Synapsis and crossing over within a paracentric inversion in the grasshopper,Camnula pellucida

A male grasshopper, Camnula pellucida (Scudder), was found to be heterozygous for a paracentric inversion occupying approximately 10% of the length of one of the two longest chromosomes. Analysis of 297 cells in pachytene revealed inversion loops, suspected inversion loops, asynapsis, and straight pairing in 1.0, 2.7, 8.4, and 87.9% of the analyzable cells, respectively. The frequency of straight pairing (87.9%) indicated a high degree of non-homologous pairing. Analysis of 603 cells in anaphase I and II, and in telophase I and II for the presence of acentric fragments and dicentric chromatid bridges indicated that crossing over within the inversion region occured in about 8% of the cells. The difference between the frequency of the observed plus suspected inversion loops in pachytene and that of the dicentric chromatid bridges and acentric fragments in anaphase I or subsequent stages was not statistically significant. The correspondence between the presence of inversion loops and crossovers within the region of the inversion is thus similar to that observed by Maguire (1966) for a short paracentric inversion in maize. The reasons for this correspondence are considered.Supported by grants GB 1585 and GB 6745 from the National Science Foundation, Washington, D.C.     

The origin of meiotic bridges by chiasma formation in heterozygous inversions in Prosimulium multidentatum (Diptera: Simuliidae)

Prosimulium multidentatum (Twinn) has three metacentric pairs in its chromosome complement. All six arms are individually identifiably in polytene nuclei. XY₁ males are heterozygous for a small basal non-conformity in section 59 of the non-pairing sex differential segments which extends from sextion 58 to section 62 of the IIL arm. XY₂ males carry an additional large heterozygous inversion in the center of this same arm. Meiosis is chiasmate in both kinds of male. In XY₂ individuals 14.2% of the pachytene nuclei show reverse loop pairing and 12.5% of the anaphase I cells form bridge-fragment configurations. A majority of these bridges persist into second division and 7.1% double sized spermatids are formed. No pachytene loops or anaphase bridges were found in XY₁ males. It is concluded therefore that the bridges and fragments of XY₂ males result from chiasma formation within the Y₂ inversion.     

Fine structure of anaphase bridges in meiotic chromosomes of the crane fly Pales

Chromatin bridges of autosomal bivalents in anaphase I were observed in spermatocytes of Pales ferruginea. The bridges are formed without simultaneous production of akinetochoric (akinetic) fragments. A bridge consists of a single fiber up to approximately. 500 Å in diameter. Filamentous substructures of approximately 100 Å diameter can be visualized. It is suggested that these bridges represent a low order coiling of the chromatid, and may be caused by non-separation of the terminal segments of the chromatids (telomeres).     

New observations on lecanoid spermatogenesis in the mealy bug,Planococcus citri

Summary A reexamination of the second spermatogenic division of the mealy bug, Planococcus citri (Risso), a lecanoid coccid, has revealed hitherto unknown spindle activity of the euchromatic set of chromosomes during anaphase II. An initial large half spindle elaborated by the heterochromatic chromosomes in early metaphase, gives way to a less pronounced, but clearly visible bipolar spindle involving both sets of chromosomes at early anaphase. There is no lengthening of the spindle or cell, but the separation of the chromosomes occurs around the periphery of the cell with the aid of interzonal activity. The active participation of the euchromatic chromosome during the separation is furthermore inferred by the formation of bridges resulting from euchromatic-heterochromatic translocations.     

Configurations resulting from iso-chromatid and iso-subchromatid unions after meiotic and mitotic prophase irradiation

Summary By making use of the chromosomes of Trillium erectum as a model, potential and actual configurations arising from presumed iso-chromatid and iso-subchromatid unions after irradiation of meiotic or mitotic prophase have been studied and analyzed. Diagrams and photographs of various recognizable types of chromatid or subchromatid rearrangements are presented. A minimum of two iso-chromatid unions within an arm of a single chromosome in meiotic prophase, if separated by a single chiasma, can give rise to a monocentric chromosome with a triplicated segment, the middle portion of which is an inversion. A minimum of two iso-subchromatid breaks within an arm at either meiotic or mitotic prophase also can result in the production of a monocentric chromosome containing a triplicated segment. The stage of appearance of dicentrics or bridges arising from chromatid or subchromatid unions in meiotic prophase is influenced by chiasma number and pattern and by the number of strands per chromosome or chromatid. Some of the rearrangements described may have genetic and evolutionary implication of considerable potential importance which has not been recognized previously.Research carried out at Brookhaven National Laboratory under the auspices of the U.S. Atomic Energy Commission.     

Chromosomal interchange inEleutherine plicata Herb

Chromosome behaviour at metaphase I and anaphase I of meiosis inEleutherine plicata Herb. (2n=14) is studied. Cells with chromosome associations comprising an association of four long chromosomes, in addition to five bivalents were observed more frequently than those with seven bivalents. it is concluded that the ring of four is due to a segmental interchange between the two long non-homologous chromosome pairs. The ring of four at anaphase I showed delayed disjunction, bridge formation and irregular separation of chromosomes in a number of cells while the behaviour of the other bivalents was normal.     

Chromosome differentiation in diploid species of Lotus (Leguminosae)

Summary Meiotic chromosome behavior was studied in seven diploid species of Lotus (L. alpinus Schleich., L. japonicus (Regel) Larsen, L. filicaulis Dur., L. schoelleri Schweinf., L. krylovii Schischk. and Serg., L. tenuis Waldst. et Kit., L. corniculatus var. minor Baker) and in 51 interspecific hybrids from 16 different crosses. Meiosis in the diploid species was quite regular. In a high proportion of the PMC's of the hybrids there was close chromosome homology with a normal association of 6 II's. However, meiotic irregularities including bridges, lagging chromosomes, univalents, and quadrivalents, occurred in a small percentage of the cells. The late separation of bivalents, the presence of quadrivalents, and inversion bridges with fragments, would indicate for some hybrids that certain chromosomes were structurally differentiated. The large number of rod bivalents observed at diakinesis was also highly suggestive that genetic nonhomology in one chromosome arm could contribute to the frequency of this type of bivalent. Therefore, the maximum number of 6 II's which occurred in a high percentage of cells may be misleading in that cryptic structural differences between chromosome arms, or segments, are not revealed. Pollen fertility in the species and hybrids was not correlated with meiotic irregularities suggesting that pollen fertility is genotypically controlled.     

小麦等离子束处理的生物学效应研究

本文以早熟品种冀麦31号和晚熟品系88—4284萌动种子为材料,报道了等离子束处理对小麦种子萌发的影响和细胞学效应。观察表明,等离子束长时间处理抑制种子发芽。在根尖细胞有丝分裂中期观察到染色体断裂和断片,冀麦31号染色体断裂的频率为0.49—1.34％,88—4284为0.21—2.14％。随着处理时间延长,染色体断裂频率逐渐提高。有丝分裂后期和末期出现大量的落后染色单体和染色体桥。在花粉母细胞减数分裂中期Ⅰ,经过等离子束处理的材料出现环状单价体。后期Ⅰ观察到染色体倒位造成的染色体桥和断片。在四分体期还有微核出现。     

Cytogenetic studies on <Emphasis Type="Italic">Metasequoia glyptostroboides</Emphasis>, a living fossil species

The chromosome morphology and meiotic pairing behavior in the pollen mother cells (PMCs) of Metasequoia glyptostroboides were investigated. The results showed that: (1) The chromosome number of the PMCs was 2n=22. (2) The PMCs developed in the successive manner, and the nucleoids in the dynamic development were similar to those of the other gymnosperms. (3) At prophase, most of the chromosomes were unable to be identified distinctively because the chromosomes were long and tangled together. The chromosome segments were paired non-synchronously. At pachytene, the interstitial or terminal regions of some bivalents did not form synapsis and the paired chromosomes showed difference in sizes, indicating that there were structure differences between the homologous chromosomes. (4) At diakinesis, the ring bivalents showed complicated configurations due to the differences in location and number of chiasmata. In addition, there were cross-linked bivalents. (5) At metaphase I, the chromosome configuration of each cell was 8.2II ⁰ + 1.1II + 1.3II ⁺ + 0.8I. Most of the chromosomes were ring bivalents, but some were cross-linked bivalents, rod bivalents, or univalents. (6) 15\% PMCs at anaphase I and 22\% PMCs at anaphase II presented chromosome bridges, chromosome fragments, micronuclei, and lagging chromosomes. Twenty seven percent microspores finally moved into one to three micronuclei. Twenty five percent pollens were abortive. The results indicated that the observed individual of M. glyptostroboideswas probably a parpcentric inversion heterozygote, and there were structural and behavioral differences between the homologous chromosomes. The chromosomal aberration of M. glyptostroboidesmay play an important role in the evolution of this relict species, which is known as a living fossil. Further evidence is needed to test whether the differences between homologous chromosomes were due to hybridization.     

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     

Mitosis with undivided chromosomes

Summary Cases of cell division with single chromatids are discussed in connection with a study on mitosis with undivided chromosomes made on living material of the endosperm of Haemanthus katharinae. Such divisions are known from certain abnormal mitoses in the microspores of a few plant species, and also from the second meiotic division, in which it is possible in numerous materials to study the behaviour of daughter univalents, and, in a few cases, also daughter chromosomes derived from chromosomes that were paired during the first division.The various cases of mitosis with single chromatids show a great variation with respect to the degree of scattering of the chromosomes over the spindle at metaphase. In a few cases there is practically no tendency to form a metaphase plate. In other cases the tendency to form such a plate is more or less pronounced, but also in these cases it is difficult for the chromosomes to form this arrangement. Some of them remain scattered over the spindle. After the metaphase a kind of anaphase usually follows in which the single chromatids, without division, move to the poles, often with other chromosomes lagging in intermediate positions.An approach of chromosomes to the poles may be caused by two different mechanisms in mitoses of this kind and only in a few cases is the information sufficient to show that active centromere movements occur during these anaphases.In many aspects of their behaviour on the spindle, single chromatids are similar to ordinary univalents of the first meiotic division. For this reason the movement mechanics of the chromosomes of the first meiotic division is briefly reviewed.The interpretation is expressed that the structure of the centromere region of a single chromatid shows some similarity to that of a univalent of the first meiotic division and that this may be the reason for their similar behaviour. The chromatid centromere would have a structural multiplicity with respect to its kinetic elements, corresponding to its subdivision in half-chromatids and also to the presence of two or three consecutive chromomeres in its longitudinal direction. As these kinetic elements are arranged close to one another on one side of the narrow cylinder of the centromere constriction, it is difficult for them to orient, towards both poles simultaneously. A single chromatid having a centromere of this kind will show orientation instability and change its orientation between the two unipolar orientations and various more or less bipolar orientations. The movements following these different orientations would cause the scattering of these single chromatids over the spindle. The orientation of ordinary mitotic metaphase chromosomes, consisting of two such chromatids, could often be the consequence of a process of co-orientation similar to that in meiotic bivalents.The anaphase movement of undivided chromosomes, which by active centromere movements are shifted in the polar directions without a separation of daughter components, is discussed with reference to a similar behaviour observed by Dietz in multivalents in Ostracods. These multivalents are stabilized in the equator during metaphase, in spite of the fact that they have two or three centromeres directed towards one pole and a single one towards the other. During anaphase their chromosomes do not separate but the whole configurations are shifted towards that pole towards which the majority of the centromeres are directed (this is followed by another type of movement which does not concern us in this connection). Undivided chromosomes that are oriented with more of their kinetic material towards one of the poles and less towards the other should by the same mechanisms as moved the multivalents be shifted in the equatorial direction during metaphase and in the polar direction during anaphase. The mechanism of these events is obscure. A change in the interpretation given by Dietz is suggested.This paper is dedicated to Professor Franz Schrader on the occasion of his seventieth birthday.     

Evidence for separate genetic control of crossing over and chiasma maintenance in maize

The meiotic cytological behavior of chromosomes in maize microsporocytes homozygous for the recessive mutant desynaptic was studied at various stages. It was found that following apparently normal pachytene synapsis there appears to be sporadic precocious desynapsis. By diakinesis bivalents heterozygous for a distal knob have often separated to pairs of univalents, each with a knob-carrying and a knobless chromatid. From the frequency of such events it is inferred that the crossover process is probably not affected by the mutant and that the genetic defect affects instead a distinct function concerned with chiasma maintenance following crossing over. Since precocious separation of dyads to monads at prophase II was also found in the desynaptic material, it is suggested that normal chiasma maintenance until anaphase I and normal dyad integrity maintenance between anaphase I and anaphase II may depend upon the same mechanism; it is also suggested that this may involve a special tendency for cohesiveness of sister chromatids during meiosis, beyond that which is ordinarily found at mitosis.     

矮牡丹小孢子母细胞减数分裂异常现象的观察

矮牡丹(Paeonia suffruticosa subsp. spontanea)(永济居群)存在多种结构杂合现象,减数分裂存在一些异常:如单价体、异形二价体、互锁四价体、六价体、后期I倒位桥、落后单价体、不均等分离、后期Ⅱ桥和微核等。统计了这些异常现象出现的频率,并对其形成的机制和对正常小孢子形成的影响进行了讨论。从细胞学水平上探讨了矮牡丹可能的濒危机制。同时结合前人的研究,对芍药属内3个组的结构杂合程度进行了比较。     

Holocentric chromosome behaviour inCuscuta (Cuscutaceae)

The present study demonstrates thatCuscuta babylonica Choisy has holocentric chromosomes. Evidence for this phenomenon comes from three different observations. (1) Mitosis: During metaphase and anaphase the sister-chromatids are situated parallel to the equatorial plane with no sign of localized kinetochore activity. (2) Inverted meiosis in microsporocytes. (3) X-rayed microsporocytes, in which the numerous chromosome fragments do not show any lagging or formation of micronuclei. We assume that only one out of the three subgenera inCuscuta, namely subg.Cuscuta, has holocentric chromosomes, while the two other subgenera have monocentric chromosomes.     

Incomplete sister chromatid separation of long chromosome arms

Chromosome segregation ensures the equal partitioning of chromosomes at mitosis. However, long chromosome arms may pose a problem for complete sister chromatid separation. In this paper we report on the analysis of cell division in primary cells from field vole Microtus agrestis, a species with 52 chromosomes including two giant sex chromosomes. Dual chromosome painting with probes specific for the X and the Y chromosomes showed that these long chromosomes are prone to mis-segregate, producing DNA bridges between daughter nuclei and micronuclei. Analysis of mitotic cells with incomplete chromatid separation showed that reassembly of the nuclear membrane, deposition of INner CENtromere Protein (INCENP)/Aurora B to the spindle midzone and furrow formation occur while the two groups of daughter chromosomes are still connected by sex chromosome arms. Late cytokinetic processes are not efficiently inhibited by the incomplete segregation as in a significant number of cell divisions cytoplasmic abscission proceeds while Aurora B is at the midbody. Live-cell imaging during late mitotic stages also revealed abnormal cell division with persistent sister chromatid connections. We conclude that late mitotic regulatory events do not monitor incomplete sister chromatid separation of the large X and Y chromosomes of Microtus agrestis, leading to defective segregation of these chromosomes. These findings suggest a limit in chromosome arm length for efficient chromosome transmission through mitosis.Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

End association and segregation of the achiasmatic X and Y chromosomes of the sand rat,Psammomys obesus

In Psammomys obesus there is no pairing between the X and Y chromosomes and no chiasma formation (Solari and Ashley, 1977). It is demonstrated that ends of the axial elements of the X and Y chromosomes come together during pachytene, and regularly form at least one end-to-end junction. This achiasmatic physical connection between the ends of the X and Y persists until anaphase I, thus assuring the normal distribution of the sex chromosomes observed by light microscopy. In addition, there are no differentiations of the axes of the X and Y similar to those observed in other mammalian species thus far examined, a fact that could influence chromatid cohesiveness and disjunction.     

Sequence of centromere separation: generation of unstable multicentric chromosmes in a rat cell line

A transformed cell line, B₁, of cerebral endothelial origin from the Wistar-Kyoto male rat has chromatid and chromosome type bridges in virtually every cell. It exhibits various dicentric and polycentric chromosomes. Most dicentrics are symmetric isochromosomes. Certain isodicentrics are present in a fair segment of the cell population; however, almost all cells have some newly arising isodicentrics. The live cells show a lengthened prometaphase. Anaphase is also retarded possibly due to the occurrence of bridges. At anaphase some multicentrics split at only one centromere. When pulled to the two poles the unsplit centromeres and the distal chromosome segment form a side arm bridge. Another mechanism appears to be a total lack of separation of daughter centromeres at meta-anaphase (meiotic-like behavior of mitotic chromosomes). This is realized by the pulling of each of the two unsplit centromeres to opposite poles and results in bridges with both sister chromatids running parallel to each other. A break at corresponding weak points in the two sister chromatids followed by rejoining can form a dicentric isochromosome. A third mechanism, the breakage-fusion-bridge cycle, is also operative but would not produce isodicentrics. In the case of the first two mechanisms some or all centromeres apparently split between telophase and onset of the following DNA synthesis rather than at the usual time at late metaphase. These observations may suggest some previously unknown behavior of multicentric chromosomes during mitosis.     

Chromosomal Behavior during Meiosis in the Progeny of Triticum timopheevii × Hexaploid Wild Oat

The meiotic behavior of pollen mother cells (PMCs) of the F₂ and F₃ progeny from Triticum timopheevii × hexaploid wild oat was investigated by cytological analysis and sequential C-banding-genomic in situ hybridization (GISH) in the present study. A cytological analysis showed that the chromosome numbers of the F₂ and F₃ progeny ranged from 28 to 41. A large number of univalents, lagging chromosomes, chromosome bridges and micronuclei were found at the metaphase I, anaphase I, anaphase II and tetrad stages in the F₂ and F₃ progeny. The averages of univalents were 3.50 and 2.73 per cell, and those of lagging chromosomes were 3.37 and 1.87 in the F₂ and F₃ progeny, respectively. The PMC meiotic indices of the F₂ and F₃ progeny were 12.22 and 20.34, respectively, indicating considerable genetic instability. A sequential C-banding-GISH analysis revealed that some chromosomes and fragments from the hexaploid wild oat were detected at metaphase I and anaphase I in the progeny, showing that the progeny were of true intergeneric hybrid origin. The alien chromosomes 6A, 7A, 3C and 2D were lost during transmission from F₂ to F₃. In addition, partial T. timopheevii chromosomes appeared in the form of univalents or lagging chromosomes, which might result from large genome differences between the parents, and the wild oat chromosome introgression interfered with the wheat homologues’ normally pairing.     

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.     

Inversion polymorphism in the midge Glyptotendipes barbipes (Staeger)

Summary The salivary gland nuclei of larvalGlyptotendipes barbipes from Stratford (Ontario) comprise 3 pairs of long metacentric chromosomes (I–III) and one pair of short acrocentrics (IV). Homologues are closely paired when homozygous, centromeres are expressed as heterochromatic drums. Each chromosome carries a nucleolus, and Chromosome IV in addition carries a Ring of Balbiani. The gross polytene idiogram is identical with that implicit inBauer's cytological analysis of the same species from Germany.In direct comparison the banding pattern of the German and Canadian larvae proved identical for at least one sequence in each chromosome arm.Three simple inversions are known in the species; one (III L-1) is endemic to Germany, a second (II L-3) has been found only once in Canada and a third (I S-1) is common in Canada but has not been reported from Germany. The remaining 2 inversions (II L-1,2 and III S-1,2) are complex and are here analysed as included types. Rearrangement II L-1,2 occurs both in Canada and in Germany and achieves heterozygous frequencies near 50% in both countries. It is the first inversion reported to be holarctic in distribution in insects not associated with man. The second complex inversion III S-1,2 has been found only in Canada where it is common. In each of the complex inversions at least one pair of breaks is near coincident.Meiosis is normal and chiasmatic in structurally homozygous males. It lacks conspicuous localization of chiasmata. No meiotic abnormalities were observed in inversion heterozygotes. It is postulated that complex heterozygous inversions interfere with pairing to such an extent that the residual ill-effects are out balanced by heterosis of co-adapted systems.A selective advantage of heterozygosis is directly indicated for II L by a significant (p=0.02) excess of II L/II L-1,2 larvae over the Hardy-Weinberg expectation.