 trend: combined sexes gained in lineages with low chromosome numbers 11. 05*** trend: separate sexes gained in lineages with low chromosome numbers 5.09 trend: change in sexual system before change in chromosome number 0.01
Chromosome number (0=odd, 1=even) 3.31 4.61 trend: change to odd chromosome numbers in lineages with combined sexes 0.36 0.34 0.06 0.88

There was inconsistent evidence for associations with seta length and spore size. Although TIPS analyses suggested that mosses with combined sexes were significantly likely to have setae that were smaller than those of mosses with separate sexes, this result was not significant when taking phylogeny into account (Table 3). TIPS analysis suggested that mosses with combined sexes had significantly larger spores than those with separated sexes (Table 3), and there was significant evidence that these did not evolve independently (using a discrete, but not a continuous coding scheme, Table 3). We also found evidence for the conditional evolution of gender dimorphism: gender dimorphism was more likely to be gained in moss lineages with small spores (Table 4).

When species were treated as independent data points there was a significant association between separate sexes and asexual propagules, although this association was not as strong as that found by Longton and Schuster (1983) , and was not robust to corrections for multiple comparisons (Tables 3, 5). However, maximum likelihood analyses revealed no association between sexual system and asexual reproduction (Table 3).

Longton and Schuster (1983) This study
Asexual propagules No asexual propagules Asexual propagules No asexual propagules
Separate sexes 87 295 99 129
Combined sexes 11 234 43 96
χ 2 =36.5, P<0.0001 χ 2 =5.161, P=0.02

Species with separate sexes generally had lower chromosome numbers than those with combined sexes (Table 3), and as predicted species with separate sexes were significantly less likely to have even chromosome numbers (Table 3 and data not shown). Maximum likelihood analyses categorizing chromosome number into discrete classes suggested that the traits did not evolve independently regardless of whether chromosome number was coded using the threshold system or as odd versus even numbers (Table 3). A significant association was found both when polyploidy was determined as n ≥ 8 and as ≥ 10 (results not shown).

We hypothesized that changes in sexual system occur simultaneous to chromosome doubling. However, we found inconclusive evidence for simultaneous transitions. Changes in one trait did not obviously result in high transition rates in the second trait (i.e., q12 +q13 ≈q24 +q34, Appendix 1). There was a marginally significant difference between q12 and q13 using threshold coding, suggesting that gender dimorphism was more likely lost before chromosome number changed. This result was not found for n= 8 and n= 10 thresholds (results not shown) and the odd/even coding scheme, suggesting more ancient polyploidy may be involved. There was also no evidence for temporal order in the gains of both polyploidy and combined sexes (i.e., q34 ≈q24, threshold coding LR = 1.18, odd/even coding LR = 1.40), suggesting the evolution of combined sexes and polyploidy could occur simultaneously.

There were significant correlations between the other life-history traits tested in this study, but most did not hold up to correction for multiple comparisons. Plants with long setae were significantly likely to have small spore size (Table 6), likely reflecting trade-offs for finite resources, or selection for increased dispersability. Plants with large chromosome numbers also had long setae, although this became nonsignificant after correction for multiple comparisons. There was a negative association between chromosome number and plant size, likely influenced by the correlation of gametophore size and chromosome number with sexual system, but again this was not significant after sequential Bonferroni correction. Surprisingly, there was no evidence for a relationship between asexual propagules and any of the continuous traits (Table 6).

Seta length Spore size Asexual propagules Chromosome number
Gametophore size 0.07 0.05 −0.07 −0.11
Seta length −0.13* −0.02 0.11
Spore size −0.08 −0.06
Asexual propagules −0.008

Results

Unpaired supernumerary chromosomes show drive correlated with mitochondrial transmission

To test the transmission of supernumerary chromosomes during meiosis we used the reference strain IPO323 (mating type mat1-1) and eight isogenic chromosome deletion strains (IPO323㥌hr14-21, mating type mat1-1) generated in a previous study (Habig et al., 2017). Each of the chromosome deletion strains differs in the absence of one supernumerary chromosome, thereby allowing us to compare the transmission of individual chromosomes in a paired and an unpaired state. We crossed these strains, in planta, with another Z. tritici isolate: IPO94269 (mat1-2) ( Figure 1B ) in three separate experiments (A, B and C) and used a combination of PCR assays, electrophoretic karyotyping and whole genome sequencing to assess the segregation of chromosomes during meiosis. IPO94269 contains six supernumerary chromosomes homologous to the IPO323 chromosomes 14, 15, 16, 17, 19, and 21 ( Figure 1𠅏igure supplement 1 ) (Goodwin et al., 2011). The experiments included a total of 39 crosses of IPO323/IPO323 chromosome deletion strains with IPO94269 resulting in different complements of paired and unpaired supernumerary chromosomes in the diploid zygote (Table 1, Supplementary file 1). We hypothesized that the inheritance of the unpaired supernumerary chromosomes could be linked to the female or male role of the parental strain. Sexual mating of heterothallic fungi of the genus Zymoseptoria involves a female partner that produces a sexual structure called the ascogonium. The ascogonium receives the spermatium with the male nucleus from the fertilizing male partner through a particular structure called the trichogyne (Crous, 1998) ( Figure 1A ). Importantly, the same strain can act as either the female or male partner (Kema et al., 2018). Mitochondrial transmission is generally associated with the female structure (Ni et al., 2011). We used specific mitochondrial PCR based markers to distinguish the mitochondrial genotype in the progeny and thereby determine which of the two parental strains (in this case IPO323 or IPO94269) acted as a female partner in a cross.

In all three experiments the ascospore progeny showed either the mitochondrial genotype of IPO94269 or IPO323. Therefore, both strains can act as the female and male partner during crosses (Supplementary file 2𠄴). However, transmission of the mitochondrial genotype varied significantly between experiments with the relative frequency of the IPO94269 mitochondrial genotype in the progeny being 80%, 11% and 65% in experiment A, B and C, respectively ( Figure 2A ). Interestingly, the transmission of unpaired chromosomes correlated to the sexual role (female/male) of the parent from which the unpaired chromosome was inherited. Unpaired chromosomes inherited from IPO94269 were underrepresented among ascospores with the IPO323 parent mitochondrial genotype ( Figure 2B ). In contrast, the unpaired supernumerary chromosomes 18 and 20, which were always inherited from the parent IPO323, were highly overrepresented among ascospores with the IPO323 mitochondrial genotype ( Figure 2B ). For ascospores with the mitochondrial type of the IPO94269 parent, this segregation distortion was reversed. Unpaired chromosomes inherited from IPO94269 (with the exception of unpaired chr14) were highly overrepresented among ascospores with the mitochondrial genotype of the IPO94269 parent. On the other hand, the unpaired supernumerary chromosomes 18 and 20, always inherited from the IPO323 parent, were underrepresented among ascospores with the mitochondrial genotype of the IPO94269 parent ( Figure 2B ).

(A) Relative frequencies of mitochondrial genotypes in random and randomized ascospores in experiments A, B and C. The mitochondrial transmission varied significantly between the three experiments. Statistical significance was inferred by Fisher’s exact test (pς.2*10 � ). (B) Relative frequencies of the presence and absence of unpaired supernumerary chromosomes in all progeny ascospores pooled for experiments A, B and C according to the mitochondrial genotype of the ascospore. Orange and blue indicate unpaired chromosomes originating from IPO94269 or IPO323, respectively. Unpaired chromosomes (with the exception of chr14) inherited from the parent that provided the mitochondrial genotype (i.e. the female parent) are overrepresented in the progeny, while the same chromosomes when originating from the male parent are not. Statistical significance was inferred by a two-sided binomial test with a probability of p=0.5. (C) Cell density affects the sexual role during mating and thereby the transmission of the mitochondria. Relative frequencies of mitochondrial genotype in random and randomized ascospores isolated from crosses of IPO94269 and IPO323㥌hr19 that were co-inoculated on wheat at different cell densities. The resulting progeny shows a correlation between cell-density and mitochondria transmission. Strains inoculated at lower density in general take the female role as observed by the mitochondrial transmission. Statistical significance was inferred by a two-sided Fisher’s exact test compared to the co-inoculation with equal cell densities of both strains. (D) The cell density affects the transmission of unpaired chromosomes. Relative frequencies in all ascospores of the presence and absence of unpaired supernumerary chromosomes 19, inherited from parent IPO94269 and unpaired chromosomes 18 and 20, inherited from IPO323 㥌hr19 are indicated according to the cell density of the parental strains IPO94269 and IPO323㥌hr19 at inoculation. Statistical significance was inferred by a two-sided Fisher’s exact test compared to the co-inoculation with equal cell density of both strains. (*=p <਀.05, **=p <਀.005, ***=p <਀.0005, see Supplementary file 5 for details on all statistical tests).

Figure 2𠅏igure supplement 1.

(A) Detailed presence/absence frequencies for all unpaired supernumerary chromosomes according to the mitochondrial genotype present in all ascospores. Data for the three experiments A, B and C are depicted separately. Statistical significance was inferred by a two-sided binomial test with a probability of p=0.5. (B) Detailed presence/absence frequencies for all unpaired supernumerary chromosome in experiment A, B and C, restricted to all co-inoculation crosses using 1*10 7 cells/mL cell density for both parental strains. Experiment A, B and C differ in the observed transmission advantage for the chromosomes originating from the parent IPO94269 (orange) or IPO323 (blue). Statistical significance was inferred by a two-sided binomial test with a probability of p=0.5. (C) Presence/absence frequencies of all unpaired supernumerary chromosomes pooled for all three experiments A, B and C, restricted to all co-inoculation crosses using the 1*10 7 cells/mL cell density for both parental strains. All supernumerary chromosomes, with the exception of chromosome 14 show a highly significant transmission advantage. Statistical significance was inferred by a two-sided binomial test with a probability of p=0.5. (*=p <਀.05, **=p <਀.005, ***=p <਀.0005, see Supplementary file 5 for details on all statistical tests).

Although the transmission of the supernumerary chromosomes was highly similar between experiments A, B and C when the mitochondrial genotype was used to group the data ( Figure 2𠅏igure supplement 1A ), the overall transmission of the supernumerary chromosomes varied considerably between the experiments due to the highly divergent mitochondrial genotype inheritance in the three experiments ( Figure 2𠅏igure supplement 1B ). However, we find a clear transmission advantage for all supernumerary chromosomes, except chromosome 14, when pooling all data from the three experiments (transmission to more than 50% of the progeny) ( Figure 2𠅏igure supplement 1C ). Based on these observations, we conclude that unpaired supernumerary chromosomes show a chromosome drive mechanism, but this drive is restricted to chromosomes inherited from the mitochondria-donating female parent.

Transmission of mitochondria is affected by the cell density

We next asked which factors determine the sexual role and thereby the mitochondrial inheritance in the sexual crosses of Z. tritici. Recently, competition between sexual and asexual modes of reproduction in Z. tritici was shown to be affected by the cell density (Suffert et al., 2018). We therefore hypothesized that the cell density of the two parental strains could also affect the sexual roles of the two parental strains. To test this, we set up crosses between the strains IPO323㥌hr19 and IPO94269 in which the cell density varied from 10 5 cells to 10 7 cells/mL of each of the parental strains. To distinguish the female and male partner in the crosses we again assessed the mitochondrial transmission frequencies. Interestingly, we find that the cell density of the two parental strains strongly correlates with the transmission of the mitochondrial genotype. Crosses with a lower cell density of IPO323㥌hr19 resulted in a higher proportion of the progeny carrying the IPO323 mitochondrial genotype ( Figure 2C ). Similarly, a lower cell density of IPO94269 resulted in a higher proportion of the progeny carrying the IPO94269 mitochondrial genotype. This illustrates that a density-dependent mechanism affects the sexual role of the Z. tritici strains during sexual mating.

Variation in the sexual role in turn affected the transmission of unpaired supernumerary chromosomes. The unpaired chromosome 19 inherited from the parent IPO94269 increased in frequency in the meiotic progeny with increasing frequency of the IPO94269 mitochondrial genotype. Unpaired chromosome 18 and chromosome 20 inherited from the parent IPO323㥌hr19 increased in frequency in the meiotic progeny with an increase in frequency of the IPO323 mitochondrial genotype ( Figure 2D ). Therefore, a clear effect of cell density is discernable and we conclude that environmental factors that affect the infection density of different Z. tritici strains also strongly affect the sexual role of strains and thereby the transmission of supernumerary chromosomes. We hypothesize that variability in environmental factors between the three experiments, which were conducted during different seasons and at different locations, may explain the observed variability in the frequency of sexual roles of the two strains among the three experiments. Accordingly, the transmission of unpaired supernumerary chromosomes may also be affected by seasonal changes in environmental factors.

Paired supernumerary chromosomes show Mendelian segregation with frequent losses

In Z. tritici, as in other ascomycetes, one meiosis produces eight ascospores by an additional mitosis following meiosis (Ni et al., 2011 Wittenberg et al., 2009). The outcome of single meiotic events can be analyzed by tetrad analyses whereby the eight ascospores of a tetrad - in ascomycetes the ascospores contained in an ascus - are isolated and genotyped. We used tetrad analyses to address how paired supernumerary chromosomes segregate during meiosis. For a total of 24 separate asci, we verified that all eight ascospores originated from the same ascus and were the products of a single meiosis using six segregating markers located on the essential chromosomes (Table 2). With these 24 asci we could identify segregation patterns and furthermore eliminate post-meiotic effects on the observed chromosomal frequencies. Each tetrad allowed the analysis of the segregation pattern for both unpaired and paired chromosomes. First, we focused on the segregation of paired chromosomes within these tetrads. In the 24 asci we could observe the transmission of paired supernumerary chromosomes in 129 instances. We could discern the segregation of each paired supernumerary chromosome using specific segregating markers for each of the supernumerary chromosomes from both parental strains. In general, the paired supernumerary chromosomes showed a Mendelian segregation pattern ( Figure 3 ). Of the 129 instances of supernumerary chromosome pairing, 120 (93%) showed a Mendelian segregation pattern with the expected 4:4 ratio ( Figure 3 , black outline) and only nine instances (7%) showed a non-Mendelian transmission pattern deviating from the 4:4 ratio ( Figure 3 , red outline). In no instances did the number of ascospores with a segregating marker for a paired supernumerary chromosome exceed the four ascospores predicted by Mendelian segregation. In two of the nine instances however, we found ascospores with two copies of supernumerary chromosome 21, representing one copy from IPO323 and another from IPO94269. Whole genome sequencing validated the presence of two copies of chromosome 21 in the genomes of these ascospores. In the two additional ascospores of the tetrad analysis the chromosome 21 was missing ( Figure 3𠅏igure supplement 1A𠄼 ) suggesting that the non-Mendelian transmission patterns are due to loss of chromosomes, non-disjunction of sister chromatids, or non-disjunction of homologous chromosomes during meiosis (Wittenberg et al., 2009).

Analysis of segregation of paired supernumerary chromosomes in 24 complete tetrads. The transmission of chromosomes 14, 15, 16, 17, 19, and 21 with homologs in both parental strains IPO94269 and IPO323 was detected using segregating markers for chromosomes inherited from the parental strains IPO94269 (orange) and IPO323/IPO323𢁬hr14-21 (blue) in the eight ascospores originating from 24 asci. In 120 of the 129 cases (black outline) we observed a 4:4 ratio in the progeny. Note: for crosses/chromosome combinations where no paired chromosome was present no ascus is shown, which reduces the number of shown asci from the 24 asci that were analyzed in both Figure 3 and Figure 4 .

Figure 3—source data 1.

Figure 3𠅏igure supplement 1.

Coverage heatmap for eight ascospores (T0325-T0332) of ascus A03-4 (A𠄼) and eight ascospores (T0801-T0808) of ascus A08-1 (D𠄿). (a) Supernumerary chromosome complement in cross resulting in ascus A03-4 between IPO323 and IPO94269. (B) Coverage heatmap of all chromosomes on from ascus A03-4 reflecting similar coverage for essential and supernumerary chromosomes. (C) Coverage heatmap of supernumerary chromosomes of ascus A03-4 indicating constant coverage of chromosome 18 and 20 in all eight ascospores, loss of paired chromosome 15 in four ascospores and non-disjunction of sister-chromatids in meiosis two resulting in two ascospores containing two chromosomes 21 and two corresponding ascospores lacking chromosome 21. (D) Supernumerary chromosome complement in cross resulting in ascus A08-1 between PO323𢁬hr14 and IPO94269. (E) Coverage heatmap of all chromosomes on from ascus A08-1 reflecting similar coverage for essential and supernumerary chromosomes. (F) Coverage heatmap of supernumerary chromosomes of ascus A08-1 indicating constant coverage of chromosome 20 in all eight ascospores, segregating unpaired chromosome 14 inherited from the male parental strain and segregating loss of coverage on right arm of chromosome 18 in four ascospores.

Figure 3𠅏igure supplement 2.

The distribution is consistent with the origin of each tetrad in one meiotic event and Mendelian segregation for core and paired supernumerary chromosomes. (A) Distribution of IPO94269 specific SNP in the eight ascospores of ascus A03-4 mapped on the IPO323 reference genome. Stretches of IPO94269 haplotype alternate with absence of IPO94269 specific SNPs. (B) Detail of distribution of IPO94269 specific SNPs exemplified on core chromosome one and paired supernumerary chromosome 14 of the eight ascospores of ascus A03-4. Recombination events leading to crossover are indicated by blue bars. A total of seven and four crossover involving all sister-chromatids did occur on the core chromosome one and paired supernumerary chromosome 14, respectively. The distribution of the haplotypes is consistent with their origin in one meiotic event. (C) Distribution of the count SNP among the eight ascospores of ascus A03-4 for core and paired supernumerary chromosomes. Mendelian segregation predicts four of the eight ascospores to contain the SNP. Increasing the fidelity of the SNP detetection increased the fraction of SNPs detected in exactly four ascospores. Including all SNP regardless of sequencing coverage in the analsysis results in 68% of the SNPs showing Mendelian segregation. Restricting the analysis to those SNP variants that showed at least in ascospore a sequecning coverage higher than eightfold increased the fraction of SNPs that showed Mendelian segregation to 94%. Further increasing the fidelity of the SNP detection by restricting the analysis to those SNPs that were always detected at a eightfold coverage further increased the fraction of SNPs that showed Mendelian segregation to 97% (ascus A03-4) or 98% (ascus A08-1). (D𠄿) Distribution of the SNPs of core and supernumerary chromosomes in the eight ascospores of the ascus A08-1, analog to A-C).

Whole genome sequencing of two tetrads allowed for the dissection of the inheritance of SNPs on paired supernumerary and core chromosomes. For both paired supernumerary and core chromosomes stretches of haplotypes of parental IPO94269 SNPs can be recognized in the ascospores as a result of recombination and crossover events ( Figure 3𠅏igure supplement 2A𠄿 ). These IPO94269 haplotypes are consistently present in four of the eight ascospores indicating a Mendelian segregation. A more detailed analyses of the SNP distribution showed that the majority of individual IPO94269 SNPs are present in four of the eight ascospores as expected (113135 of a total 167346 SNPs in ascus A03-4 and 115612 of a total 170537 SNPs in ascus A08-1) ( Figure 3𠅏igure supplement 2C and F , Supplementary file 5). However a number of SNPs was found to be present only in three (10.5% and 10.7% of all SNPs in ascus A03-4 and A08-1, respectively), two (8.9% and 9.0% of all SNPs) or one ascospores (12.7% and 12.8% of all SNPs) of a tetrad, which could indicative non-Mendelian transmission for a subset of SNPs located on the core and supernumerary chromosomes. To further investigate the occurrence of non-Mendelian SNPs we restricted our analyses to include only SNPs in regions of the genome alignment with high read coverage. Restricting the analysis to SNPs with ϨX read coverage in at least one ascospore substantially increased the relative frequency of SNPs detected in exactly four ascospores to 94% (96498 of a total 102797 SNPs in ascus A03-4 and 98954 of a total 105024 SNPs in ascus A08-1). Further increasing the fidelity of SNP detection to SNPs at positions with ϨX read coverage at all occurrences further increased the proportion of SNPs detected in exactly four ascospores to 97% or 98% in ascus A03-4 and A08-1, respectively ( Figure 3𠅏igure supplement 2C and F , Supplementary file 5). Very few SNPs on paired supernumerary chromosomes are present in more than four ascospores (69 and 130 SNPs in ascus A03-4 and A08-1, respectively), which could be the result of gene conversion. In conclusion, a similar pattern of the transmission of SNPs was detected for core chromosomes and paired supernumerary chromosomes consistent with Mendelian segregation for the vast majority of SNPs.

We next extended the analysis of transmission fidelity to include all ascospores isolated in experiments A, B and C to compare the rate of loss of paired supernumerary chromosomes. We assessed the rate of chromosome loss from 10078 instances of paired supernumerary chromosomes in isolated meiotic progenies. In 377 cases (3.7%) we found evidence for supernumerary chromosome loss in the ascospores based on the absence of specific chromosome markers (Supplementary file 6). Interestingly, the frequency of loss of paired supernumerary chromosomes varies significantly between the individual chromosomes (χ 2 -Test: exp. A: p=1.96휐 � , exp. B: p=1.72휐 � , exp. C: p=4.18휐 𢄤 ) (Supplementary file 6) with chromosome 16 showing the lowest rate of loss in all three experiments. The frequency of chromosome loss, however, shows no correlation to particular chromosome characteristics like chromosome size or the extent of homology between the chromosomes from IPO323 and IPO94269 ( Figure 1𠅏igure supplement 1B ).

Unpaired supernumerary chromosomes inherited from the female parent show meiotic drive

Using the same 24 complete tetrads we dissected the fate of unpaired chromosomes during single meiotic events. Each tetrad contained between one to three unpaired chromosomes with chromosome 18 and 20 being solely inherited from IPO323 and chromosome 14, 15, 16, 17, and 19 being solely inherited from IPO94269 in crosses performed with the five IPO323 whole਌hromosomeꃞletion strains. In contrast to paired supernumerary chromosomes, unpaired supernumerary chromosomes show distinct segregation distortion ( Figure 4 ) that correlates with mitochondrial transmission unpaired chromosomes originating from the female parent show a strong meiotic chromosome drive. On the other hand, unpaired chromosomes originating from the male parent (i.e. the parent that did not provide the mitochondria) often show a Mendelian segregation pattern and are frequently lost.

Analysis of segregation of unpaired supernumerary chromosomes in 24 complete tetrads according to the mitochondrial genotype. The transmission of chromosomes unique to one of the parental strains and the mitochondrial genotype was detected using chromosomal or mitochondrial markers originating from IPO94269 (orange) and IPO323 (blue) in eight ascospores derived from 24 asci. When IPO323 was the female parent (i.e. the ascospores inherited the mitochondrial genotype of the IPO323 parent) unpaired chromosomes 18 and 20 originating from IPO323 show a strong chromosome drive and are overrepresented in the ascospores. When IPO94269 was the female parent unpaired chromosomes originating from IPO94269 show a strong chromosome drive. Unpaired chromosomes originating from the male parent (i.e. the one not donating the mitochondrial genotype) show Mendelian segregation pattern or are lost. Note: for crosses/chromosome combinations where no unpaired chromosome was present no ascus is shown, which reduces the number of shown asci from the 24 asci that were analyzed in both Figure 3 and Figure 4 .

In the 24 tetrads dissected here, all eight ascospores originating from the same ascus showed the same mitochondrial genotype (Supplementary file 3 & 4) confirming previous results on the uniparental inheritance of mitochondria in Z. tritici (Kema et al., 2018). Isolated ascospores had the mitochondrial genotype of the parent IPO323 in 18 asci, while the ascospores of the remaining six asci showed the IPO94269 genotype, confirming that both parental strains, IPO323 and IPO94269, can act as the female parent during sexual mating with no significant difference between the two strains (two sided binomial test (p=0.5), p=0.25) (Kema et al., 2018). In 18 asci that showed the IPO323 mitochondrial genotype, supernumerary chromosomes 18 and 20, inherited from IPO323 (the female), were unpaired in 15 and 16 meioses, respectively ( Figure 4 ). Chromosome 18 was present in all eight ascospores in 11 of the 15 asci (ascus #1�) instead of the expected four ascospores. In two asci, the chromosome was present in six ascospores (ascus #12�). Chromosome 20 was present in all eight ascospores (ascus #1�) in 15 of the 16 asci and in one ascus in six ascospores (ascus #16). This transmission pattern was reversed for the six asci exhibiting the IPO94269 mitochondrial genotype ( Figure 4 ). Here, the female-inherited unpaired chromosomes from IPO94269 show meiotic drive while the male-inherited unpaired chromosomes from IPO323 show Mendelian segregation pattern or were lost ( Figure 4 ). We validated our PCR-karyotyping by sequencing the genomes of 16 ascospore isolates originating from two asci and mapped the resulting reads to the reference genome of IPO323. For all 16 ascospores we find similar coverage for all essential chromosomes and supernumerary chromosomes ( Figure 3𠅏igure supplement 1 ) and all 16 ascospores show similar coverage for chromosome 18 and 20 verifying the transmission of these chromosomes to the eight ascospores of each ascus instead of the expected four ascospores.

The meiotic drive of the unpaired supernumerary chromosome could imply an additional amplification step that only affects unpaired chromosomes derived from the female parent. We found however that in one cross, this additional amplification of an unpaired chromosome was incomplete: In the ascus A08-1, unpaired chromosome 18 was transmitted to all eight ascospores, but four of the ascospores contained only a partial chromosome 18, ( Figure 3𠅏igure supplement 1F ). The partial chromosomes 18 showed a Mendelian segregation pattern, indicating that the additional amplification step of the unpaired chromosome 18 occurred prior to the first meiotic division, which in this rare case was incomplete.


© 2014 The Authors. Published by the Royal Society under the terms of the Creative Commons Attribution License http://creativecommons.org/licenses/by/4.0/, which permits unrestricted use, provided the original author and source are credited.

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Conclusions

Evolutionary transitions from obligatory outbreeding to predominantly self-fertilising are prominent in both plants and animals. The resulting changes in selective regime confers predictable changes on populations, as hermaphrodites adapt to optimise resource allocation trade-offs for selfing fitness combined with the effects of excess homozygosity and relaxed sexual selection and sexual conflict. In particular, selfing yields a ‘selfing syndrome’ in plants and in animals, with both phenotypic and genomic characteristics. Nevertheless, botanical and zoological traditions have emphasised distinct perspectives to framing evolutionary shifts between biparental and uniparental reproduction: the ‘evolution of selfing’ viewpoint of reproductive assurance, automatic selection transmission advantage, and inbreeding depression vs the ‘evolution of sex’ viewpoint of the cost of males, host−pathogen co-evolution, rates of adaptation and mutation accumulation. A conceptual separation in these traditions is the explicit consideration by plant biologists of how much pollen discounting co-sexual hermaphrodites experience vs an implicit presumption of complete ‘pollen’ discounting in many animals systems. These distinct lenses on reproductive-mode evolution are complementary, with much potential for their synergy to provide more general tests of theory across the tree of life. The genomic and phenotypic divergence resulting from reproductive-mode evolution also intersects with the process of speciation, setting the stage for reproductive isolation. Key among the characters important in selfing-associated speciation is relaxed sexual selection and sexual conflict, in which postmating prezygotic interactions can play a crucial role in both plant and animal systems.


Watch the video: GCSE Biology - Sexual vs Asexual Reproduction - What is Asexual Reproduction? #46 (May 2022).