THE ULTRASTRUCTURE OF KINETOCHORES OF UNPAIRED CHROMOSOMES IN A WHEAT HYBRID

1973 ◽  
Vol 15 (4) ◽  
pp. 801-806 ◽  
Author(s):  
E. B. Wagenaar ◽  
D. F. Bray

The kinetochore region of unpaired chromosomes (univalents) consists of two kinetochores, each belonging to a sister chromatid, that are located adjacent to one another on the surface of the univalent chromosome. This condition results in a movement by the univalent towards one of the polar regions at the onset of metaphase I. Once arrived in this region, one of the sister kinetochores obtains attachments of microtubules from the opposite pole. This results in a gradual return of the univalent to the equatorial plate, where it reaches an equilibrium. The sister kinetochores remain adjacent during the movement, but once arrived at the metaphase plate they develop a typical mitotic appearance, in which the sister kinetochores have opposite positions on the chromosomes.

1994 ◽  
Vol 107 (4) ◽  
pp. 785-798 ◽  
Author(s):  
B.C. Williams ◽  
M.L. Goldberg

We have examined several issues concerning how the Drosophila l(1)zw10 gene product functions to ensure proper chromosome segregation. (a) We have found that in zw10 mutant embryos and larval neuroblasts, absence of the zw10 protein has no obvious effect on either the congression of chromosomes to the metaphase plate or the morphology of the metaphase spindle, although many aberrations are observed subsequently in anaphase. This suggests that activity of the zw10 protein becomes essential at anaphase onset, a time at which the zw10 protein is redistributed to the kinetochore region of the chromosomes. (b) The zw10 protein appears to bind to kinetochores in mitotically arrested cells, eventually accumulating to high levels within the chromosome mass. Our results imply that zw10 may act as part of a novel feedback pathway that normally renders sister chromatid separation dependent upon spindle integrity. (c) The localization of zw10 protein is altered by two mitotic mutations, rough deal and abnormal anaphase resolution, that specifically disrupt anaphase. These findings indicate that the zw10 protein functions as part of a multicomponent mechanism ensuring proper chromosome segregation at the beginning of anaphase.


1999 ◽  
Vol 112 (17) ◽  
pp. 2957-2969 ◽  
Author(s):  
J.A. Suja ◽  
C. Antonio ◽  
A. Debec ◽  
J.S. Rufas

Sister-chromatid arm cohesion is lost during the metaphase I/anaphase I transition to allow homologue separation. To obtain needed information on this process we have analysed in grasshopper bivalents the sequential release of arm cohesion in relation to the behaviour of chromatid axes. Results show that sister axes are associated during early metaphase I but separate during late metaphase I leading to a concomitant change of chromosome structure that implies the loss of sister-kinetochore cohesion. Afterwards, homologues initiate their separation asynchronously depending on their size, and number and position of chiasmata. In all bivalents thin chromatin strands at the telomeres appeared as the last point of contact between sister chromatids. Additionally, we have analysed the participation of phosphoproteins recognised by the MPM-2 monoclonal antibody against mitotic phosphoproteins in arm cohesion in bivalents and two different kinds of univalents. Results show the absence of MPM-2 phosphoproteins at the interchromatid domain in mitotic chromosomes and meiotic univalents, but their presence in metaphase I bivalents. These phosphoproteins are lost at the onset of anaphase I. Taken together, these data have prompted us to propose a ‘working’ model for the release of arm cohesion during meiosis I. The model suggests that MPM-2 phosphoproteins may act as cohesive proteins associating sister axes. Their modification, once all bivalents are correctly aligned at the metaphase plate, would trigger a change of chromosome structure and the sequential release of sister-kinetochore, arm, and telomere cohesions.


1961 ◽  
Vol 39 (1) ◽  
pp. 81-108 ◽  
Author(s):  
E. B. Wagenaar

In two hybrids between Triticum timopheevi Zhuk. and T. durum Desf., which have irregular meioses, metaphase I was subdivided into four developmental stages, early, medium, late, and very late. This subdivision was based on the presence in the anthers of other stages that occurred together with metaphase I. It was then discovered that in metaphase I cell populations there was a progressive increase of univalents from the early and medium stages to the very late stage. This phenomenon can be explained on the assumption that metaphase I is of shorter duration in the less irregular cells which pass into anaphase I earlier than the more irregular cells. As a consequence of this developmental phenomenon at metaphase I, the anaphase I and telophase I cells in the late anthers contained fewer lagging chromosomes than the anaphase I and telophase I cells in the very late anthers. Despite these numerical differences, the degrees of lagging were remarkably similar in both stages; approximately 70% of these univalents lagged at late and very late stages in both hybrids.During metaphase I many univalents of the irregular cells moved towards the equatorial plate, became oriented, and lagged at anaphase I and telophase I. It was found that the univalents of the least irregular cells accumulated somewhat faster at the plates than those of the more irregular cells.Considering the relationships between all of the available data, the hypothesis is advanced that when a certain number of univalents have accumulated at the equatorial plate a state of equilibrium is established and anaphase I is initiated. On the basis of this hypothesis an explanation of the trends observed at metaphase I is given.


1988 ◽  
Vol 90 (3) ◽  
pp. 493-499
Author(s):  
GEOFFREY K. RICKARDS ◽  
WENDY A. BAKER

The position of an interchange quadrivalent at metaphase I of Allium triquetrum was modelled so as to simulate the original placement of the quadrivalent in the spindle and the preparation of linear spreads through squashing. In this way an expected distribution for the quadrivalent in linear spreads was generated. The procedure used polar views of metaphase I to which the quadrivalent was assigned pairs of positions normally occupied by bivalents. The positions of the bivalents and assigned quadrivalent were then into a linear array and analysed as ‘real’ data. Comparisons with observed distributions showed that a general bias in favour of marginal placement of the quadrivalent in the linear array is expected; and also showed that the quadrivalent is positioned non-randomly in the metaphase plate prior to slide preparation.


1990 ◽  
Vol 110 (1) ◽  
pp. 81-95 ◽  
Author(s):  
C L Rieder ◽  
S P Alexander

During mitosis in cultured newt pneumocytes, one or more chromosomes may become positioned well removed (greater than 50 microns) from the polar regions during early prometaphase. As a result, these chromosomes are delayed for up to 5 h in forming an attachment to the spindle. The spatial separation of these chromosomes from the polar microtubule-nucleating centers provides a unique opportunity to study the initial stages of kinetochore fiber formation in living cells. Time-lapse Nomarski-differential interference contrast videomicroscopic observations reveal that late-attaching chromosomes always move, upon attachment, into a single polar region (usually the one closest to the chromosome). During this attachment, the kinetochore region of the chromosome undergoes a variable number of transient poleward tugs that are followed, shortly thereafter, by rapid movement of the chromosome towards the pole. Anti-tubulin immunofluorescence and serial section EM reveal that the kinetochores and kinetochore regions of nonattached chromosomes lack associated microtubules. By contrast, these methods reveal that the attachment and subsequent poleward movement of a chromosome correlates with the association of a single long microtubule with one of the kinetochores of the chromosome. This microtubule traverses the entire distance between the spindle pole and the kinetochore and often extends well past the kinetochore. From these results, we conclude that the initial attachment of a chromosome to the newt pneumocyte spindle results from an interaction between a single polar-nucleated microtubule and one of the kinetochores on the chromosome. Once this association is established, the kinetochore is rapidly transported poleward along the surface of the microtubule by a mechanism that is not dependent on microtubule depolymerization. Our results further demonstrate that the motors for prometaphase chromosome movement must be either on the surface of the kinetochore (i.e., within the corona but not the plate), distributed along the surface of the kinetochore microtubules, or both.


2003 ◽  
Vol 160 (5) ◽  
pp. 657-670 ◽  
Author(s):  
Maureen Eijpe ◽  
Hildo Offenberg ◽  
Rolf Jessberger ◽  
Ekaterina Revenkova ◽  
Christa Heyting

In meiotic prophase, the sister chromatids of each chromosome develop a common axial element (AE) that is integrated into the synaptonemal complex (SC). We analyzed the incorporation of sister chromatid cohesion proteins (cohesins) and other AE components into AEs. Meiotic cohesin REC8 appeared shortly before premeiotic S phase in the nucleus and formed AE-like structures (REC8-AEs) from premeiotic S phase on. Subsequently, meiotic cohesin SMC1β, cohesin SMC3, and AE proteins SCP2 and SCP3 formed dots along REC8-AEs, which extended and fused until they lined REC8-AEs along their length. In metaphase I, SMC1β, SMC3, SCP2, and SCP3 disappeared from the chromosome arms and accumulated around the centromeres, where they stayed until anaphase II. In striking contrast, REC8 persisted along the chromosome arms until anaphase I and near the centromeres until anaphase II. We propose that REC8 provides a basis for AE formation and that the first steps in AE assembly do not require SMC1β, SMC3, SCP2, and SCP3. Furthermore, SMC1β, SMC3, SCP2, and SCP3 cannot provide arm cohesion during metaphase I. We propose that REC8 then provides cohesion. RAD51 and/or DMC1 coimmunoprecipitates with REC8, suggesting that REC8 may also provide a basis for assembly of recombination complexes.


Development ◽  
1996 ◽  
Vol 122 (12) ◽  
pp. 3775-3784 ◽  
Author(s):  
B. de Saint Phalle ◽  
W. Sullivan

Sex in Sciara coprophila is determined by maternally supplied factors that control the number of paternal X chromosomes eliminated during the syncytial embryonic divisions. Confocal microscopy and FISH demonstrate that the centromeres of the X chromosomes separate at anaphase and remain functional during the cycle in which the X chromosomes are eliminated. However, a region of the sister chromatids fails to separate and the X chromosomes remain at the metaphase plate. This indicates that failure of sister chromatid separation is the mechanism of chromosome elimination. Elimination of the X chromosomes requires the presence of a previously discovered Controlling Element that acts in cis during male meiosis. Using an X-autosome translocation, we demonstrate that the Controlling Element acts at-a-distance to prevent sister chromatid separation in the arm of an autosome. This indicates that the region in which sister chromatid separation fails is chromosome-independent. Although chromosome elimination occurs in all somatic nuclei and is independent of location of the nuclei within the embryo, the decision to eliminate is made at the level of the individual nucleus. Programmed X chromosome elimination occurs at different cycles in male and female embryos. These observations support a model in which elements on the X chromosome are titrating maternally supplied factors controlling the separation of sister X chromatids.


2011 ◽  
Vol 22 (18) ◽  
pp. 3465-3477 ◽  
Author(s):  
Jibak Lee ◽  
Sugako Ogushi ◽  
Mitinori Saitou ◽  
Tatsuya Hirano

In many eukaryotes, condensins I and II associate with chromosomes in an ordered fashion during mitosis and play nonoverlapping functions in their assembly and segregation. Here we report for the first time the spatiotemporal dynamics and functions of the two condensin complexes during meiotic divisions in mouse oocytes. At the germinal vesicle stage (prophase I), condensin I is present in the cytoplasm, whereas condensin II is localized within the nucleus. After germinal vesicle breakdown, condensin II starts to associate with chromosomes and becomes concentrated onto chromatid axes of bivalent chromosomes by metaphase I. REC8 “glues” chromosome arms along their lengths. In striking contrast to condensin II, condensin I localizes primarily around centromeric regions at metaphase I and starts to associate stably with chromosome arms only after anaphase I. Antibody injection experiments show that condensin functions are required for many aspects of meiotic chromosome dynamics, including chromosome individualization, resolution, and segregation. We propose that the two condensin complexes play distinctive roles in constructing bivalent chromosomes: condensin II might play a primary role in resolving sister chromatid axes, whereas condensin I might contribute to monopolar attachment of sister kinetochores, possibly by assembling a unique centromeric structure underneath.


2001 ◽  
Vol 276 (50) ◽  
pp. 47575-47582 ◽  
Author(s):  
Heather C. Gregson ◽  
John A. Schmiesing ◽  
Jong-Soo Kim ◽  
Toshiki Kobayashi ◽  
Sharleen Zhou ◽  
...  

The cohesin multiprotein complex containing SMC1, SMC3, Scc3 (SA), and Scc1 (Rad21) is required for sister chromatid cohesion in eukaryotes. Although metazoan cohesin associates with chromosomes and was shown to function in the establishment of sister chromatid cohesion during interphase, the majority of cohesin was found to be off chromosomes and reside in the cytoplasm in metaphase. Despite its dissociation from chromosomes, however, microinjection of an antibody against human SMC1 led to disorganization of the metaphase plate and cell cycle arrest, indicating that human cohesin still plays an important role in metaphase. To address the mitotic function of human cohesin, the subcellular localization of cohesin components was reexamined in human cells. Interestingly, we found that cohesin localizes to the spindle poles during mitosis and interacts with NuMA, a spindle pole-associated factor required for mitotic spindle organization. The interaction with NuMA persists during interphase. Similar to NuMA, a significant amount of cohesin was found to associate with the nuclear matrix. Furthermore, in the absence of cohesin, mitotic spindle asters failed to formin vitro. Our results raise the intriguing possibility that in addition to its well demonstrated function in sister chromatid cohesion, cohesin may be involved in spindle assembly during mitosis.


1992 ◽  
Vol 116 (4) ◽  
pp. 967-976 ◽  
Author(s):  
G Maldonado-Codina ◽  
D M Glover

Maternally contributed cyclin A and B proteins are initially distributed uniformly throughout the syncytial Drosophila embryo. As dividing nuclei migrate to the cortex of the embryo, the A and B cyclins become concentrated in surface layers extending to depths of approximately 30-40 microns and 5-10 microns, respectively. The initiation of nuclear envelope breakdown, spindle formation, and the initial congression of the centromeric regions of the chromosomes onto the metaphase plate all take place within the surface layer occupied by cyclin B on the apical side of the blastoderm nuclei. Cyclin B is seen mainly, but not exclusively, in the vicinity of microtubules throughout the mitotic cycle. It is most conspicuous around the centrosomes. Cyclin A is present at its highest concentrations throughout the cytoplasm during the interphase periods of the blastoderm cycles, although weak punctate staining can also be detected in the nucleus. It associates with the condensing chromosomes during prophase, segregates into daughter nuclei in association with chromosomes during anaphase, to redistribute into the cytoplasm after telophase. In contrast to the cycles following cellularization, neither cyclin is completely degraded upon the metaphase-anaphase transition.


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