Incipient Genome Differentiation in Gossypium. III. Comparison of Chromosomes of G. HIRSUTUM and Asiatic Diploids Using Heterozygous Translocations

Hybrids between upland cotton (G. hirsutum, genome constitution 2A_hD_h) and either A-genome or D-genome diploid species exhibit 26 paired and 13 unpaired chromosomes at metaphase I. The A_h and D_h genomes are therefore considered homoeologous with those of the respective diploids. Previous studies, nevertheless, revealed a low level of ("incipient") differentiation between D_h and various diploid D genomes. The diploid A genomes have been regarded as more closely homologous to A_h on the basis of low preferential pairing and autotetraploid segregation ratios in allohexaploids.—The present study addressed the following questions: Are the diploid A genomes differentiated from A_h in meiotic homology? If so, is the differentiation manifested equally by all 13 chromosomes or is it localized in certain chromosomes?—Three diploid A-genome lines representing G. herbaceum and G. arboreum were hybridized by in ovulo culture of embryos (1) with a standard line of G. hirsutum, which differs from G. herbaceum by two and from G. arboreum by three naturally occurring reciprocal translocations involving chromosomes 1–5, and (2) with six lines homozygous for experimental translocations involving chromosomes 6, 7, 10, 11, 12 and 13. Chiasma frequencies in hybrids were compared with those in appropriate G. hirsutum controls. In every comparison overall chiasma frequencies were slightly lower in the hybrids. Therefore A_h appears to be differentiated from the diploid A genomes. No localized differentiation was detected in chromosomes marked by experimental translocations. The differentiation may be localized mainly in chromosomes 4 and 5.     

Spatial and temporal expression of the two sucrose synthase genes in maize: immunohistological evidence

Summary The DNAs of two diploid species of Gossypium, G. herbaceum var. africanum (A₁ genome) and G. raimondii (D₅ genome), and the allotetraploid species, G. hirsutum (A_h and D_h genomes), were characterized by kinetic analyses of single copy and repetitive sequences. Estimated haploid genome sizes of A₁ and D₅ were 1.04 pg and 0.68 pg, respectively, in approximate agreement with cytological observations that A genome chromosomes are about twice the size of D genome chromosomes. This differences in genome size was accounted for entirely by differences in the major repetitive fraction (0.56 pg versus 0.20 pg), as single copy fractions of the two genomes were essentially identical (0.41 pg for A₁ and 0.43 pg for D₅). Kinetic analyses and thermal denaturation measurements of single copy duplexes from reciprocal intergenomic hybridizations showed considerable sequence similarity between A₁ and D₅ genomes (77% duplex formation with an average thermal depression of 6 °C). Moreover, little sequence divergence was detectable between diploid single copy sequences and their corresponding genomes in the allotetraploid, consistent with previous chromosome pairing observations in interspecific F₁ hybrids.Journal paper No. 4461 of the Arizona Agricultural Experiment Station     

Incipient Genome Differentiation in Gossypium. II. Comparison of 12 Chromosomes in G. HIRSUTUM, G. MUSTELINUM and G. TOMENTOSUM Using Heterozygous Translocations

Eight homozygous translocation lines (TT) of G. hirsutum marking 3 chromosomes of the A genome and 9 chromosomes of the D genome were crossed with G. hirsutum, G. mustelinum and G. tomentosum, all homozygous for the standard end arrangements (tt). Chiasma frequencies in the G. hirsutum Tt controls were compared with those in the G. hirsutum x G. mustelinum and the G. hirsutum x G. tomentosum Tt hybrids. Both nucleus-wide and region-specific chiasma frequencies were compared.—Some genome differentiation appears to have arisen between G. hirsutum and G. mustelinum. The G. hirsutum x G. mustelinum hybrids had a 1.8 to 1.9% reduction in the nucleus-wide chiasma frequency. Four of the eight TT lines showed a 3.4 to 10.5% reduction in chiasmata in the hybrid translocation quadrivalents, suggesting that chromosomes 1, 21, 23 and 24 may have undergone localized genome differentiation. The two species may differ naturally in the end arrangement of two chromosomes, since a quadrivalent not due to experimentally introduced translocations was observed in 13% of the PMC's of two G. hirsutum x G. mustelinum hybrids.—Very little genome differentiation has occurred between G. hirsutum and G. tomentosum. In the G. hirsutum x G. tomentosum hybrids, the nucleus-wide estimates showed only a very small (0.1 to 0.2%), though statistically significant, lowering of the chiasma frequency, and there was no reduction in chiasma frequency in the more sensitive readings for specific translocation quadrivalents.     

Chromosomal Locations of 5S and 45S rDNA in Gossypium Genus and Its Phylogenetic Implications Revealed by FISH

We investigated the locations of 5S and 45S rDNA in Gossypium diploid A, B, D, E, F, G genomes and tetraploid genome (AD) using multi-probe fluorescent in situ hybridization (FISH) for evolution analysis in Gossypium genus. The rDNA numbers and sizes, and synteny relationships between 5S and 45S were revealed using 5S and 45S as double-probe for all species, and the rDNA-bearing chromosomes were identified for A, D and AD genomes with one more probe that is single-chromosome-specific BAC clone from G. hirsutum (A₁D₁). Two to four 45S and one 5S loci were found in diploid-species except two 5S loci in G . incanum (E₄), the same as that in tetraploid species. The 45S on the 7th and 9th chromosomes and the 5S on the 9th chromosomes seemed to be conserved in A, D and AD genomes. In the species of B, E, F and G genomes, the rDNA numbers, sizes, and synteny relationships were first reported in this paper. The rDNA pattern agrees with previously reported phylogenetic history with some disagreements. Combined with the whole-genome sequencing data from G . raimondii (D₅) and the conserved cotton karyotype, it is suggested that the expansion, decrease and transposition of rDNA other than chromosome rearrangements might occur during the Gossypium evolution.     

Transmission of duplication-deficiencies from cotton translocations is unrelated to map lengths of the unbalanced segments

Adjacent-1 duplication-deficiencies (dp-dfs) are readily recovered from most heterozygous translocations in Gossypium hirsutum L., but frequencies of specific cytotypes differ widely in progenies from heterozygote (♀) x standard crosses. Surprisingly, these frequencies seem to be unrelated to the primary (postmeiotic) frequencies predicted by metaphase I configurations or to the proportion of the chromosome arm that is duplicate or deficient. Deficiencies and duplications from different translocations involving the same arm, as well as the two complementary dp-dfs from the same translocation, seldom exhibit similar frequencies. We conclude that the frequency of each of 101 different adjacent-1 cytotypes is largely idiosyncratic and may depend in part on interactions between the specific chromosome regions that are respectively trisegmental and monosegmental. Few, if any, of these interactions can be between homoeologues of the A_h and D_h genomes. Adjacent-2 dp-dfs are seldom recovered, even if they involve chromosomes that are readily tolerated in monosomic condition. Comparison of monosomes and telosomes with deficiencies suggests that some chromosomes and chromosome regions may be more dosage-sensitive than others, but their identification is not strongly supported by these data.     

Chromosome relationships between Lolium and Festuca (Gramineae)

With a view to eclucidating chromosome relationships between Lolium perenne (Lp), L. multiflorum (Lm) and Festuca pratensis (Fp), chromosome pairing in different diploid (2n=14), auto-allotriploid (2n=3x=21), trispecific (2n=3x=21), amphidiploid (2n=4x=28) and auto-allohexaploid (2n=6x=42) hybrids between them was analysed. At all these levels of ploidy there was very good chiasmate pairing between the chromosomes of the three species and, on the whole, there was little evidence of preferential pairing of the chromosomes of a particular species in the triploid, tetraploid and hexaploid hybrids. A critical test for this also came from the synaptic ability of the chromosomes of the single genome with those of the duplicated genome in the auto-allotriploids which formed predominantly trivalents with 2, 3 or even 4 chiasmata. Moreover, the homology between the Lp and Lm chromosomes seems strong enough to pass the discrimination limits of the B-chromosomes which do not suppress homoeologous pairing in the Lp LmLm triploid and LpLm diploid hybrids. — The triploids having two genomes of a Lolium species and one of F. pratensis had some male and female fertility which suggested genetic compatibility of the parental chromosomes resulting, presumably, in compensation at the gametic level. Also, the occurrence of comparable chiasma frequencies in the auto-allotriploids and trispecific hybrids showed that they were not markedly affected whether two doses of one genome and one of the other or all the three different genomes from the three species were present. From the trend of chromosome pairing in all these hybrids it is concluded that there is little structural differentiation between the chromosomes of the three species, no effective isolation barrier to gene-flow between them, and that they are closely related phylogenetically, having possibly evolved from a common progenitor. Taxonomic revision of the two Lolium species is suggested.     

The genomic relationship between Glycine max (L.) Merr. and G. soja Sieb. and Zucc. as revealed by pachytene chromosome analysis

Summary This study was conducted with the objective of determining the genomic relationship between cultivated soybean (Glycine max) and wild soybean (G. soja) of the subgenus Soja, genus Glycine. Observations on cross-ability rate, hybrid viability, meiotic chromosome pairing, and pollen fertility in F ₁ hybrids of G. max × G. soja and reciprocals elucidated that both species hybridized readily and set mature putative hybrid pods, generated vigorous F₁ plants, had a majority of sporocytes that showed 18II + 1IV chromosome association at diakinesis and metaphase I, and had a pollen fertility that ranged from 49.2% to 53.3%. A quadrivalent was often associated with the nucleolus, suggesting that one of the chromosomes involved in the interchange is a satellited chromosome. Thus, G. max and G. soja genetic stocks used in this study have been differentiated by a reciprocal translocation. Pachytene analysis of F₁ hybrids helped construct chromosome maps based on chromosome length and euchromatin and heterochromatin distribution. Chromosomes were numbered in descending order of 1–20. Pachytene chromosomes in soybean showed heterochromatin distribution on either side of the centromeres. Pachytene analysis revealed small structural differences for chromosomes 6 and 11 which were not detected at diakinesis and metaphase I. This study suggests that G. max and G. soja carry similar genomes and validates the previously assigned genome symbol GG.Research supported in part by the Illinois Agricultural Experiment Station and U.S. Department of Agriculture Competitive Research Grant (85-CRCR-1-1616)     

Spatiotemporal Asymmetry of the Meiotic Program Underlies the Predominantly Distal Distribution of Meiotic Crossovers in Barley

Meiosis involves reciprocal exchange of genetic information between homologous chromosomes to generate new allelic combinations. In cereals, the distribution of genetic crossovers, cytologically visible as chiasmata, is skewed toward the distal regions of the chromosomes. However, many genes are known to lie within interstitial/proximal regions of low recombination, creating a limitation for breeders. We investigated the factors underlying the pattern of chiasma formation in barley (Hordeum vulgare) and show that chiasma distribution reflects polarization in the spatiotemporal initiation of recombination, chromosome pairing, and synapsis. Consequently, meiotic progression in distal chromosomal regions occurs in coordination with the chromatin cycles that are a conserved feature of the meiotic program. Recombination initiation in interstitial and proximal regions occurs later than distal events, is not coordinated with the cycles, and rarely progresses to form chiasmata. Early recombination initiation is spatially associated with early replicating, euchromatic DNA, which is predominately found in distal regions. We demonstrate that a modest temperature shift is sufficient to alter meiotic progression in relation to the chromosome cycles. The polarization of the meiotic processes is reduced and is accompanied by a shift in chiasma distribution with an increase in interstitial and proximal chiasmata, suggesting a potential route to modify recombination in cereals.     

Multidisciplinary approach to genome analysis in the diploid species,Thinopyrum bessarabicum and Th. elongatum (Lophopyrum elongatum), of the Triticeae

Summary The J and E genome species of the Triticeae are invaluable sources of salt tolerance. The evidence concerning the phyletic relatedness of the J genome of diploid Thinopyrum bessarabicum and the E genome of diploid Th. elongatum (=Lophopyrum elongatum) is discussed. Low level of chromosome pairing between J and E at different ploidy levels, suppression of J-E pairing by the Ph1 pairing regulator that inhibits homoeologous pairing, complete sterility of the diploid hybrids (JE), karyotypic divergence of the two genomes, differences in total content and distribution of heterochromatin along their chromosomes, and marked differences in gliadin proteins, isozymes, 5S DNA, and rDNA indicate that J and E are distinct genomes. Well-defined biochemical markers have been identified in the two genomes and may be useful in plant breeding. The level of distinction between J and E is comparable to that among the universally accepted homoeologous genomes A, B, and D of wheat. Therefore, the J and E genomes are homoeologous and not homologous, although some workers continue to call them homologous. The previous workers' data on chromosome pairing in diploid hybrids and/ or karyotypic differences in the conventionally stained chromosomes do not provide sufficient evidence for the proposed merger of J and E genomes (and, hence, of the genera Thinopyrum and Lophopyrum) specifically and for establishing genome relationships generally. Extra precautions should be exercised before changing the designation of an established genome and before merging two genera. A uniform, standardized system of genomic nomenclature for the entire Triticeae is proposed, which should benefit cytogeneticists, plant breeders, taxonomists, and evolutionists.Cooperative investigations of the USDA-Agricultural Research Service and the Utah Agricultural Experiment Station, Logan, UT 84322, USA. Approved as Journal Paper no. 3832     

Chromosome-Specific Painting in Cucumis Species Using Bulked Oligonucleotides

Chromosome-specific painting is a powerful technique in molecular cytogenetic and genome research. We developed an oligonucleotide (oligo)-based chromosome painting technique in cucumber (Cucumis sativus) that will be applicable in any plant species with a sequenced genome. Oligos specific to a single chromosome of cucumber were identified using a newly developed bioinformatic pipeline and then massively synthesized de novo in parallel. The synthesized oligos were amplified and labeled with biotin or digoxigenin for use in fluorescence in situ hybridization (FISH). We developed three different probes with each containing 23,000–27,000 oligos. These probes spanned 8.3–17 Mb of DNA on targeted cucumber chromosomes and had the densities of 1.5–3.2 oligos per kilobases. These probes produced FISH signals on a single cucumber chromosome and were used to paint homeologous chromosomes in other Cucumis species diverged from cucumber for up to 12 million years. The bulked oligo probes allowed us to track a single chromosome in early stages during meiosis. We were able to precisely map the pairing between cucumber chromosome 7 and chromosome 1 of Cucumis hystrix in a F₁ hybrid. These two homeologous chromosomes paired in 71% of prophase I cells but only 25% of metaphase I cells, which may provide an explanation of the higher recombination rates compared to the chiasma frequencies between homeologous chromosomes reported in plant hybrids.     

All paired up with no place to go: pairing, synapsis, and DSB formation in a balancer heterozygote

The multiply inverted X chromosome balancer FM7 strongly suppresses, or eliminates, the occurrence of crossing over when heterozygous with a normal sequence homolog. We have utilized the LacI-GFP: lacO system to visualize the effects of FM7 on meiotic pairing, synapsis, and double-strand break formation in Drosophila oocytes. Surprisingly, the analysis of meiotic pairing and synapsis for three lacO reporter couplets in FM7/X heterozygotes revealed they are paired and synapsed during zygotene/pachytene in 70%–80% of oocytes. Moreover, the regions defined by these lacO couplets undergo double-strand break formation at normal frequency. Thus, even complex aberration heterozygotes usually allow high frequencies of meiotic pairing, synapsis, and double-strand break formation in Drosophila oocytes. However, the frequencies of failed pairing and synapsis were still 1.5- to 2-fold higher than were observed for corresponding regions in oocytes with two normal sequence X chromosomes, and this effect was greatest near a breakpoint. We propose that heterozygosity for breakpoints creates a local alteration in synaptonemal complex structure that is propagated across long regions of the bivalent in a fashion analogous to chiasma interference, which also acts to suppress crossing over.     

Comparative hybridization reveals extensive genome variation in the AIDS-associated pathogen Cryptococcus neoformans

Background

Genome variability can have a profound influence on the virulence of pathogenic microbes. The availability of genome sequences for two strains of the AIDS-associated fungal pathogen Cryptococcus neoformans presented an opportunity to use comparative genome hybridization (CGH) to examine genome variability between strains of different mating type, molecular subtype, and ploidy.

Results

Initially, CGH was used to compare the approximately 100 kilobase MAT a and MATα mating-type regions in serotype A and D strains to establish the relationship between the Log2 ratios of hybridization signals and sequence identity. Subsequently, we compared the genomes of the environmental isolate NIH433 (MAT a) and the clinical isolate NIH12 (MATα) with a tiling array of the genome of the laboratory strain JEC21 derived from these strains. In this case, CGH identified putative recombination sites and the origins of specific segments of the JEC21 genome. Similarly, CGH analysis revealed marked variability in the genomes of strains representing the VNI, VNII, and VNB molecular subtypes of the A serotype, including disomy for chromosome 13 in two strains. Additionally, CGH identified differences in chromosome content between three strains with the hybrid AD serotype and revealed that chromosome 1 from the serotype A genome is preferentially retained in all three strains.

Conclusion

The genomes of serotypes A, D, and AD strains exhibit extensive variation that spans the range from small differences (such as regions of divergence, deletion, or amplification) to the unexpected disomy for chromosome 13 in haploid strains and preferential retention of specific chromosomes in naturally occurring diploids.     

Genetic Lengths and Break Points in Twelve Chromosomes of GOSSYPIUM HIRSUTUM Involved in Ten Reciprocal Translocations

Chromosome configurations were recorded in about 5500 pollen mother cells (PMC's) in 2n and 2n-1 (missing the intact A-genome chromosome) heterozygotes of ten reciprocal translocations involving six A-genome chromosomes (H1, H2, H3, H4, H6 and H7) and six D-genome chromosomes (H14, H15, H16, H19, H20 and H21) of Gossypium hirsutum. From these records, chiasma frequencies at each of six positions were determined for nine translocations and at two positions for one. These frequencies were used to calculate recombination frequencies in different chromosome regions, and from these distances the breakpoints in 15 chromosome arms were mapped relative to each other and to their respective centromeres, insofar as the data permitted. The karyotype so derived for twelve chromosomes is in reasonably good agreement with data from genetic mapping, telosome and monosome mapping, and the mitotic idiogram.     

SUPERNUMERARY CHROMOSOMES IN TRADESCANTIA II. EFFECT OF COLCHICINE ON MEIOTIC ISOCHROMOSOME CONFIGURATIONS

The effect of colchicine on meiotic pairing and configuration frequencies of three homologous isosupernumerary chromosomes was investigated. In the absence of colchicine, the three isochromosomes displayed a high degree of interchromosomal pairing and chiasma formation. As a consequence, a high frequency of bivalents and trivalents were observed at diakinesis-metaphase I. The unique structure of isochromosomes enables them to pair intrachromosomally (i.e., foldback pairing) yet the preferential occurrence of interchromosomal pairing suggests that all six arms of the three isosupernumerary chromosomes were in close association prior to or upon initiation of synapsis. Supernumerary chromosomes in microsporocytes treated during presynapsis or early synapsis with colchicine exhibited a significant reduction (P < 0.001) in the number of bivalents and trivalents at diakinesis. However, there was no reduction in overall chiasma frequency among supernumeraries due to the induction of increased intrachromosomal pairing and chiasma formation. A colchicine-sensitive association or alignment of homologues preceding effective pairing has been demonstrated in standard chromosomes of a number of plant species. This study provides the first evidence to indicate that at least certain supernumerary chromosomes may display presynaptic association as well. The results also support the strongly held contention that colchicine is not directly preventing or inhibiting the actual formation of chiasmata, since no reduction in chiasma frequency was observed in the isochromosomes.     

Chromosome pairing relationships among the A,B, and D genomes of bread wheat

Summary Chromosome pairing and chiasma frequency were studied in bread wheat euhaploids (2n = 3x = 21; ABD genomes) with and without the major pairing regulatorPh1. This constitutes the first report of chromosome pairing relationships among the A, B, and D genomes of wheat without the influence of an alien genome. AllPh1 euhaploids had very little pairing, with 0.62–1.05 rod bivalents per cell; ring bivalents were virtually absent and mean arm-binding frequency (c) values ranged from 0.050 to 0.086. In contrast, theph1b euhaploids had extensive homoeologous pairing, with chiasma frequency 7.5–11.6 times higher than that in thePh1 euhaploids. They had 0.53–1.16 trivalents, 1.53–1.74 ring bivalents, and 2.90–3.57 rod bivalents, withc from 0.580 to 0.629. N-banding of meiotic chromosomes showed strongly preferential pairing between chromosomes of the A and D genomes; 80% of the pairing was between these genomes, especially in the presence of theph1b allele. The application of mathematical models to unmarked chromosomes also supported a 21 genomic structure of theph1b euhaploids. Numerical modeling suggested that about 80% of the metaphase I association was between the two most related genomes in the presence ofph1b, but that pairing under Ph1 was considerably more random. The data demonstrate that the A and D genomes are much more closely related to each other than either is to B. These results may have phylogenetic significance and hence breeding implications.This paper is dedicated to the memory of the late Ernest R. SearsCooperative investigations of the USDA-Agricultural Research Service and the Utah Agricultural Experiment Station, Logan, UT 84322, USA. Approved as Journal Paper No. 3986     

Comparative Genome Analyses Highlight Transposon-Mediated Genome Expansion and the Evolutionary Architecture of 3D Genomic Folding in Cotton

Transposable element (TE) amplification has been recognized as a driving force mediating genome size expansion and evolution, but the consequences for shaping 3D genomic architecture remains largely unknown in plants. Here, we report reference-grade genome assemblies for three species of cotton ranging 3-fold in genome size, namely Gossypium rotundifolium (K₂), G. arboreum (A₂), and G. raimondii (D₅), using Oxford Nanopore Technologies. Comparative genome analyses document the details of lineage-specific TE amplification contributing to the large genome size differences (K₂, 2.44 Gb; A₂, 1.62 Gb; D₅, 750.19 Mb) and indicate relatively conserved gene content and synteny relationships among genomes. We found that approximately 17% of syntenic genes exhibit chromatin status change between active (“A”) and inactive (“B”) compartments, and TE amplification was associated with the increase of the proportion of A compartment in gene regions (∼7,000 genes) in K₂ and A₂ relative to D₅. Only 42% of topologically associating domain (TAD) boundaries were conserved among the three genomes. Our data implicate recent amplification of TEs following the formation of lineage-specific TAD boundaries. This study sheds light on the role of transposon-mediated genome expansion in the evolution of higher-order chromatin structure in plants.     

The relation between synapsis and chiasma formation in Rhoeo spathacea

Rhoeo spathacea (Schwartz) Stearn (2n=12) is a complete translocation heterozygote in which a ring of 12 chromosomes is often observed from diakinesis through metaphase I. A new hypotonie bursting technique on primary microsporocytes made possible visualization of the complete pattern of pachytene synapsis in R. spathacea by light and electron microscopy. Analysis showed that most of the length of a chromosome never synapsed and that relational twisting of the lateral elements was prominent in the distal segments of most synaptonemal complexes (SCs). The most common situation was 12 relatively short SCs representing the 24 synapsed terminal segments of the 12 chromosomes; 11 and 10 SCs were also observed but less frequently, indicating one and two synaptic failures, respectively. Similarly, in diakinesis a ring of 12 chromosomes was most common, but a chain of 12 chromosomes and two chains of chromosomes could also occur. These situations indicated zero, one, and two chiasma failures, respectively. The frequencies of synaptic failure (0.021) and of chiasma failure (0.022) were essentially the same as were the distributions of cells with different numbers of synaptic and chiasma failures. These comparisons indicate that if synapsis occurs between two distal segments of the chromosomes of R. spathacea, a chiasma will almost certainly follow. A diagram to illustrate the arrangement of mid-pachytene chromosomes and a model to explain the origin of relational twisting of lateral elements in SCs are presented.     

Synapsis with and without recombination in the male meiosis of the leaf-footed bug <Emphasis Type="Italic">Holhymenia rubiginosa</Emphasis> (Coreidae,Heteroptera)

In organisms with chiasmatic meiosis two different relationships have been described between crossing over and synapsis: in one group of organisms synapsis depends on the initiation of meiotic recombination while in the other group it is independent of this initiation. These patterns have been observed mainly in organisms where all meiotic bivalents in the set have similar behaviors. In some heteropteran insects a pair of chromosomes named m chromosomes is known to behave differently from autosomes regarding synapsis and recombination. Here we used immunodetection of a synaptonemal complex component and acid-fixed squashes to investigate the conduct of the small m chromosome pair during the male meiosis in the coreid bug Holhymenia rubiginosa. We found that the m chromosomes form a synaptonemal complex during pachytene, but they are not attached by a chiasma in diakinesis. On the other hand, the autosomal bivalents synapse and recombine regularly. The co-existence of these variant chromosome behaviors during meiosis I add further evidence to the absence of unique patterns regarding the interdependence of synapsis and recombination.     

The quantitative analysis of chromosome pairing and chiasma formation based on the relative frequencies of M I configurations. II. Primary trisomics

On an earlier occasion the estimation of the “crossing-over potentials” of the two arms of a chromosome from the relative frequencies of the different types of bivalent was discussed. When in normal diploids the bivalents in M I can not be distinguished from each other, primary trisomics may be used to mark particular chromosomes. For the estimation of the “crossingover potentials” of the two arms the only type of configuration available (trivalent) is not sufficient. The required additional information may be obtained from the average probability (b) of the chromosome arms to be bound. Using the satellite chromosome trisomic ofSecale cereale it is shown that the use ofb introduces an excessive error, especially with highb values. Consequently, the primary trisomics can not be recommended for the estimation of the “crossing-over potentials” of the two arms of particular chromosomes. The telocentric trisomics permit the recognition of more configurations and are to be preferred. Partner exchange and nucleolus may both interfere with chiasma formation. The data suggest that the three homologous chromosomes may not be equal in respect to pairing and chiasma formations. There are indications that the initiation of pairing is localized in one locus on each arm.