Influence of a rare unstable B-chromosome on chiasma frequency and nonhaploid sperm production in Dichroplus pratensis (Melanoplinae,Acrididae)

Three B-containing individuals of Dichroplus pratensis were found among 300 males collected in several Argentine localities. The B-chromosome is a partially euchromatic short telocentric and it is mitotically unstable. The B-chromosome number of testis cells ranged from 0 to 4 in two males and from 0 to 3 in the remaining one; this variation was inter- as well as intrafollicular. Meiotic behaviour of the Bs was very regular, association of Bs was usually chiasmate and no lagging univalents were observed during first division. The presence of Bs increased the production rate of abnormal spermatids and this effect was more pronounced in follicles with odd numbers of Bs. Chiasma frequency and between-cell variance were also increased in B-containing cells. A possible interaction between the Bs and the polymorphic centric fusions present in the species is discussed in relation to chiasma frequency determination.     

Morphology and Meiotic/Mitotic Behavior of B Cromosomes in a Japanese Harvestman, Metagagrella tenuipes (Arachnida: Opiliones): No Evidence for B Accumulation Mechanisms

Earlier, it has been demonstrated that wild populations of a Japanese harvestman Metagagrella tenuipes (Arachnida: Opiliones) are polymorphic for B chromosomes. In this paper, we present results of a study of the morphology and mitotic and meiotic behavior of the Bs. The B chromosomes varied considerably in size and proportion of eu- and heterochromatin. The single nucleolus organizing region, found in males, was located on a chromosome of the A complement. Some intercell variation in number of Bs may be explained by accidental chromosome losses during chromosome preparation. We also found no intertissue variation in number of Bs. There were also no differences in mean number of B chromosomes per individual among males and females, adult and subadult harvestmen. Segregation of Bs in mitotic and meiotic divisions was nonrandom; B chromosomes tended to segregate equally between daughter cells. The results obtained provide no support for the hypothesis of existence of B accumulation mechanism in this species.     

Mechanism of chromosome elimination in the hybridogenetic spermatogenesis of allotriploid males between Japanese and European water frogs

Of 21 allotriploid males that possessed two genomes of Rana nigromaculata and one genome of Rana lessonae 10 produced a large number of spermatozoa in their testes. When 4 of these males were backcrossed with a female of R. nigromaculata, all of the resulting froglets were diploid in chromosome number and were completely R. nigromaculata type in appearance. These allotriploid males proved to have produced spermatozoa with one R. nigromaculata genome hybridogentically. Therefore, their germ line cells were investigated for the mechanism of elimination of their R. lessonae chromosomes. In histologicla sections of testes, the great majority of spermatogonia (approximately 10⁴ cells) between mitotic prometaphase and anaphase appeared normal in chromosome behavior, whereas 17 spermatogonia showed several chromosomes whose behavior deviated from the normal course during the same period. These deviant chromosomes concentrated together near the equatorial plate and remained stationary at anaphase. In metaphase chromosome preparations made from spermatogonia, 67 and 185 of the 477 chromosome spreads were diploid and triploid, respectively. The rest were aneuploid. Notably, 8 triploid spreads consisted of 26 or more normal chromosomes and 13 or fewer degenerate chromosomes. From these results it is concluded that a set of R. lessonae chromosomes is eliminated from some, but not all spermatogonia by becoming degenerate during the mitotic period.by H.C. Macgregor     

Mitotic instability leading to an accumulation of B-chromosomes in grasshoppers

The study of mitotically unstable B-chromosomes (supernumeraries) of two grasshopper species confirmed a suggestion made earlier (Nur, 1963) that the instability should always be associated with a tendency of the B's to increase in frequency. Among 780 Camnula pellucida (Scudder) males from California, 105 had B's. In the testes of these males the number of B's varied from follicle to follicle and ranged between 0 and 4. Because of this variation, the number with which each male started to develop could not be determined. However, the relatively low frequency of males with B's and the regular meiotic behavior of the latter suggested that most of the 105 males started with a single B. Cytological analysis of the cells of the gastric caeca of 31 males whose testes contained B's confirmed this suggestion by showing that only one male had two B's in these cells; all the rest had one. In the testes of the 74 other males the mean number of B's ranged from 0.89–2.50, but only two males had means higher than 2.00. The observed ratio of one male with two B's to 30 with one, suggested that only the two males with the highest means started to develop with two B's and that the other 72 males all started with one. Since the mean for the 72 males was 1.37 B's per male, it was concluded that during the development of the testes of these males the mean increased by 37%. The males with B's had fewer follicles in their testes and apparently had also a lower frequency of normal sperm. — The analysis of the testes of Locusta migratoria L. males from Japan gave results which agreed with those from C. pellucida.Supported by grants GB 1585 and GB 6745 from the National Science Foundation, Washington, D.C.     

A mitotically unstable supernumerary chromosome with an accumulation mechanism in a grasshopper

Summary A population of the grasshopper Calliptamus palaestinensis Bdhr. from near Jerusalem, Israel was analyzed cytologically. Twenty-two percent of the males had extra heterochromatic chromosome segments or supernumerary chromosomes. The extra segments were present in either a heterozygous or a homozygous condition and in frequencies similar to those expected according to the Hardy-Weinberg law.The supernumeraries were heteropycnotic in the first meiotic division but paired and segregated regularly. In almost all the males with supernumeraries the number of supernumeraries varied between follicles in the testes but was constant within each follicle. Only 12.3% of the males had supernumeraries, but among those with supernumeraries the majority had means higher than 1.5. A calculation based on the frequency of the supernumeraries in the zygotes which are expected in the next generation indicated that of the 31 males with supernumeraries, probably not more than 3 males originated from zygotes with more than a single supernumerary. The variation in the number of supernumeraries and the high means probably resulted from preferential non-disjunction of the supernumeraries in the early mitoses prior to the differentiation of the follicles.Mitotic non-disjunction of supernumeraries as the cause of intra-individual variation is considered. It is suggested that variation due to non-disjunction would be maintained only in association with accumulation mechanisms.Postdoctoral Trainee in Biology, U. S. Public Health Service, 1962–1963.     

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.     

Inverted meiosis and its place in the evolution of sexual reproduction pathways

Inverted meiosis is observed in plants (Cyperaceae and Juncaceae) and insects (Coccoidea, Aphididae) with holocentric chromosomes, the centromeres of which occupy from 70 to 90% of the metaphase chromosome length. In the first meiotic division (meiosis I), chiasmata are formed and rodlike bivalents orient equationally, and in anaphase I, sister chromatids segregate to the poles; the diploid chromosome number is maintained. Non-sister chromatids of homologous chromosomes remain in contact during interkinesis and prophase II and segregate in anaphase II, forming haploid chromosome sets. The segregation of sister chromatids in meiosis I was demonstrated by example of three plant species that were heterozygous for chromosomal rearrangements. In these species, sister chromatids, marked with rearrangement, segregated in anaphase I. Using fluorescent antibodies, it was demonstrated that meiotic recombination enzymes Spo11 and Rad5l, typical of canonical meiosis, functioned at the meiotic prophase I of pollen mother cells of Luzula elegance and Rhynchospora pubera. Moreover, antibodies to synaptonemal complexes proteins ASY1 and ZYP1 were visualized as filamentous structures, pointing to probable formation of synaptonemal complexes. In L. elegance, chiasmata are formed by means of chromatin threads containing satellite DNA. According to the hypothesis of the author of this review, equational division of sister chromatids at meiosis I in the organisms with inverted meiosis can be explained by the absence of specific meiotic proteins (shugoshins). These proteins are able to protect cohesins of holocentric centromeres from hydrolysis by separases at meiosis I, as occurs in the organisms with monocentric chromosomes and canonical meiosis. The basic type of inverted meiosis was described in Coccoidea and Aphididae males. In their females, the variants of parthenogenesis were also observed. Until now, the methods of molecular cytogenetics were not applied for the analysis of inverted meiosis in Coccoidea and Aphididae. Evolutionary, inverted meiosis is thought to have appeared secondarily as an adaptation of the molecular mechanisms of canonical meiosis to chromosome holocentrism.     

Body size and symbiotic status influence gonad development in <Emphasis Type="Italic">Aiptasia pallida</Emphasis> anemones

Pale anemones (Aiptasia pallida) coexist with dinoflagellates (primarily Symbiodinium minutum) in a mutualistic relationship. The purpose of this study was to investigate the role of these symbionts in gonad development of anemone hosts. Symbiotic and aposymbiotic anemones were subjected to light cycles that induced gametogenesis. These anemones were then sampled weekly for nine weeks, and gonad development was analyzed histologically. Anemone size was measured as mean body column diameter, and oocytes or sperm follicles were counted for each anemone. Generalized linear models were used to evaluate the influence of body size and symbiotic status on whether gonads were present and on the number of oocytes or sperm follicles produced. Body size predicted whether gonads were present, with larger anemones being more likely than smaller anemones to develop gonads. Both body size and symbiotic status predicted gonad size, such that larger and symbiotic anemones produced more oocytes and sperm follicles than smaller and aposymbiotic anemones. Overall, only 22 % of aposymbiotic females produced oocytes, whereas 63 % of symbiotic females produced oocytes. Similarly, 6 % of aposymbiotic males produced sperm follicles, whereas 60 % of symbiotic males produced sperm follicles. Thus, while gonads were present in 62 % of symbiotic anemones, they were present in only 11 % of aposymbiotic anemones. These results indicate that dinoflagellate symbionts influence gonad development and thus sexual maturation in both female and male Aiptasia pallida anemones. This finding substantiates and expands our current understanding of the importance of symbionts in the development and physiology of cnidarian hosts.     

The B-chromosome system of Hypochoeris maculata

Populations of Hypochoeris maculata (Compositae) over a wide area of Europe contain a small proportion of individuals with mitoticallystable B-chromosomes. The telocentric Bs are about one sixth the length of the smallest A-chromosome and are heterochromatic. Offspring grown from seeds of natural populations have one or at most two B-chromosomes although plants with up to five Bs have been produced in experiment. The Bs rarely pair during meiotic prophase and the univalents usually undergo centric division at anaphase I. Despite lagging, the Bs are rarely lost during male meiosis. The inheritance of the B-chromosomes differs between the two sides with accumulation in the egg and random inheritance or loss on the pollen side. Although meiotic drive may play some part in the maintenance of the B-polymorphism there is a strict upper limit on the number of Bs in natural populations.     

Structure,function, and evolutionary significance of the reproductive system in males of Hebrus ruficeps and H. pusillus (heteroptera,gerromorpha, hebridae)

In Holland, bugs of the species Hebrus pusillus and H. ruficeps have one generation per year and overwinter as unmated adults. Males have two testes with two follicles + vasa efferentia each, paired vasa deferentia and seminal vesicles, an ejaculatory duct, and a protrusible phallus comprising an articulatory apparatus, phallotheca, endosoma, and paired claspers. The skeletomusculature of this system is described (it has 12 paired and four unpaired muscles) and its functions in generating and transferring sperm (summarized in Figs. 70–75) are reconstructed from study of living bugs, dissections, whole mounts, and serial sections. Males of both species produce sperm >2 mm long from stem spermatogonia in the germarium of each follicle. Initial definitive spermatogonia divide synchronously three times to form clones of eight, interconnected, primary spermatocytes. These enlarge up to 43-fold in males of H. pusillus and 78-fold in those of H. ruficeps, undergo meiosis, and, after adult emergence, complete their differentiation into bundles of 32 sperm which coil transversely about the periphery of each follicle at its base. These begin to enter the vasa efferentia in mid August, rupture, and release their sperm into the seminal vesicles where they are stored overwinter. Most spermatocyte and spermatid cysts remaining in the testes degenerate in fall, leaving only stem spermatogonia and a few early spermatocysts in the germaria. Males of H. pusillus begin to mate the first warm days of spring but only the most persistent succeed. A male jumps on the back of a female, induces her to lower her ovipositor, and, within 12 min (@ 18–24°C), introduces the endosoma of his phallus up its shaft and fills his seminal duct with sperm. The female draws this into her gynatrial sac at the end of copulation and transfers it into her spermatheca in about 30 min, the sperm reversing themselves within it so that their heads face towards its mouth. The male may stay on her back for up to 2 hours and may copulate again up to three times before leaving to mate with other females. Males of H. pusillus may be sexually active for months after overwintering, because spermatogonia in their germaria reactivate in spring to produce additional sperm. Those of H. ruficeps do not and males mate successfully only until their supply of overwintered sperm is exhausted. The chromosome complement of H. pusillus males is 2N = 22 + XY. The X and Y chromosomes are of unequal length, form a pseudo pair at metaphase I, and segregate to opposite poles at anaphase I—the first instance of pre-reductional segregation of sex chromosomes to be recorded in the Gerromorpha. The chromosome complement of H. ruficeps may be 2N = 24 + XO but the nature of two chromosomes was not resolved. The single X segregates to half the spermatids at anaphase II.     

Study on Genetic Behaviour of B Chromosomes at Meiosis in Agropyron Gaertn

The chromosomal behaviour at meiosis in pollen mother cells of 33 sample materials from 4 species in Agropyron Gaerm. distributed over various regions in China were observed. We. found that B chromosomes were mainly-present in A. mongolicum Keng (2n=2x=14) and were absent in the tetraploid ,A. cristarum (L.) Gaertn. (2n=4x=28) and ,A. michnoi Roshev. (2n=4x=28). Of all A. desertorum (Fisch.) Schult. (2n=4x=28) populations, they were present in 40%. The pairing between even numbers of B chromosomes usually occured at diakinesis where as uneven numbers of B chromosomes at metaphase 1. B chromosomes at anaphase I separated at random. The Fagging Bs tended to reach two poles by chromatids through precocious division of Bs. The pairing and chiasma frequencies between A chromosomes were increased by the larger numbers of Bs. The relationship between B chromosomes and species ploidy, the homeology among Bs, etc were discussed.     

Morphological and molecular data for larval stages of four species of Petasiger Dietz, 1909 (Digenea: Echinostomatidae) with an updated key to the known cercariae from the Palaearctic

Recent molecular phylogenetic studies on fish tapeworms of the genus Caryophyllaeus Gmelin, 1790 (Cestoda: Caryophyllidea), parasites of cyprinid fishes in the Palaearctic Region, have revealed unexpected phenotypic plasticity that seems to be related to definitive hosts. In the present paper, Caryophyllaeus brachycollis Janiszewska, 1953 is redescribed and its two morphotypes are circumscribed on the basis of newly-collected specimens. Morphotype 1 from barbels [Barbus spp. including the type-host Barbus barbus (L.); Barbinae] and chubs (Squalius spp.; Leuciscinae) is characterised by a more robust body with spatulate scolex, which is only slightly wider than a very short neck region, and the anterior position of the testes and vitelline follicles, which begin immediately posterior to the scolex. Specimens of Morphotype 2 from breams (Abramis spp., Ballerus spp. and Blicca spp.; Abraminae), which have been previously misidentifed as Caryophyllaeus laticeps (Pallas, 1781), possess a more slender body with a flabellate scolex, which is much wider than a long neck, and the first testes begin at a considerable distance posterior to the first vitelline follicles. Despite conspicuous differences in the scolex morphology and the anterior extent of the testes and vitelline follicles, both morphotypes are identical in the morphology of the posterior end of the body, in particular that of the cirrus-sac, which is large, thick-walled, elongate-pyriform, and contains a long cirrus, and in the distribution of the vitelline follicles, which surround medially vas deferens near the cirrus-sac. A specimen of Morphotype 1 from B. barbus from the Argens River, France, is designated as neotype of C. brachycollis. The presence of phenotypic plasticity in morphological characteristics previously used for differentiation of species of Caryophyllaeus may confound species identification, which is crucial for biodiversity, ecological and evolutionary studies. To avoid these potential problems, combination of morphological and molecular data is strongly recommended.     

Cytogenetic studies on natural populations of grasshoppers with special reference to B-chromosomes

In Atractomorpha bedeli the frequencies of males with B-chromosomes in the males from the Tofuro-ato population were 21, 30, 32, 40, and 26%, respectively, over five years. Numbers of B's (0 to 7) in the primary spermatocytes were usually constant within the follicles but were variable among the follicles within the individuals. In males from the Okazyo-ato population, 35% contained B's in their primary spermatocytes. The cells of the gastric caeca were observed in 8 males with B's in the primary spermatocytes but no cells of the gastric caeca contained B's. Cells of ovariole walls and those of gastric caeca were analysed in 80 females from the same population; none of them contained B's. These facts indicate that B's are present in the germ line but absent in the somatic line. The difference between the germ line and the somatic one was explained on the basis of elimination of B's from the somatic cells. The inter-follicular variation in number of B's was ascribed to mitotic non-disjunction in the germ line prior to differentiation of the follicles.Contribution No. 86 from the Institute of Biology, Oita University.     

Reproductive performance of partial gonadectomized male African catfish, Clarias gariepinus broodstocks

This study was conducted to evaluate the effect of partial gonadectomy on reproductive performance of male Clarias gariepinus broodstock. Testes from C. gariepinus broodstock were surgically removed; 25% of the testes (Treatment 1), 50% of the testes (Treatment 2), 75% of the testes (Treatment 3), and removal of the sperm from the testes sac using syringe after the abdominal cavity had been cut open (Treatment 4) {control}. The incisions were sutured and the fish kept inside separate concrete tanks for 4 mo. The incisions closed up within 8 to 9 wk of surgery. The postsurgical survival of C. gariepinus was 100%, indicating the efficiency of the surgical procedure. There was no significant difference (P > 0.05) in sperm production, percentage fertilization, hatchability and survival of the larvae using sperm derived from regenerated testes of the partially gonadectomized C. gariepinus and nongonadectomized C. gariepinus. It also reveals that partial gonadectomy could not alter the quality of sperm production of C. gariepinus. Sperm derived from regenerated testes performed effectively for fertilization of eggs. Based on the results of this study, the removal of 75% of testes during partial gonadectomy proved to be the best as the total number of spermatozoa was more than that of other methods and the sperm was able to fertilize more eggs. Hence the removal of 75% of testis during partial gonadectomy of C. gariepinus is recommended based on the results of this study.     

Analysis of the testes of H-Y negative XOSxrb mice suggests that the spermatogenesis gene (Spy) acts during the differentiation of the A spermatogonia

H-Y antigen negative XOSxrb mice, like their H-Y positive XOSxra counterparts, have testes; but, in contrast to XOSxra males, XOSxrb testes almost totally lack meiotic and postmeiotic stages of spermatogenesis. The quantitative analysis of the testes of XOSxrb males and their XY +/- Sxrb sibs, described in the present study, identified two distinct steps in this spermatogenic failure. First, there was a reduction in mitotic activity among T1 prospermatogonia, so that approximately half the normal number of T2 prospermatogonia were produced. Second, there was a dramatic decrease in the number of A3 and A4 spermatogonia and no Intermediate or B spermatogonia. These reductions were also largely due to decreased mitotic activity, there being a shortage of A1 and A2 spermatogonial divisions and no divisions among A3 or A4 spermatogonia. Mitotic activity among the T2 prospermatogonia and the undifferentiated A spermatogonia was normal. This means that the spermatogonial stem cells, which are a subset of the undifferentiated A spermatogonia, are unaffected in XOSxrb mice. Sxrb is now known to have arisen by deletion of DNA from Sxra. It is clear from the present findings that a gene (or genes) present in the deleted DNA plays a major role in the survival and proliferation of the differentiating A spermatogonia.     

Crossing barriers of zoogeographical regions: Molecular evidence of vicariance of the only cestode parasite of loaches (Cobitidae) in the Indomalayan region,Paracaryophyllaeus lepidocephali (Cestoda: Caryophyllidea), and its redescription

Tapeworms of the genus Paracaryophyllaeus Kulakovskaya, 1961 (Cestoda: Caryophyllidea) are specific parasites of loaches (Cypriniformes: Cobitoidea) and occur almost exclusively in the Palaearctic region. The only exception and example of vicariance over the borders of two zoogeographical regions is Paracaryophyllaeus lepidocephali (Kundu, 1985), an insufficiently known species described from the Indomalayan region, with uncertain generic allocation. In the present paper, the species is redescribed based on new material collected from the type host, Lepidocephalichthys guntea, in West Bengal, India. Molecular data reveal this species as a member of Paracaryophyllaeus, within which it is the most closely related to P. cf. gotoi from Misgurnus anguillicaudatus in China and Japan on the basis of large subunit of ribosomal nuclear DNA (28S rDNA). Generic position of P. lepidocephali examined herein is also confirmed by morphology including cross sections, in particular, by a small, cylindrical body, medullary testes with testicular fields crossing the anterior margin of the cirrus sac, medullary vitelline follicles, with some follicles paramuscular, a shallow common genital atrium, short vagina and uterus not extending anteriorly beyond the cirrus sac. It differs from all Palaearctic congeners but Paracaryophyllaeus vladkae Scholz, Oros and Aydogdu, 2014 by the testicular field crossing the anterior margin of the cirrus sac. It differs from P. vladkae by more anterior position of the first vitelline follicles compared to the first testes. This species is a unique example of a fish tapeworm crossing the borders of the Palaearctic and Indomalayan zoogeographical regions.     

On the Origin and Evolution of the Extant System of B Chromosomes in Oryzomyini Radiation (Rodentia,Sigmodontinae)

Heterogeneous supernumerary chromosomes (Bs) are recognized in the oryzomyines Holochilus brasiliensis, Nectomys rattus, N. squamipes, Oligoryzomys flavescens and Sooretamys angouya, representing about 10% of all known B-containing rodent species. They provide an outstanding model for understanding the origin, evolution and diversity of Bs in a phylogenetic context. Therefore, whole chromosome-specific probes were generated from flow-sorted Holochilus brasiliensis (HBR) autosomes 11 and 25+26 and chromosomes X, Y and Bs. Hybridizations were performed on male metaphases of 15 Oryzomyini species of which 3 are B-containing species. The results reveal that among the species sampled, 12 of them, belonging to a monophyletic Oryzomiyini subclade, are positive for an anonymous Oryzomyini shared heterochromatic region (OSHR) on both sex chromosomes. The OSHR is also present on Bs of Holochilus brasiliensis, Nectomys rattus and N. squamipes but not on Bs of O. flavescens and S. angouya. Two distinct additional OSHR/autosome associations are observed on S. angouya. The three species that are OSHR negative belong to an outgroup. Molecular dating suggests that the OSHR originated between 7.8 and 3 Mya on ancestral sex chromosomes. A tentative explanation for the OSHR-positive nature of B regions in three species could be that transposable elements (TEs) from this specific sex chromosome region may have invaded existing B chromosomes. The presence of the OSHR on entire Xp and Yp adjacent to interstitial telomeric sequences at pericentromeric positions, as observed in Drymoreomys albimaculatus, show a similar organization as on B chromosomes in Nectomys squamipes. The diversity of the Oryzomyini Bs in number, size, morphology and genetic content may be explained by the independent origin of B chromosomes in different subgroups of species, with Bs in Holochilus brasiliensis, Nectomys squamipes and N. rattus sharing the OSHR with sex chromosomes, and those in Oligoryzomys flavescens and Sooretamys angouya lacking OSHR in Bs. The species-specific pattern of Bs is probably a consequence of their independent evolutionary origin.     

Cytogenetics of the parthenogenetic grasshopper Warramaba (formerly Moraba) virgo and its bisexual relatives

Hybrids of both sexes were obtained from the reciprocal crosses, carried out in the laboratory, between the species believed to have given rise, by hybridization some thousands of years ago, to the parthenogenetic species W. virgo. All males with a Y-chromosome derived from P196 died before reaching the adult stage, but two males from the reciprocal cross (i.e., with a Y from P169) survived up to the adult stage. Their testes were small but normal in structure and histology. At the first meiotic division almost all the chromosomes were univalents, 0–2 bivalents being formed. Some chiasmata between non-homologous chromosomes were present. Segregation at first anaphase is irregular so that sperms with 3–15 chromosomes are formed. Such hybrids appear to be entirely sterile.     

Cohesin is needed for bipolar mitosis in human cells

Multi-polar mitosis is strongly linked with aggressive cancers and it is a histological diagnostic of tumor-grade. However, factors that cause chromosomes to segregate to more than two spindle poles are not well understood. Here we show that cohesins Rad21, Smc1 and Smc3 are required for bipolar mitosis in human cells. After Rad21 depletion, chromosomes align at the metaphase plate and bipolar spindles assemble in most cases, but in anaphase the separated chromatids segregate to multiple poles. Time-lapse microscopy revealed that the spindle poles often become split in Rad21-depleted metaphase cells. Interestingly, exogenous expression of non-cleavable Rad21 results in multi-polar anaphase. Since cohesins are present at the spindle poles in mitosis, these data are consistent with a non-chromosomal function of cohesin.