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H T Imai

Publications and source records attributed to H T Imai.

At least 19 recordsLinked to original sources

Integrative study on chromosome evolution of mammals, ants and wasps based on the minimum interaction theory.

There is well-known evidence that in many eukaryotes, different species have different karyotypes (e.g. n=1-47 in ants and n=3-51 in mammals). Alternative (fusion and fission) hypotheses have been proposed to interpret this chromosomal diversity. Although the former has long been accepted, accumulating molecular genetics evidence seems to support the latter. We investigated this problem from a stochastic viewpoint using the Monte Carlo simulation method under the minimum interaction theory. We found that the results of simulations consistently interpreted the chromosomal diversity observed in mammals, ants and wasps, and concluded that chromosome evolution tends to evolve as a whole toward increasing chromosome numbers by centric fission. Accordingly, our results support the fission hypothesis. We discussed the process of chromosome evolution based on the latest theory of the molecular structure of chromosomes, and reconfirmed that the fission burst is the prime motive force in long-term chromosome evolution, and is effective in minimizing the genetic risks due to deleterious reciprocal translocations and in increasing the potential of genetic divergence. Centric fusion plays a biological role in eliminating heterochromatin (C-bands), but is only a local reverse flow in contrast to the previously held views.

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Cytological, genetic and evolutionary functions of chiasmata based on chiasma graph analysis.

The nature of the chiasma as a cytological parameter for analysing cross-over was reexamined quantitatively by an improved chiasma graph method. It was reconfirmed in Mus platythrix (n =13) that interstitial chiasmata at diakinesis are distributed randomly and almost uniformly along bivalents except for the centromere and telomere regions. The size of these chiasma blank regions was consistently 0.8% of the total length of haploid autosomes in all chromosomes. There was a minimum value of chiasma interference distance between two adjacent chiasmata, which was constantly 1.8% in all chromosomes. The chiasma frequency at diakinesis was 20.1+/-2. 0 by the conventional method including terminal chiasmata. However, the primed in situ labeling technique revealed that terminal chiasmata were mostly telomere-telomere associations. From these data and also from recent molecular data we concluded that the terminal chiasma is cytologically functional for ensuring the normal disjunction of bivalents at anaphase I, but genetically non-functional for shuffling genes. The chiasma frequency excluding terminal chiasmata was 14.6+/-1.8. Reexamination of the chiasma frequency of 106 animal species revealed that the chiasma frequency increased linearly in proportion to the haploid chromosome number in spite of remarkable difference in their genome size. The increase in chiasma frequency would be evolution-adaptive, because gene shuffling is expected to be accelerated in species with high chromosome numbers.

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Genomic dispersion of 28S rDNA during karyotypic evolution in the ant genus Myrmecia (Formicidae)

The chromosomal localization of 28S rDNA was investigated in 16 speices of the Australian ant genus Myrmecia, with 2n numbers ranging from 4 to 76, using the fluorescence in situ hybridization method and karyographic analysis. A unique phenomenon was observed: the number of chromosomes carrying 28S rDNA increases from 2 in species with low chromosome numbers to 19 in species with high chromosome numbers. This is termed rDNA dispersion. Centric fission and a reciprocal translocation that occurs in C-bands were detected as the major mechanisms involved in rDNA dispersion.

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Mitochondrial-DNA sequence evidence on the phylogeny of Australian jack-jumper ants of the Myrmecia pilosula complex.

Australian ants of the Myrmecia pilosula species complex include some individuals (in M. croslandi) with the lowest possible metazoan chromosome number of 2n = 2. Others in this cluster of sibling species have much higher numbers, the known maximum being 2n = 32. Two species (M. pilosula and M. 'banksi') are believed on cytogenetic and morphological grounds to have hybridized over a long period. To investigate the phylogeny and age of this group relative to the congeneric outgroup species M. gulosa, we sequenced part of the cytochrome b gene and the intergenic sequence between it and a primer anchored on the nearby tRNA(UCNSer) gene and analyzed the coding region using bootstrapped parsimony and neighbor-joining trees using the numbers of synonymous and nonsynonymous codons per site. The intergenic space demonstrated a profusion of repeated sequences, and only very closely related sequences (as judged by that for cytochrome b) showed detectable similarity at this almost 100% A+T region. In agreement with predictions from karyotype studies, the phylogenetic analyses showed that M. croslandi is the sister group to the other siblings; the time of separation of M. croslandi from the rest of the pilosula group is unexpectedly ancient. Other relationships were poorly resolved, but the results suggest that M. 'banski' and M. pilosula cluster together, as expected on cytogenetic grounds, and the tentative suggestion of close affinity of the M. pilosula samples and two "PB" samples supports derivation of PB from female M. pilosula and male M. 'banksi.'

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FISH analysis of the telomere sequences of bulldog ants (Myrmecia: formicidae).

Chromosomes from several species of ants from the genus Myrmecia were hybridized with deoxyoligomer probes of either (T2AG2)7, the putative insect telomere repeat sequence, or (T2AG3)7, the vertebrate telomere repeat sequence. While both sequences hybridized over a range of stringency conditions, (T2AG2)n was clearly the predominant sequence at the termini of the Myrmecia chromosomes. No interstitial sites of either sequence were detected. The genus Myrmecia has a wide range of karyotypes, with chromosome numbers ranging from 2n=2-84. It has been hypothesized that the ancestral karyotype was 2n=4 and karyotype evolution proceeded with an increase in chromosome number. In the absence of detectable interstitial sites of telomere sequence, it is interesting to speculate on the origin of the new telomeres as the chromosome numbers increased.

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Theoretical analyses of chiasmata using a novel chiasma graph method applied to Chinese hamsters, mice, and dog.

Some basic concepts of chiasma (including chiasma distribution, chiasma frequency, interstitial and terminal chiasmata, and chiasma interference) are reexamined theoretically in the light of gene shuffling, and a new method for chiasma analysis termed the chiasma graph is proposed. Chiasma graphs are developed for three mammals with greatly different chromosome numbers: Chinese hamster (with n = 11), mice (n = 20), and a dog (n = 39). The results demonstrate that interstitial chiasmata can contribute both to gene shuffling and to the binding of bivalents, but that so-called terminal chiasmata are in fact mostly achiasmatic terminal associations, the main function of which is to bind bivalents. For this reason, terminal chiasmata should be excluded when chiasma frequency is estimated. It is also demonstrated that interstitial chiasmata distribute on bivalents randomly and uniformly, except at the centromere and telomere. Interference distance fluctuates almost randomly above a minimum value equivalent to about 1.8% of total bivalent length at diakinesis. These results indicate that chiasma formation in mammals is principally a random event. The demonstrated minimum interference distance seems consistent with the polymerization model for chiasma formation. Some cytological aspects of crossing-over are discussed with reference to the minimum interaction theory for eukaryotic chromosome evolution.

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Multiplication of 28S rDNA and NOR activity in chromosome evolution among ants of the Myrmecia pilosula species complex.

Chromosomal localization of rDNA in samples of five taxa of the Mymecia pilosula species complex (Hymenoptera: Formicidae: Myrmeciinae) with 2n = 3 (M. croslandi), 8 (M. imaii), 10 (M. banksi), 18 (M. haskinsorum), and 27 (M. pilosula) was carried out by fluorescence in situ hybridization (FISH) using cloned M. croslandi rDNA (pMc.r2) including the coding region for 28S rRNA. Results show that (1) the 28S rDNA in the genome of these ants is repetitive and is localized in pericentromeric C-bands, (2) the number of chromosomes carrying rDNA is two in M. croslandi, M. imaii and M. banksi, six in M. haskinsorum and ten in M. pilosula, and (3) only one or two clusters of rRNA genes generate nucleoli in each species. We suggest that the rDNA in the ancestral stock of the M. pilosula complex was localized originally in a pericentromeric C-band, and multiplied by chance with time during saltatory increases in C-banding following episodes of centric fission. Most rDNA multiplied on various chromosomes seems to be inactivated and eliminated from the genome, together with C-bands, by AM-inversion or centric fusion, with the remnant rDNAs dispersed in the genome by centric fission and AM-inversion.

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A theoretical approach to chromosome banding pattern analysis.

Based on a schematic model of karyotype evolution, a new methodology for G-, R-, or Q-banding pattern analysis was investigated. Banding pattern analysis essentially depends on the unidirectional alteration and the randomness of the exchange sites of the AM-inversion. In karyotypes that evolved by AM-inversion and Robertsonian rearrangement, two matching patterns appear; (1) tandem and (2) complementary matching patterns. The former is characteristic of a single lineage sharing the same AM-inversions, and the latter appears in different lineages sharing different AM-inversions, by which it is theoretically possible to detect the ancestral karyotype and to reconstruct the karyotype phylogeny (cladogram). In contrast, the evolutionary pathway cannot always be perceived if karyotypes evolve only by Robertsonian rearrangement. The tandem matching pattern does not always mean tandem fusion, but can be interpreted as 'tandem fission' by a combination of AM-inversion and centric fission. Tandem fusion and MM-inversion often cause entangled matching patterns, and thus they interfere with banding pattern analysis. Some methodological problems inherent in the conventional banding pattern analysis are highlighted, and suggested that such problems can be minimized by using the karyograph method. The methodology of banding pattern analysis proposed in the present paper will be applicable for matching the chromosome map of genetic markers among different species.

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An increased level of sperm abnormalities in mice with a partial deletion of the Y chromosome.

Two congenic lines of mice, one with a partial deletion of the Y chromosome, differ in the percentage of spermatozoa with abnormal heads: B10.BR/SgSn males give 22.6% and B10.BR-Ydel/Ms males give 64.2% abnormal sperm. The F1s resulting from crosses of B10.BR/SgSn males with females of five common inbred strains exhibited significantly lower levels of abnormal sperm than the parental strains, as opposed to F1 hybrids sired by B10.BR-Ydel/Ms mutant males, where very high levels of abnormal spermatozoa were found. About 30% of abnormal spermatozoa, produced by males with deletion on the Y chromosome, were characterized by a flat acrosomal cap. This class of abnormality was never observed in non-mutant males, suggesting a mutant-specific defect. These results demonstrate the important role of the Y chromosome in spermatogenesis.

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On the robertsonian polymorphism found in the Japanese raccoon dog (Nyctereutes procyonoides viverrinus).

Karyotypes of 39 Japanese raccoon dogs (NPV) which appeared in the literature and of 7 previously unreported specimens were examined. Thirty four individuals showed the standard karyotype 2K = 26M + 10A + (M)X + (A)Y + Bs (2n = 38 + Bs), where Bs are supernumerary chromosomes. The remaining 11 individuals had 2K = 25M + 12A + XY + Bs (2n = 39 + Bs) and one was 2K = 23M + 16A + XY + Bs (2n = 41 + Bs). The G- and C-banding analyses of both somatic and germ cells revealed that these karyotypes with odd numbers are heterozygous (M/A) for a single Robertsonian rearrangement of chromosomes 2, 5, 6, 8, or 11, and one is M/A heterozygous for three autosomes: 5, 6, and 11.

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Mutability of constitutive heterochromatin (C-bands) during eukaryotic chromosomal evolution and their cytological meaning.

A quantitative analysis of the alterations of constitutive heterochromatin in eukaryotic chromosomal evolution was attempted using the accumulated C-banding data available for mammals, amphibians, fish, ants, grasshoppers, and plants. It was found that these eukaryotes could be classified into two types by their C-banding patterns: 1) Type I included mammals, fish, and ants, and 2) Type II included amphibians, grasshoppers, and plants. C-bands were rather scarce in Type I eukaryote chromosomes and were found around the pericentromeric region when present at all, whereas the predominance of interstitial or terminal C-bands was found in Type II eukaryote chromosomes. The Type I and II C-banding patterns can best be interpreted by assuming that in the former group of eukaryotes the saltatory increase in constitutive heterochromatin occurs preferentially at the pericentromeric regions of telocentric chromosomes induced by centric fission, with C-bands being eliminated almost completely by centric fusion and/or pericentric inversion. On the other hand, C-bands appear in the Type II eukaryotes both interstitially and in the telomeric regions of chromosomes, and there may be no effective mechanism to eliminate these bands once they are integrated.

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Modes of spontaneous chromosomal mutation and karyotype evolution in ants with reference to the minimum interaction hypothesis.

Aspects of chromosomal mutation and karyotype evolution in ants are discussed with reference to recently accumulated karyological data, and to detailed karyotype analyses of several species or species complexes with low chromosome number and unusual chromosomal mutations (the complexes of Myrmecia pilosula (Smith) (n = 1, 5 or 9 to 16); M. piliventris Smith (n = 2, 3-4, 17 or 32), and Ponera scabra Wheeler (n = 3 or 4, 2n = 7 or 8). Translocations and Robertsonian polymorphisms are confirmed to be non-randomly distributed among ants -the former are found at high frequencies in species with low chromosome numbers (n less than or equal to 12), while the latter predominate in those with high numbers (n greater than 12). This situation is consistent with the minimum interaction hypothesis of Imai et al. (1986), under which translocations are expected to occur most frequently in low-numbered karyotypes, and that the resulting genetic risks are minimized by increases in chromosome and/or arm numbers through centric fission and pericentric inversion. Centric fusion is considered to be a transient event in karyotype evolution, resulting from telomere instability in acrocentric chromosomes.

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Theoretical bases for karyotype evolution. II. The fusion burst in man and mouse.

As a theoretical standard for evaluating the high incidence of centric fusion in man and mouse, the relative probabilities of occurrence of reciprocal translocation (Tr), inversion (In) and centric fusion (Fu) were estimated based on the random-contact-and-exchange model. It was shown by this model that centric fusion was extremely rare (Fu = 0.0002, In = 0.0521 and Tr = 0.9477 for a human haploid karyotype). On the other hand, the occurrence rate of centric fusion in human newborn babies and European feral mice was about 500-1,000 times higher than the theoretically expected values, which is termed here the "fusion burst". We suggest that the fusion burst may be induced by the physical proximity of telomeres on the nuclear membrane, and the exchange of DNA strands by errors of telomere replication mechanisms. The cytogenetical significance of the fusion burst is discussed with regard to the minimum interaction hypothesis proposed by Imai et al. (1986). We suggest two closely linked possibilities that (1) the fusion burst in man and mouse can theoretically be placed in karyotype evolution as a transitional phase in the main stream of the fission-inversion cycle, and (2) it may be accelerated by some unknown (mutagenic) factors other than ionizing radiation.

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C-banding analysis of six species of lung flukes, Paragonimus spp. (Trematoda: Platyhelminthes), from Japan and Korea.

We examined C-banded karyotypes of six species of lung flukes from Japan and Korea; diploid and triploid Paragonimus westermani, P. miyazakii, P. ohirai, P. iloktsuenensis and P. sadoensis, with special reference to their karyotypic diversification. C-band analysis between the diploid and the triploid westermani revealed that two of three homologues of the triploid resembled those of the diploid in C-band pattern, while the remaining chromosome showed a different pattern from any species examined here. This karyological evidence indicates that the triploid is allotriploid probably induced by interspecific hybridization between the diploid westermani and an unknown species; we, therefore, suggest that the triploid westermani is an independent species and synonymous with P. pulmonalis (Miyazaki 1978). As the morphologically similar three species, ohirai, iloktsuenensis and sadoensis, had the same C-band polymorphism in chromosome No. 4, these species are classified as the local races of P. ohirai. Paragonimus miyazakii has one common C-band (5q) with the diploid westermani, but other bands (1q, 4q, 6q, 7p and 7q) are different. From these observations, the six species examined are phylogenetically divided into three groups: (1) westermani group containing diploid and triploid (= pulmonalis) species, (2) miyazakii and (3) ohirai including two geographic races, iloktsuenensis and sadoensis.

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Modes of inheritance of X-Y dissociation in inter-subspecies hybrids between BALB/c mice and Mus musculus molossinus.

Genetic analysis of the high frequency of X-Y chromosome dissociation found in primary spermatocytes of F1 hybrids between Japanese wild mice (Mus musculus molossinus) and inbred laboratory mice (BALB/c) was attempted. The frequency of X-Y dissociation (X//Y) in both BALB/c and M. m. molossinus was lower than 30% (Low X//Y), while the value was more than 70% (High X//Y) in their F1 hybrids. Two types of progeny (High X//Y and Low X//Y) appeared in the backcross between BALB/c and High X//Y males, although the frequency of Low X//Y progeny decreased with increasing numbers of backcross generations (26.5% at N2, 13.2% at N3, 5.3% at N4, and 0% at N5). Low X//Y sires produced only Low X//Y mice. We hypothesize that at least one heritable factor which is responsible for the end-to-end association of the sex chromosomes (temporally symbolized as Sxa) is located on the common part of the X and Y chromosomes. The Sxa allele of BALB/c is Sxaa and that of M. m. molossinus is Sxab. The genotype expected in High X//Y males is Sxaa/Sxab and in Low X//Y males and their parental stocks either Sxaa/Sxaa or Sxab/Sxab. The repeated segregation of Low X//Y progeny from High X//Y sires is interpreted simply by assuming that crossing-over has occurred between the X and Y chromosomes. The gradual decrease in the recombinant type mice (Low X//Y) during sequential backcrosses suggests the presence of some autosomal factors that suppress the crossing-over of the sex chromosomes and that do not seem to function in the inter-subspecies hybrids.

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A re-examination of chiasma terminalization and chiasma frequency in male mice.

The distribution and frequency of chiasmata have been analyzed in male BALB/c mice. Bivalents were classified in terms of the number of interstitial chiasmata (CH) and terminal associations (TA) present as follows: 1CH, 2CH, 1TA, 2TA, 1CH X 1TA, 1CH X 2TA, 2CH X 1TA, 2CH X 2TA and 0. We provide evidence that the TA frequently dissociates during 1st meiotic prophase. Consequently six of the observed bivalents may be derived from three basic bivalent types (namely 2CH X 2TA, 1CH X 2TA and 2TA) by dissociation of the TA according to the following schemas: (1) 2CH X 2TA leads to 2CH X 1TA leads to 2CH, (2) 1CH X 2TA leads to 1CH X 1TA leads to 1CH, and (3) 2TA leads to 1TA leads to 0. We also provide evidence that interstitial chiasmata do not move, which implies that a TA can not be formed by chiasma terminalization. The chiasma frequency estimated by assuming that terminal associations do not result from terminalized chiasmata is 17.2+/-2.4 compared to a value of 25.4+/-2.2 calculated on the assumption of chiasma terminalization.

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