Role of rye chromosome 2R from wheat-rye substitution line 2R(2D)1 ( Triticum aestivum L. cv. Saratovskaya 29- Secale cereale L. cv. Onokhoiskaya) in genetic regulation of meiotic restitution in wheat-rye polyhaploids

A study was made of the role of rye chromosome 2R from the wheat-rye substitution line 2R(2D)₁ (Triticum aestivum L. cv. Saratovskaya 29-Secale cereale L. cv. Onokhoiskaya) in genetic regulation of meiotic restitution in wheat-rye polyhaploids 2R(2D)₁ × S. cereale L. cv. Onokhoiskaya. Rye chromosome 2R proved to affect the completeness of the meiotic program, suppressing the formation of restitution gametes. This was evident from the reductional division of univalent chromosomes in AI and the occurrence of the second meiotic division. The interrelationships between the type of chromosome division in AI and the two-step character of meiosis are discussed. The structural and functional organization of the centromeric regions of chromosomes undergoing reductional division is assumed to determine the two-step character of division. Original Russian Text ? O.G. Silkova, A.I. Shchapova, V.K. Shumny, 2007, published in Genetika, 2007, Vol. 43, No. 7, pp. 971–981.     

FORMATION OF 2N MEGAGAMETOPHYTES IN DIPLOID TUBER-BEARING SOLANUMS

Development of megaspores and megagametophytes was analyzed for several diploid potato clones (Solanum spp.) that exhibit either high (HI) or low (LO) seed set when crossed as female with the tetraploid cultivated potato S. tuberosum Group Tuberosa. The objectives were to determine the relationship between ploidy and diam of nuclei and nucleoli, and to determine the mechanism(s) and frequencies of 2n megagametophyte formation. Sizes of nuclei and nucleoli were found to depend on ploidy. For HI clones, the distributions of sizes indicated that doubling occurred during meiosis, and that 30 to 50% of the megaspores and megagametophytes were 2n rather than haploid. Omission of the second meiotic division led to formation of second division restitution (SDR) 2n megagametophytes. Only one HI clone had abnormal meiosis I, in addition to omission of meiosis II in some meiocytes; this clone seemed to produce not only 1n and 2n, but also 4n megagametophytes. The results indicated that high crossability of the HI clones as female with tetraploids largely was due to formation of SDR 2n megagametophytes, a finding strongly supporting the hypothesis that sexual polyploidization is the driving force behind polyploidization of Solanums. The results contribute to increasing evidence that meiotic mutants and abnormalities play an important role in angiosperm evolution.     

The B-chromosome system of Hypochoeris maculata

The meiotic process in PMCs of Hypochoeris maculata is progressively disrupted in the presence of two or more B-chromosomes. Bivalent formation and chiasma conditions are unaffected by up to 3Bs although some univalence occurs with higher numbers. Spindle behaviour, however, is inefficient at both first and second division in the presence of two or more Bs. At metaphase I, regular equatorial alignment breaks down and A-bivalents sometimes show amphitelic or monosyntelic orientation. Anaphase I is characterised by irregular segregation, equatorial laggards and centric division products. The proportion of normal anaphase segregations declines by 20% for every B more than one. A-chromosome laggards, but not Bs, can induce nuclear restitution at telophase I. Following centric division at anaphase II poleward movement can fail leading to further nuclear restitution. Telophase II nuclei thus can be approximately haploid, diploid or tetraploid with aneuploid variation around the haploid and diploid levels. The frequency of numerical mutants in the offspring indicates that EMC meiosis is much less susceptible to the presence of Bs than PMC meiosis: only 4- and 5B^t plants have an enhanced frequency of numerically-aberrant offspring. The deleterious effects of Bs on meiotic efficiency will contribute to setting an upper limit on B-numbers in natural populations of this species.     

Meiotic restitution in amphihaploids in the tribe Triticeae

In haploid and diploid organisms of the plant kingdom, meiotic division of diploid cells proceeds in two consecutive stages, with DNA replicating only once. In amphihaploids (interspecific or intergeneric hybrids), where homologs are absent, the reduction of the chromosome number does not occur, meiosis is abnormal, and the plants are sterile. Gamete viability in F₁ hybrids is ensured by a single division when chromosomes are separated into sister chromatids in either the first or the second division. Such gametes ensure partial fertility of amphihaploids, thereby facilitating their survival and stabilization of the polygenome. The frequency of the formation of viable gametes varies from a few cases to 98.8% in different anthers of the hybrids. Here, studies on the cytological mechanisms and genetic control of chromosome unreduction or restitution in different amphihaploids of the tribe Triticeae are reviewed. The current notions on the control of formation of restitution nuclei based on the principles of a prolonged metaphase I and different types of meiocytes. The main terms used for systematization of restitution mechanisms are first-division restitution (FDR), single-division meiosis (SDM), and unreductional meiotic cell division (UMCD). It has been assumed that archesporial cells of wide hybrids may have two cell division programs, the meiotic and the mitoyic ones The possible approaches to the analysis of the genetic control of chromosome restitution in amphihaploids are discussed.     

Features of the regulation of meiotic restitution in androgenic haploids of wheat-rye substitution lines 2R(2D)1, 2R(2D)3, and 6R(6A) (Triticum aestivum L., cultivar Saratovskaya 29/Secale cereale L., cultivar Onokhoiskaya)

Regulation of meiotic restitution in androgenic haploids generated by cultivation of isolated anthers of three wheat-rye substitution lines 2R(2D)₁, 2R(2D)₃, and 6R(6A) (Triticum aestivum L., cultivar Saratovskaya 29/Secale cereale L., cultivar Onokhoiskaya) was studied. The presence of rye chromosomes and the absence of homeologous wheat chromosomes in the haploid plant genome was shown to cause meiotic restitution, as observed in the case of androgenic haploids 6R(6A), or to inhibit it—in meiosis of haploids 2R(2D)₁ and 2R(2D)₃. In haploids of lines 2R(2D)₁ and 2R(2D)₃, the reductional type of division of univalent chromosomes was observed, leading to preferential formation of tetrads. In haploids of line 6R(6A), the equational type of division of univalents into sister chromatids, resulting in the block of the second division and formation of diads in approximately 50% of cells, was detected. These results confirm data on the effect of the genotype of line 2R(2D)₁ on the induction of reductional type division of univalents and two-phase meiosis, which were earlier obtained in studies of meiosis in polyhaploids 2R(2D)₁ × S. cereale L., cultivar Onokhoiskaya.     

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     

The occurrence and frequency of 2n pollen in three diploid solanums from Northwest Argentina

Summary A group of wild, tuber-bearing species from Northwest Argentina, belonging to the series Tuberosa, Solarium spegazzini Bitt. (spg, 2n=2x=24), S. gourlayi Hawkes (grl, 2n=2x=24 and 2n=4x=48) and S. oplocense Hawkes (opl, 2n=6x=72), and Cuneolata, S. infundibuliforme Phil (ifd, 2n=2x=24), is being used to investigate the mode of origin of polyploids in the genus Solanum. 2n gametes have been detected in the diploid species ifd and spg and in a diploid race of grl, using cytological and breeding approaches. Twenty-two introductions of spg, 8 of grl and 26 of ifd have been tested for 2n pollen; 59%, 63% and 54% of them, respectively, had at least one 2n pollen producing plant. These introductions comprised 238, 76 and 235 plant respectively, of which 20, 16, and 32 plant produced 5% or more 2n pollen. The mechanism of 2n pollen formation was determined in several plant of 2x spg, 2x grl and 2x ifd. All of them were found to form diplandroids via parallel spindles. This mechanism, which gives meiotic products genetically equivalent to first division restitution gametes, is under control of the Mendelian recessive ps. The results suggest that the allele ps is widely distributed in natural populations of the three diploids, and that its frequency is very high. These species are seen as valuable material for population genetic studies, and for the eventual incorporation into a breeding scheme involving sexual polyploidization via 2n gametes.     

Formation of first division restitution (FDR) 2n-megaspores through pseudohomotypic division in ds-1 (desynapsis) mutants of diploid potato: routine production of tetraploid progeny from 2xFDR × 2xFDR crosses

Summary The level and mode of 2n megaspore formation was studied in full-sib diploid potato clones with either normal or desynaptic (ds-1ds-1) meiosis. Cytological analysis revealed that functional 2n megaspores produced by normal and desynaptic clones originate exclusively from second division restitution (SDR) and first division restitution (FDR), respectively. SDR 2n megaspores resulted from the omission of the second meiotic division following chromosome doubling after anaphase I, whereas FDR 2n megaspores resulted from a direct equational division of univalent chromosomes at anaphase I (pseudohomotypic division). Comparative data strongly indicated that the observed mechanisms of SDR and FDR 2n megaspore formation are extremes of a continuum that is being brought about by common genes for precocious chromosome division. Depending on the relative timing of cell cycle and chromosome division, this precocious chromosome division may impose postreductional (SDR) or prereductional (FDR) restitution of the sporophytic chromosome number under normal synaptic and desynaptic conditions, respectively. The observed frequencies of 2n megaspores closely correlated with seed set, following pollination by tetraploid varieties and by desynaptic diploid clones with exclusive FDR 2n pollen formation. Up to 54.0 and 21.5 seeds/ fruit were obtained from normal synaptic (SDR) and desynaptic (FDR) progeny, respectively. The high frequency of segregants with either SDR or FDR 2n megaspore formation (78.0 and 45.2%, respectively) supports the hypothesis that sexual polyploidization is the driving force behind the origin and evolution of polyploid Solanum species. The present identification of diploid potato clones with consistent FDR 2n megaspore formation extends the opportunities for direct transfer of enhanced diploid germ plasm to tetraploids, and particularly advocates the feasibility of 2x(ds-1; FDR)×2x(ds-1; FDR) breeding schemes in cultivar development and the production of relatively vigorous and uniform true potato seed (TPS) varieties. Its potential value and limitations for breeding and the experimental induction of diplosporic apomixis are discussed.     

Two novel meiotic restitution mechanisms in haploid maize (<Emphasis Type="Italic">Zea mays</Emphasis> L.)

Two original mechanisms of nuclear restitution related to different processes of meiotic division of pollen mother cells (PMCs) have been found in male meiosis of the lines of maize haploids no. 2903 and no. 2904. The first mechanism, which is characteristic of haploid no. 2903, consists in spindle deformation (bend) in the conventional metaphase-anaphase I. This leads to asymmetric incomplete cytokinesis with daughter cell membranes in the form of incisions on the mother cell membrane. As a result, the chromosomes of the daughter nuclei are combined into a common spindle during the second meiotic division, and a dyad of haploid microspores is formed at the tetrad stage. The frequency of this abnormality is about 50%. The second restitution mechanism, which has been observed in PMCs of haploid no. 2904, results from disturbance of the fusion of membrane vesicles (plastosomes) at the moment of formation of daughter cell membranes and completion of cytokinesis in the first meiotic division. This type of cell division yields a binuclear monad. In the second meiotic division, the chromosomes of the daughter nuclei form a common spindle, and meiosis results in a dyad of haploid microspores. The frequency of this abnormality is as high as 15%. As a result, haploid lines no. 2903 and no. 2904 partly restore fertility.     

Two novel meiotic restitution mechanisms in haploid maize (Zea mays L.)

Two original mechanisms of nuclear restitution related to different processes of meiotic division of pollen mother cells (PMCs) have been found in male meiosis of the lines of maize haploids no. 2903 and no. 2904. The first mechanism, which is characteristic of haploid no. 2903, consists in spindle deformation (bend) in the conventional metaphase-anaphase I. This leads to asymmetric incomplete cytokinesis with daughter cell membranes in the form of incisions on the mother cell membrane. As a result, the chromosomes of the daughter nuclei are combined into a common spindle during the second meiotic division, and a dyad of haploid microspores is formed at the tetrad stage. The frequency of this abnormality is about 50%. The second restitution mechanism, which has been observed in PMCs of haploid no. 2904, results from disturbance of the fusion of membrane vesicles (plastosomes) at the moment of formation of daughter cell membranes and completion of cytokinesis in the first meiotic division. This type of cell division yields a binuclear monad. In the second meiotic division, the chromosomes of the daughter nuclei form a common spindle, and meiosis results in a dyad of haploid microspores. The frequency of this abnormality is as high as 15%. As a result, haploid lines no. 2903 and no. 2904 partly restore fertility.     

Role of rye chromosome 2R from wheat-rye substitution line 2R(2D)1 (Triticum aestivum L. cv. Saratovskaya 29-Secale cereale L. cv. Onokhoiskaya) in genetic regulation of meiotic restitution in wheat-rye polyhaploids

A study was made of the role of rye chromosome 2R from the wheat-rye substitution line 2R(2D)1 (Triticum aestivum L. cv. Saratovskaya 29-Secale cereale L. cv. Onokhoiskaya) in genetic regulation of meiotic restitution in wheat-rye polyhaploids 2R(2D)1 x S. cereale L. cv. Onokhoiskaya. Rye chromosome 2R proved to affect the completeness of the meiotic program, suppressing the formation of restitution gametes. This was evident from the reductional division of univalent chromosomes in AI and the occurrence of the second meiotic division. The interrelationships between the type of chromosome division in AI and the two-step character of meiosis are discussed. The structural and functional organization of the centromeric regions of chromosomes undergoing reductional division is assumed to determine the two-step character of division.     

AFLP-based estimation of 2n gametophytic heterozygosity in two parthenogenetically derived dihaploids of Rosa hybrida L

Two dihaploid Rosa hybrida L. genotypes, derived through parthenogenesis by using irradiated pollen, were crossed with clonally propagated plants of the diploid species Rosa rugosa Thunb. and Rosa wichuraiana Crép., respectively. Three progeny groups were obtained which contained numerous polyploids, as determined by flow cytometry. Production of fertile 2n female gametes is apparently very common in one of these R. hybrida dihaploid derivatives, whereas the other one is able to produce fertile 2n pollen. Hence, an amplified fragment length polymorphism (AFLP) study was performed on the parental plants and the resulting hybrid offspring in order to estimate (1) the respective genomic parental contributions, and (2) the level of heterozygosity transmitted by the 2n unreduced gametes. Comparison of the levels of transmitted parental heterozygosity revealed that two types of 2n gametes were produced simultaneously, presumably resulting from restitution at the first and at the second meiotic division, respectively. Received: 15 February 2001 / Accepted: 22 May 2001     

Female reproductive development and pollen tube growth in diploid genotypes of<Emphasis Type="Italic"> Solanum cardiophyllum</Emphasis> Lindl.

We have characterized female gametophyte (megagametophyte) development and the kinetics of pollen tube growth in self-pollinated diploid genotypes (2n=2x=24) of Solanum cardiophyllum Lindl. that show normal seed formation. In this species megasporogenesis and megagametogenesis give rise to a female gametophyte of the Polygonum type composed of two synergids, an egg cell, a binucleated central cell and three antipodals; however, asynchronous abnormalities resembling mechanisms that prevail during the formation of second division restitution gametes were observed. In self-pollinated pistils at least 1–2% of germinating pollen tubes were able to reach the megagametophyte 60–84 hours after pollination (hap). Although the egg cell acquired a zygote-like morphology 60–84 hap, division of the primary endosperm nucleus was only observed 84 hap. The analysis of genetic variability in full-sib progeny confirmed that seeds are derived from sexual reproduction. These observations suggest that diploid genotypes of S. cardiophyllum can serve as an ideal system to genetically investigate true seed formation in a tuber-bearing Solanum species.     

Effects of several meiotic mutations on female meiosis in maize

A modified enzyme digestion technique of ovary isolation followed by staining and squash preparation has allowed us to observe female meiosis in normal maize meiotically dividing megaspore mother cells (MMCs). The first meiotic division in megasporogenesis of maize is not distinguishable from that in mi-crosporogenesis. The second female meiotic division is characterized as follows: (1) the two products of the first meiotic division do not simultaneously enter into the second meiotic division; as a rule, the chalazal-most cell enters division earlier than the micropylar one, (2) often the second of the two products does not proceed with meiosis, but degenerates, and (3) only a single haploid meiotic product of the tetrad remains alive, and this cell proceeds with three rounds of mitoses without any intervening cell wall formation to produce the eight-nucleate embryo sac. This technique has allowed us to study the effects of five meiotic mutations (aml, aml-pral, afdl, dsy *-9101, and dvl) on female meiosis in maize. The effects of the two alleles of the aml gene (aml and aml-pral) and of the afdl and dsy *-9101mutations are the same in both male and female meiosis. The aml allele prevents the entrance of MMCs into meiosis and meiosis is replaced by mitosis; the aml-pral permits MMCs to enter into meiosis, but their progress is stopped at early prophase I stages. The afdl gene is responsible for substitution of the first meiotic (reductional) division by an equational division including the segregation of sister chromatid centromeres at anaphase I. The dsy * -9101 gene exhibits abnormal chromosome pairing; paired homologous chromosomes are visible at pachytene, but only univalents are observed at diakinesis and metaphase I stages. These mutation specific patterns of abnormal meiosis are responsible for the bisexual sterility of these meiotic mutants. The abnormal divergent shape of the spindle apparatus and the resulting abnormal segregation of homologous chromosomes observed in micro-sporogenesis in plants homozygous for the dv1 mutation have not been found in meiosis of megasporogenesis. Only male sterility is induced by the dv1 gene in the homozygous condition. © 1993 Wiley-Liss, Inc.     

Production and characterization of intertribal somatic hybrids of Raphanus sativus and Brassica rapa with dye and medicinal plant Isatis indigotica

Intertribal somatic hybrids of Raphanus sativus (2n = 18, RR) and Brassica rapa spp. chinensis (2n = 20, AA) with the dye and medicinal plant Isatis indigotica (2n = 14, I I) were firstly obtained by polyethylene glycol-induced symmetric fusions of mesophyll protoplasts. One mature hybrid with R. sativus established in field had intermediate morphology but was totally sterile. It had the expected chromosome number (2n = 32, RRI I) and parental chromosomes were distinguished by genomic in situ hybridization (GISH) analysis, and these chromosomes were paired as 16 bivalents in pollen mother cells (PMCs) at diakinesis and mainly segregated equally as 16:16 at anaphase I (A I), but the meiotic disturbance in second division was obvious. Five mature hybrids with B. rapa established in field were morphologically intermediate but showed some differences in phenotypic traits and fertility, two were partially fertile. Cytological and GISH investigations revealed that these hybrids had 2n = 48 with AAIIII complement and their PMCs showed normal pairing of 24 bivalents and mainly equal segregation 24:24, but meiotic abnormalities of lagging chromosomes and micronuclei appeared frequently during second divisions. AFLP analysis showed that all of these hybrids had mainly the DNA banding pattern from the addition of two parents plus some alterations. Some hybrids should be used for the genetic improvement of crops and the dye and medicinal plant.     

Chromosome association and pollen fertility of parental and interspecific hybrids of Lycopersicon esculentum X L. hirsutum and L. pennellii

Cytological studies were carried out on two wild species (L. hirsutum and L. pennellii) and the cultivated species (L. esculentum) of tomato and their F1 hybrids. Both parents and hybrids show a diploid chromosome number of 2n=24. The meiotic behaviour of the cultivated species showed a high degree of chromosome homology resulting in a high level of chiasmata frequency per bivalent. In contrast, the two wild species showed a slight increase in uniyalent frequency and a decrease in bivalent formation and chiasmata frequency. The meiotic behaviour of the hybrids showed a high level of univalents and low levels of bivalents as well as trivalents. Highly significant decreases in chiasmata frequency and increases in meiotic abnormalities, especially in the L. esculentum X L. pennellii hybrid, also were detected. The high meiotic irregularity and low chiasmata frequency recorded in the second hybrid indicated the disharmony and difference between its parental genomes and also served to predict its sterility. With regard to degree of pairing recorded in the hybrids, there is a possibility that sterility in such cases may refer to genetic factors in addition to the previously mentioned reasons. Pollen fertility showed no great difference between L. esculentum and L. hirsutum and their F1 hybrid, but a significant decrease was recorded in the L. esculentum X L. pennellii hybrid, which was clearly associated with high meiotic irregularity, low chiasmata frequency and chromosome association.     

Cytomorphological study of Scutellaria platystegia (Lamiaceae) in Iran

Chromosome number, meiotic behaviour and morphological characters related to habit were studied in 10 populations of Scutellaria platystegia Juz. from S. sect. Lupulinaria native to Iran. All populations are diploid and has the chromosome number 2n = 2x = 22, which is consistent with the proposed base number of x = 11. This taxon displayed regular bivalent pairing and chromosome segregation at meiosis. However, some meiotic abnormalities observed included various degrees of fragmented chromosomes, laggards and bridges in anaphase I to telophase II, precocious division of centromeres in metaphase I or II, asynchronous nucleus and cytomixis. We evaluated and determined the population limits within S. platystegia, employing multivariant statistics. We found a striking association between meiotic behaviour and gross morphology. (© 2013 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Abnormal meiosis in tetraploid genotypes ofBrachiaria brizantha (Poaceae) induced by colchicine: its implications for breeding

Meiotic behavior was analyzed in 6 progenies from 3 artificially induced tetraploid (2n = 4x = 36) sexual genotypes (C31, C41, and C48) of the normally apomicticBrachiaria brizantha (Hochst. ex A. Rich.) Stapf., syn.Urochloa brizantha (Hochst. ex A. Rich.) R. Webster. These are key plants to allow intraspecific hybridization of this important forage species, widely used for pastures in the tropics. The percentage of abnormal cells among the plants ranged from 39.8% to 63.2%. In the single plant derived from C48, only the common meiotic abnormalities typical of polyploids were observed, while in plants derived from C31 and C41, a distinct behavior was found. In the majority of cells of those plants, the chromosomes remained scattered in the cytoplasm in the first division, without forming a metaphase plate. This abnormality blocked chromosome movements at anaphase I. Several micronuclei of various sizes were formed and, after the occurrence of an irregular first cytokinesis, the meiocytes progressed normally to the second division, generating polyads with unbalanced microspores. Pollen viability was not correlated with meiotic abnormalities. The importance of these findings to theBrachiaria breeding program is discussed. The sexual progeny of C48 seems most suitable as female parents to be used in intra-and interspecific hybridization.     

Puzzling Chromosome Behavior in Triploid Females of Drosophila melanogaster

Based on a particular formation of the chromocenter and trivalents in triploid Drosophila females, as well as on asynapsis in pericentromeric regions (which is a result of trivalent competition), an explanation for the increased frequency of crossing over and nonrandom segregation of the X chromosomes and autosomes in the first meiotic division is suggested. It is proposed that a delay in pairing of the pericentromeric heterochromatic chromosome regions combined into a single chromocenter leads to the following: (1) formation of the heteroduplex structures (X structures) takes more time and, consequently, their number and the frequency of crossing over in the paired chromosome regions increases; (2) in nonhomologous chromosomes, the chromocentral connections, which normally degrade in prometaphase, are retained to fulfill a function of coorientation during the first meiotic division.     

A cinematographic study of meiosis in salamander spermatocytes in vitro

Summary Salamander spermatocytes were isolated in a modified Eagle's medium in Rose Chambers. The behavior of the spermatocytes during meiosis was recorded on a time lapse, phase contrast film. The two meiotic divisions progressed without visible irregularities in freshly isolated spermatocytes. Times required for the various meiotic events were obtained. Spermatocytes four days in vitro carried out the first meiotic division, but there were many abnormalities and the second meiotic division did not occur. At first meiotic metaphase, whole bivalent oscillations were accompanied by a relatively higher frequency oscillatory movement of the two homologous kinetochore regions. Oscillations of the kinetochore region were independently variable in magnitude and frequency. A system is proposed by which the metaphase bivalent movements are explained in terms of two pulling forces acting with variable intensity and frequency in opposite directions at the two homologous kinetochores. Meiosis in heavily compressed spermatocytes was blocked at the first meiotic metaphase, apparently because of the absence of a bipolar meiotic apparatus. In compressed spermatocytes, the centrosome divided but the two resulting centrosomes failed to reach their definitive polar positions. After about two hours of separation, the two centrosomes reversed their movement and fused to form a single centrosome from which a unipolar half-spindle radiated.This investigation was supported by grant GB-15 from the National Science Foundation and by Public Health Service Research Grant GB 12431-02 from the Division of General Medical Sciences.Deceased June 17, 1964.