Wheat univalent orientation at anaphase I in wheat-rye derivatives

The anaphase I behaviour of wheat univalents in plants with the chromosome constitution (0–7)A(0–7)BRR was analyzed using the C-banding technique, which allows to distinguish between wheat and rye chromosomes. The equational division frequencies of univalents observed in the six plants analyzed show a large variation (0.21–0.83). Within each plant syntelic univalents segregate to the poles at random. The frequency distribution of amphitelically dividing univalents does not conform to a random distribution. The lack of fit is attributed to environmental factors which differentially affect the probability of equational division for the univalents in different PMCs. Two other possible causes of the lack of adjustment, namely, each wheat univalent has a different probability of equational division, and wheat univalents do not move independently to the equator to divide equationally, are also discussed. The latter seems improbable in view of the independent behaviour of univalents dividing reductionally. A correlation observed between the behaviour of chromosome 6B and the rest of wheat univalents is attributed to variation between cells due to external causes.     

Differential anaphase-I behaviour between wheat and rye univalents in triticale-wheat hybrid plants

The chromosomes of the D genome of wheat and the genome R of rye can be distinguished at meiosis by C-banding in triticale-wheat hybrid plants. All members of both genomes almost exclusively formed univalents at metaphase I. However, at anaphase I the frequencies of equationally dividing chromosomes were higher for rye than for wheat chromosomes. The differential centromere behaviour at anaphase I is ascribed to differences in the time at which wheat and rye univalents are formed during the first meiotic prophase.     

Meiosis in Mesostoma ehrenbergii ehrenbergii (Turbellaria,Rhabdocoela)

At metaphase I of meiosis in spermatocytes of Mesostoma ehrenbergii ehrenbergii [2n=10] three bivalents and four univalents form. The same two chromosome pairs always form the univalents. Analysis of metaphase I, anaphase I and metaphase II configurations in fixed testis material suggested that the distribution of the four univalents is not a random process but the correct segregation of one member of each pair to each pole is actively achieved before the end of metaphase I. In live preparations of testis material univalents were observed to move between the poles of metaphase I cells, eventually reaching the correct segregation. All cells observed to enter anaphase I had the correct segregation of univalents. It is proposed that the univalent movement during metaphase I is directed towards obtaining the correct segregation of univalents before the cells enter anaphase.     

Meiotic studies in Acanonicus hahni (Stål) (Coreidae,Heteroptera) I. Behaviour of univalents in desynaptic individuals

Male meiosis was studied in a population of Acanonicus hahni (Stål), and nine of the sixteen individuals analyzed showed desynapsis. The frequency of univalents varied from one to seven percent in eight of them, while in the ninth the percentage of cells with univalents was higher (12%). The univalents auto-orientate at metaphase I in the center of the ring formed by autosomal bivalents and divide equationally at anaphase I; at metaphase II they show touch-and-go pairing, and lie in the center of the ring of autosomes.A desynaptic origin of the univalents is proposed, and the arrangement of the chromosomes in the first and second metaphase plate in the normal and desynaptic individuals is compared and discussed. The meiotic characteristics of these desynaptic individuals are also compared with those described in other insects with holocentric and monocentric chromosomes. It is suggested that any achiasmatic chromosome, whether a univalent, m or sex chromosome, will induce the formation of a ring and with some or all of them lying in its centre.     

What determines whether chromosomes segregate reductionally or equationally in meiosis?

Normal meiosis consists of a single round of DNA replication followed by two nuclear divisions. In the 1st division the chromosomes segregate reductionally whereas in the 2nd division they segregate equationally (as they do in mitosis). In certain yeast mutants, a single-division meiosis takes place, in which some chromosomes segregate reductionally while others divide equationally. This autonomous segregation behaviour of individual chromosomes on a common spindle is determined by the centromeres they carry. The relationship between reductional segregation of a pair of chromosomes and their earlier recombinational history is also discussed.     

Studies on triploid Allium triquetrum

This work describes the relationship between the univalents seen at metaphase I and the distribution of dyads at anaphase I in the pollen mother cells of triploid Allium triquetrum. The orientation of the centromeres within the trivalents and bivalents at metaphase I towards the two poles of the pollen mother cells is random. The distribution of polar univalents towards the two poles at metaphase I is also random, as is the distribution of dyads at anaphase I in low univalent frequency collections. However, in a high univalent frequency collection, the distribution of dyads at anaphase I is non-random. There is an excess of cells with the most equal dyad distribution (13–14) and a paucity of cells with a 12–15 distribution. In low univalent frequency collections, the equatorial univalents are believed to remain in the equatorial region during anaphase I and are seen as laggards at late anaphase I. The remaining chromosomes move according to the metaphase I orientation of their centromeres to give a random distribution of dyads at anaphase I. In high univalent frequency collections it is argued that the non-random dyad distribution seen at anaphase I is the result of non-random movement of some of the equatorial univalents away from the equatorial region during anaphase I. The remaining equatorial univalents remain in the equatorial region and are seen as laggards at late anaphase I.     

Desynaptische Mutanten mit Unterschieden im Univalentenverhalten

After application of neutrons on dry seeds ofPisum sativum three recessive mutants were isolated showing irregularities in the course of meiosis. A cytogenetical analysis showed that at metaphase I, a varying number of univalents are formed, most likely as a result of reduced chiasma frequencies. At anaphase I, some univalents divide precociously in mutant 2982 but none do so in mutants 2989 and 2552. As a consequence, most cells of 2989 and 2552 built up more than two spindles at anaphase II, whereas the majority of cells in 2982 form two spindles. This situation is reflected in the frequency distribution of gones per PMC at tetrad stage. The discussion deals with the possible causes of univalent formation at meiotic prophase and the variability of univalent behaviour at anaphase I.     

A comparative study of orientation at behavior of univalent in living grasshopper spermatocytes

Orientational movements and modes of segregation at anaphase I were analyzed in three different types of univalents in living spermatocytes of the grasshopper species Eyprepocnemis plorans, namely the sex univalent, three types of accessory chromosomes and spontaneous and induced autosomal univalents. When two or more univalents were present in the same spindle, their dynamics were directly compared. Chromosomes may show variable velocity and number of reorientations: the X and the most common B types (B₁ and B₂) are slow and rarely reorient, a more geographically restricted B (B₅) is faster and reorients more often, and autosomal univalents are the fastest and show the highest frequency of reorientations. Nonetheless, the X and the accessories are rigorously reductional at anaphase I whereas autosomal univalents often fail to migrate or divide equationally. This indicates that orientational and segregational behavior are controlled mainly by chromosomal rather than cellular characteristics and that chromosomes may display a great variety of strategies to achieve regular segregation.     

Meiotic sister chromatid cohesion in holocentric sex chromosomes of three heteropteran species is maintained in absence of axial elements

It has been suggested that in species with monocentric chromosomes axial element (AE) components may be responsible for sister chromatid cohesion during meiosis. To test this hypothesis in species with holocentric chromosomes we selected three heteropteran species with different sex-determining mechanisms. We observed in surface-spreads and sections using transmission electron microscopy that the univalent sex chromosomes form neither AEs nor synaptonemal complexes (SCs) during pachytene. We also found that a polyclonal antibody recognizing SCP3/Cor1, a protein present at AEs and SC lateral elements of rodents, labels the autosomal SCs but not AEs or SC stretches corresponding to the sex chromosomes. Cytological analysis of the segregational behaviour of the sex univalents demonstrates that although these chromosomes segregate equationally during anaphase I they never show precocious separation of sister chromatids during late prophase I or metaphase I. These results suggest that AEs are not responsible for sister cohesion in sex chromosomes. The segregational behaviour of these chromosomes during both meiotic divisions also indicates that different achiasmate modes of chromosome association exist in heteropteran species. Received: 22 September 1999; in revised form: 20 December 1999 / Accepted: 21 December 1999     

Die biometrische Bewertung des Univalenten-Verhaltens in konjugationsgestörten Mutanten von Pisum sativum

Zusammenfassung In konjugationsgestörten Mutanten treten im Verlaufe der Meiosis Univalente oder Univalente und Bivalente in wechselnden Häufigkeitsverhältnissen zueinander auf. Ihre räumliche Anordnung im Spindelraum der Metaphase I wurde zytologisch untersucht und die Befunde statistisch ausgewertet. Der überwiegende Teil unseres Materials ließ sich einheitlich durch eine Polya-Verteilung charakterisieren. Für deren Parameter wurden — für jeden Genotyp und für jede Univalentenklasse getrennt —Punkt-und Intervallschätzungen nach der Maximum Likelihood-Methode durchgeführt. Aus der Lage der Schätzwerte konnten wir schließen, daß bei allen untersuchten Mutanten die Wahrscheinlichkeit für die Univalente, sich in der Metaphaseplatte einzuordnen, um so größer wird, je mehr Bivalente in der Zelle vorliegen. Die gegenseitige Beeinflussung der Univalente ist bei unseren Mutanten unterschiedlich. Bei einer geringen Anzahl von Bivalenten erhöht jedes zufällig in der Metaphaseplatte angeordnete Univalent die Wahrscheinlichkeit für die übrigen, sich ebenfalls dort einzuordnen. Ist die Zahl der Bivalente erhöht, so nimmt die gegenseitige Beeinflussung der Univalente in einigen Fällen ab, in anderen Fällen bleibt sie lediglich erhalten.Es wurde zum Schluß versucht, die erarbeiteten Befunde als univalentenbedingte strukturelle und funktioneile Änderungen im Aufbau des Spindelapparates zu erklären.

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Biometrical evaluation of univalent behaviour in meiotic mutants of Pisum sativum

Summary Asynaptic and desynaptic mutants are characterized by the appearance of univalents in the course of meiosis. Their spatial arrangement in the spindle apparatus of metaphase I was cytologically investigated and the findings were statistically analysed. The great majority of our material could be fitted to a Polya-distribution. For the parameters of this distribution, point- and intervalestimations were performed by the maximum likelihood method, separately for each genotype and each univalent class. From the position of the estimates in the different classes, it can be concluded that in all genotypes under consideration a high number of bivalents increases the probability of the remaining univalents being arranged in the metaphase plate. The mutual interference of the univalents themselves differs from genotype to genotype: when the number of bivalents is low, each univalent that happens to be arranged in the metaphase plate raises the probability that the remaining univalents will migrate into the metaphase plate, too. In some cases, the mutual interference of the univalents is reduced when the number of bivalents increases; in other cases this interference is merely maintained. Finally, we tried to interpret the cytological and statistical findings by structural and functional changes of the spindle apparatus brought about by the existence of univalents.

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Angenommen durch H. Stubbe     

Mixed Segregation of Chromosomes during Single-Division Meiosis of Saccharomyces Cerevisiae

Normal meiosis consists of two consecutive cell divisions in which all the chromosomes behave in a concerted manner. Yeast cells homozygous for the mutation cdc5, however, may be directed through a single meiotic division of a novel type. Dyad analysis of a cdc5/cdc5 strain with centromere-linked markers on four different chromosomes has shown that, in these meioses, some chromosomes within a given cell segregate reductionally whereas others segregate equationally. The choice between the two types of segregation in these meioses is made individually by each chromosome pair. Different chromosome pairs exhibit different segregation tendencies. Similar results were obtained for cells homozygous for cdc14.     

Asynapsis in Lolium perenne

Eighteen plants displaying varying degrees of asynapsis ranging from weak to very strong were found among four out of six populations of Lolium perenne L. (2n=14) which had been subjected to three cycles of directional phenotypic selection for productivity of green material. No plants were found displaying univalents in the original generation but the incidence increased with cycles of selection, indicating the genetic control and differential distribution of asynaptic genes among these populations. — The analysis of univalents and chiasma frequency of pollen mother cells (PMC) of six partially asynaptic plants chosen for detailed study revealed that univalents occurred throughout all PMC chiasma classes irrespective of chiasma frequency, but the higher the chiasma frequency of any PMC the less the likelihood of univalents occurring. The relationship between chiasma frequency and univalent frequency per PMC per plant was negative. — Mean chiasma frequency per bivalent increased for the asynaptic cells in comparison with the normal in both the weak and medium asynaptic groups which was explained by the availability of additional chiasmata for redistribution.     

Chromosome pairing in meiosis of partially fertile wheat/rye hybrids

The normal course of meiosis depends on regular pairing of homologous chromosomes. In intergeneric hybrids, including those of wheat, there is no chromosome pairing because there are no homologs. In F₁ wheat/rye hybrids, pairing is largely prevented by the pairing homoeologous1 (Ph1) gene. In its presence, there are only rare instances of pairing; most chromosomes are univalent, and their orientation at metaphase I initiates different pathways of the meiotic cycle. The meiotic-like pathway includes a combination of the reductional and the equational + reductional steps at AI followed by the second division. The resulting gametes are mostly non-functional. The mitotic-like pathway involves equational division of univalents at AI and the absence of the second division. Any fertility of wheat/rye hybrids depends on the production of unreduced gametes arising from meiotic restitution (mitotic-like division). We examined the meiotic pairing in wheat/rye hybrids created from wheat lines with single rye chromosome substitutions and Ph1 present. This guaranteed F₁ meiosis with one pair of rye homologs. All hybrids formed bivalents, but proportions of meiocytes with bivalents varied. In the meiocytes where bivalents were present, there was a higher tendency for the meiotic-like pathway, while in meiocytes where bivalent pairing failed, the tendency was stronger for the mitotic-like pathway. Among the equationally dividing cells, we observed more than 90 % of meiocytes without bivalents, where rye homologs did not form bivalents, too. The data indicate a potential application of wheat/rye lines in producing genetic stocks of amphidiploids with designated genomic constitutions.     

Sex-dependent frequency and type of autosomal univalency at the first meiotic metaphase in mouse germ cells

Univalents at the first meiotic metaphase in mouse spermatocytes occur mainly in the XY pair, making it difficult to compare the amounts of univalency in males and females. In this study, the amounts of autosomal univalency in male and female meiosis were compared using the model strain CBA-T6, in which univalency of the small marker autosome pair T6 has been shown to occur very frequently in spermatocytes. Mice from inbred CBA and DBA strains were also analysed. The total frequencies of univalency (sex chromosomes plus autosomes) in metaphase I spermatocytes were 45.6% in CBA, 36.9% in CBA-T6, and 37.3% in DBA males. The aneuploidy in metaphase II spermatocytes ranged from 1.4 to 3% in these strains, which was in agreement with previous findings that most primary spermatocytes with abnormal chromosome configurations are arrested in their development before metaphase II. In the CBA-T6 strain, autosomal univalency at metaphase I mostly involved chromosome pair T6; however, its frequency differed significantly between the sexes, amounting to 18.9% in spermatocytes and 4.3% in oocytes. In the CBA strain, autosomal univalents at metaphase I were seen in 7.7% of the spermatocytes and 1.4% of the oocytes and, in DBA mice, in 4.9% of the spermatocytes and 3.8% of the oocytes. However, in DBA oocytes, when univalency occurred it usually concerned a greater number of bivalents in one cell (range: 2-19 disjoined bivalents), a phenomenon very rare in males of this strain. This study shows that univalent formation differs between the male and female types of meiosis.     

Meiotic segregation of chromosomes in Drosophila melanogaster oocytes

A new technique was developed for a light microscopic analysis of meiosis in Drosophila oocytes. — When the nuclear envelope breaks down the bivalents, till then compressed into a karyosome, separate in early prometaphase. The homologues remain associated by chiasmata except for the fourth chromosomes which are no longer associated. Non-homologous chromosomes regularly segregating from each other in genetic experiments are also unconnected after karyosome disintegration but during metaphase I the fourth chromosomes and the heterologous pairs coorient on the same arc of the spindle and move precociously towards opposite poles. Nondisjunction and other irregularities are not infrequent in oocytes having an uneven number of achiasmatic elements. The fourth chromosomes and the Xs or the large autosomes, when lacking chiasmata, may be involved in non-homologous segregation. In c3G homozygotes all chromosomes appear as univalents in prometaphase. Segregation is variable but the observations suggest the polar distribution of equal numbers of chromosomes in variable combinations irrespective of the size. — Coorientation of univalents may be accounted for if the centromeres, whether homologous or non-homologous, are associated in pairs during early meiotic prophase, and that in the karyosome these pairing relationships are preserved until spindle organization at the onset of prometaphase.     

Involvement of chromatid cohesiveness at the centromere and chromosome arms in meiotic chromosome segregation: A cytological approach

Kinetochores and chromatid cores of meiotic chromosomes of the grasshopper species Arcyptera fusca and Eyprepocnemis plorans were differentially silver stained to analyse the possible involvement of both structures in chromatid cohesiveness and meiotic chromosome segregation. Special attention was paid to the behaviour of these structures in the univalent sex chromosome, and in B univalents with different orientations during the first meiotic division. It was observed that while sister chromatid of univalents are associated at metaphase I, chromatid cores are individualised independently of their orientation. We think that cohesive proteins on the inner surface of sister chromatids, and not the chromatid cores, are involved in the chromatid cohesiveness that maintains associated sister chromatids of bivalents and univalents until anaphase I. At anaphase I sister chromatids of amphitelically oriented B univalents or spontaneous autosomal univalents separate but do not reach the poles because they remain connected at the centromere by a long strand which can be visualized by silver staining, that joins stretched sister kinetochores. This strand is normally observed between sister kinetochores of half-bivalents at metaphase II and early anaphase II. We suggest that certain centromere proteins that form the silver-stainable strand assure chromosome integrity until metaphase II. These cohesive centromere proteins would be released or modified during anaphase II to allow normal chromatid segregation. Failure of this process during the first meiotic division could lead to the lagging of amphitelically oriented univalents. Based on our results we propose a model of meiotic chromosome segregation. During mitosis the cohesive proteins located at the centromere and chromosome arms are released during the same cellular division. During meiosis those proteins must be sequentially inactivated, i.e. those situated on the inner surface of the chromatids must be eliminated during the first meiotic division while those located at the centromere must be released during the second meiotic division.by D.P. Bazett-Jones     

Chromosome distribution and catenation in Rhoeo spathacea var. concolor

The twelve chromosomes of Rhoeo spathacea variety concolor are arranged in a definite sequence in a ring at meiosis. Identification of all the 12 chromosomes was possible in 119 diakinesis and metaphase I cells. — Pollen viability was measured to be 36.54% by cotton blue staining procedure. Forty five of 56 metaphase I cells (80.36%) had adjacent distribution. Each of the 12 chromosomes was equally likely to be involved in adjacent distribution regardless of their sizes and heterobrachialness. Adjacent distribution occurred randomly at each arm-position in the ring regardless of the lengths of the arm-pairs. — The most frequent chromosome configuration at diakinesis and metaphase I was a chain-of-12 chromosomes (41.18%). Cells with 1 to 4 chains of chromosomes were observed. The observed frequencies of various configurations were in good agreement with the calculated frequencies. The mean number of chiasmata was 10.90 per cell and 0.908 per pair of chromosome arms. The 131 chiasma failures were distributed at random among the 12 arm-positions. Since the lengths of arm-pairs in the ring vary, the randomness may mean that chiasma formation was limited to short terminal segments on all chromosomes.     

Meiotic chromosomal aberrations in wild populations of Podophyllum peltatum

Meiotic chromosomal aberrations observed in wild populations of the plant Podophyllum peltatum include incomplete homologous pairing, non-homologous pairing, and inversion heterozygosity in pachytene; univalents, asymmetrical bivalents, and translocation heterozygosity in metaphase-I; bridge and fragments in anaphase-I; and non-disjunction as detected in anaphase-II. Incomplete homologous pachytene pairing is believed to result in non-homologous pairing and in the formation of metaphase-I univalents. The unequal distribution and precocious division of univalents in anaphase-I leads to non-disjunction. Non-disjunction chromosomes (varying in frequency from 0.0 to 24.6%) appear to be distributed among the genome on the basis of chromosome length. Asymmetrical bivalents and anaphase-I side-arm bridges are believed to be caused by chromatid breakage and fusion rather than inversion heterozygosity. Of the 135 clones examined, 20 were found to be heterozygous for translocations. The possibility of widespread distribution of some translocations is suggested.     

Maternal ageing and nondisjunction: A comparative study of two chromosomal techniques on the formation of univalents in first meiotic metaphase oocytes of the mouse

The incidence of univalents was compared between slides prepared according to two clearly different chromosomal methods, i.e. Tarkowski's method and ours, in order to examine whether a univalent pair could be formed artifactually at the first meiotic metaphase (MI). The oocytes used were obtained from young (2–3 months) and old (12–15 months) age groups of both C57BL/6 and dd mice. In Tarkowski's method only a single fixative was used, while in our method three different fixatives were used successively in order to fix oocytes without their being ruptured. Despiralized, fuzzy and loosely associated chromatids were seen frequently in the slides prepared by Tarkowski's method, while such features were seen less frequently in the slides prepared by our method. The incidence of oocytes with univalents in the slides made by Tarkowski's method was much higher than in those made by ours in both age and strain groups (P<0.05–0.001). Thus, it was confirmed that the so-called univalents could be produced artifactually. The results did not support the production line hypothesis of Henderson and Edwards (1968) which was based on their observation of an increased incidence of univalents in MI oocytes from aged female mice.     

Indeterminate meiotic restitution (IMR): a novel type of meiotic nuclear restitution mechanism detected in interspecific lily hybrids by GISH

A detailed analysis of microsporogenesis was carried out in three diploid lily cultivars (2n=2x=24) and three diploid interspecific hybrids (2n=2x=24) using DNA in situ hybridisation methods (GISH and FISH). In cvs. Gelria (Lilium longiflorum; L genome), Connecticut King and Mont Blanc (both Asiatic hybrids; Agenome) meiosis was regular and only haploid gametes were formed while the three interspecific hybrids between L. longiflorum×Asiatic hybrid (LA) showed a variable frequency of meiotic nuclear restitution and stainable 2n-pollen formation ranging from 3% to 30%. An analysis of meiotic chromosome behaviour of the LA hybrids through GISH and FISH revealed that: (1) the parental chromosomes could be clearly discriminated into univalents, half-bivalents and bivalents in the PMCs; (2) in some of the PMCs the entire complement was present either as univalents or half-bivalents which had the potential to divide equationally (following centromere division) during the first division leading to first division restitution (FDR) gametes; (3) more frequently, however, in one and the same PMC the univalents and half-bivalents divided equationally whereas the bivalents disjoined reductionally at the same time giving rise to 2n-gametes that could vary from the well-known FDR or SDR 2n-gametes. We indicate this novel type of restitution mechanism as Indeterminate Meiotic Restitution (IMR). In order to confirm the occurrence of IMR gametes, the chromosome constitutions of eight triploid BC₁ progenies derived from backcrossing the 2n-gamete producing the LAhybrids to the Asiatic hybrid parents were analysed through in situ hybridisation. The results indicated that there were seven BC₁ plants in which FDR 2n-gametes, with or without homoeologous recombinations, were functional, whereas in one case the 2n-gamete resulting from IMR was functional. In the latter, there was evidence for the occurrence of genetic recombination through homoeologous crossing-over as well as through the assortment of homoeologous chromosomes. A singular feature of the IMR 2n-gamete was that although it transmitted a euploid number of 24 chromosomes to the BC₁ progeny, the number of chromosomes transmitted from the two parental species was dissimilar: 9 L-genome chromosomes and 15 A-genome chromosomes instead of 12 of each. Received: 15 May 2000 / Accepted: 4 December 2000