Ring chromosomes in barley (Hordeum vulgare L.)

A plant with 2n = 14 + 1 ring chromosomes was obtained in the progeny of a primary trisomie for chromosome 7 of a two-rowed cultivar, Shin Ebisu 16. The morphological characteristics of the trisomic plants with an extra ring chromosome were similar to the primary trisomic for chromosome 7 (Semierect), which suggests that it originated from this chromosome. The ring chromosomes were not completely stable in mitotic cells because of abnormal behavior. Chromosome complements varied in different plants and in different roots within a plant. Root tip cells and spikes with 2n = 14 and 14 + 2 ring chromosomes were observed on plants with 14 + 1 ring chromosomes. Breakage-fusion-bridge cycle was inferred. The ring chromosome was associated with two normal homologues forming a trivalent in 17.6% sporocytes at metaphase I. The transmission of the extra ring chromosome was 23.1% in the progeny of the plant with 14 + 1 ring chromosomes. Trivalent formation may have been much higher at early prophase stages which were difficult to analyze in barley; only 4 of 120 sporocytes analyzed showed an isolated ring at pachytene. The ring chromosome moved to one pole without separation in 24.7% of the sporocytes at AI, and divided in 27.1% sporocytes giving rise to 8-8 separation. Only 10% of the sporocytes showed bridge formation at AI.     

Cytogenetics of a new type of barley with 16 chromosomes

In the progeny of a trisomic type for chromosome 6, Purple, a 16-chromosome type was obtained, which had a pair of new metacentric chromosome 6 in excess. The new metacentric chromosome 6 was shorter than any of the 14 chromosomes of normal barley complement and showed a heteropycnotic nature at late prophase in somatic mitosis. At metaphase I in the plants with 14+one metacentric chromosome 6 (2n=15) the chromosome configuration was exclusively 7_II+1_I indicating that the extra metacentric chromosome 6 could not associate with the normal chromosome 6. At diakinesis and metaphase I in the new 16-chromosome plants most of the sporocytes showed 8_IIor 7_II+2_I. Neither tetravalents nor trivalents were observed at meiosis. The chromosome behaviour at anaphase I and later stages of meiosis was regular in general, resulted in a fairly high pollen fertility of about 61 per cent. Seed fertility however, was very low. The transmission rate of the new metacentric chromosome 6 through the pollen was extremely low in 16-chromosome plants. Possible origin of new basic number and B-chromosome in diploid level through trisomic condition was suggested (Summary see p. 138).Contribution No. 141 of the Department of Plant Science, University of Manitoba.     

Production and identification of primary trisomics in diploid Agropyron cristatum (crested wheatgrass).

Six of the seven possible primary trisomics in Agropyron cristatum were produced. Based on morphology, arm length ratios, and C-banding patterns, they were identified as primary trisomics for chromosomes A, B, C, D, E, and G. Agropyron cristatum is one of several species constituting the crested wheatgrass complex. All species in this complex contain one basic genome (P). A study was conducted to produce and identify a primary trisomic series that will be used to map genes to individual chromosomes. A population of 157 plants were generated by crossing autotriploids (PPP) with diploid (PP) A. cristatum: 58 were diploid (2n = 14), 76 were primary trisomies (2n = 15), 17 were double trisomic (2n = 16), 4 were triple trisomics (2n = 14 + 3), 1 was telocentric trisomic (2n = 14 + 1 telo), and 1 was tetratrisomic (2n = 14 + 4). Karyotype analysis of acetoorcein-stained chromosomes was carried out using the CHROMPAC III computer program; for analysis of C-banded karyotypes, the computer imaging analysis program PCAS (Plant Chromosome Analysis System) was used to identify the primary trisomics. Of the 47 primary trisomics analyzed, 21 plants had one extra satellited chromosome E, 18 with the satellited D chromosome, 3 each for chromosomes B and G, and 1 each for chromosomes C and A. Chromosome pairing was studied in trisomies B, D, E, and G. Trisomics for chromosomes B and G were similar in their mieotic behavior. Each had a trivalent frequency of about 60% and pollen stainability of less than 40%. Trisomics for chromosomes D and E had a trivalent frequency of about 30% and pollen stainability of over 70%.     

Increasing the transmission rate of the extra chromosome in a trisomic Nicotiana sylvestris line by modifying the means of pollination

Summary Trisomies of primary trisomic line B220 of Nicotiana sylvestris, which contain an extra chromosome shown to be a satellite chromosome, can be readily identified by their larger flower and leaf sizes. In seed-propagated species, the low transmission of the extra chromosome has prevented such plants from becoming agriculturally useful cultivars. In line B220, the transfer of the extra chromosome in 2n×2n+1 crosses was very low (13.5%), although n and n + 1 pollen grains were produced in equal quantities, as was confirmed by anther culture. This was due to the delayed development of n + 1 pollen grains, which are not at full maturity at the time of an thesis. The transfer of the extra chromosome in 2n×2n+1 crosses was increased by a 1 day delay in pollination and also by pollination of small pollen grains selected through nylon meshes. The delayed pollination increased the frequency of trisomics by 9%, whereas pollen selected by using 30 and 25 n nylon meshes induced an extremely high transfer of the extra chromosome, namely 51.9% and 70.4%, respectively. The observed frequencies of trisomics and tetrasomics in artificial selfing of 2n+1 plants with selected small pollen grains were lower than those expected from the data of reciprocal crosses between 2n and 2n+1 plants. This discrepancy seems to indicate a disadvantage of the n+1 pollen in fertilization due to the longer style of the trisomics relative to that of the diploids.     

Trisomy in Aedes aegypti.

A trisomic (2n=6+1) pupa of the yellow fever mosquito Aedes aegypti has been found. The trisomy involved chromosome 3 which is intermediate in size between 1 and 2. The extra chromosome formed a univalent or a trivalent during meiosis.     

Effect of substituting chromosome 5A with an supernumary chromosome 5B in offspring of a somaclonal common wheat line monosomic for 5A

Cytogenetic analysis was employed in studying the cause of generation of fertile awned forms in the progeny of plants, which were selected from a speltoid somaclonal wheat line monosomic for chromosome 5A (2n = 41 = 20II + I), had speltoid spikes, and were reproduced by self pollination. On cytogenetic and genetic evidence, chromosome 5A was eliminated and the copy number of chromosome 5B increased in the plants examined. The appearance of an extra chromosome 5B is probably caused by nondisjunction of bivalent 5B in the presence of a telocentric originating from the long arm of chromosome 5A. A difference in meiotic segregation was observed for univalent chromosomes 5A and 5B.     

Aneuhaploids and tetrasomics in rice (Oryza sativa L.) derived from anther culture of trisomics.

Eight types of aneuhaploids (Aneuhaplo 4, 5, 6, 8, 9, 10, 11, and 12) and eight types of tetrasomics (Tetraplo 4, 5, 6, 7, 8, 9, 10, and 12) of rice have been obtained from anther culture of trisomics. This paper reports the plant morphology of these aneuploids and their chromosome behavior at metaphase I. Aneuhaploids for different chromosomes are distinguishable from each other and are morphologically similar to the parental trisomics, suggesting that the extra chromosome has similar genetic effects on plant morphology at the haploid level as at the diploid level. Similarly, tetrasomics with different extra chromosomes are distinguishable from each other and are similar morphologically to the parental trisomic. However, stronger changes in morphological characters were observed in tetrasomics compared with trisomics having the same extra chromosome, as a result of a dosage effect of the extra chromosomes. Comparing plant size between aneuhaploid, tetrasomic, and trisomic with the same extra chromosome, it was shown that the trisomic was the largest, the tetrasomic was of medium size, and the aneuhaploid was the smallest, except for those plants with an extra chromosome 8 in which plant size is dramatically decreased in both the aneuhaploid and the tetrasomic. At metaphase I, aneuhaploids showed chromosome configurations of 1 II + 11 I and 13 I. The frequency of the 1 II + 11 I configuration is higher than 70%, indicating that homologous chromosomes in aneuhaploids tend to stay associated in meiosis. Intragenome chromosome pairing (2 II + 9 I), so called secondary association, was observed in the aneuhaploid for chromosome 5. Tetrasomic plants showed 5 kinds of chromosome configurations: 1 IV + 11 II, 1 III + 11 II + 1 I, 13 II, 12 II + 2 I, and 11 II + 4 I. A chromosome configuration of 13 II was often observed in tetrasomics with shorter extra chromosomes and a chromosome configuration of 1 IV + 11 II was often observed in tetrasomics with longer extra chromosomes. Aneuhaploids had complete seed sterility. Tetrasomics showed very poor pollen fertility and complete seed sterility, except for a few shriveled seeds that were observed in Tetraplo 6 and 9. This is the first report in rice where many aneuhaploids and tetrasomics have been characterized. This information will help to further unravel rice aneuploidy and cytogenetics. The aneuploids obtained here will be very useful tools for the study of genetics and breeding in rice.     

Maize tertiary trisomic stocks derived from B-A translocations

Reciprocal translocations between supernumerary B chromosomes and the basic complement of A chromosomes in maize have resulted in a powerful set of tools to manipulate the dosage of chromosomal segments. From 15 B-A reciprocal translocation stocks that have the B-A chromosome genetically marked we have developed tertiary trisomic stocks. Tertiary trisomics are 2n + 1 aneuploids where the extra chromosome is a translocation element, in this case a B-A chromosome. Whereas B-A translocations produce aneuploidy in the sperm, the tertiary trisomic plant efficiently transmits hyperploid gametes maternally. Because the B-A tertiary trisomic stocks and the B-A translocation stocks from which they were derived are introgressed into the W22 inbred line, the effects of maternally and paternally transmitted trisomic B-A chromosomes can be compared. Data are presented on both the male and female transmission rates of the B-A chromosomes in the tertiary trisomic stocks.     

Meiotic studies on a Brassica campestris-alboglabra monosomic addition line and derived B. campestris primary trisomics.

Diakinesis chromosomes were studied in pollen mother cells of Brassica campestris (2n = 20, genome AA), B. alboglabra (2n = 18, genome CC), a B. campestris-alboglabra monosomic addition line (AA + 1 chromosome from the C genome), and four derived B. campestris primary trisomics. The nucleolar chromosomes of B. campestris were distinguishable by their morphology at diakinesis. The alien C-genome chromosome in the addition line paired preferentially with the nucleolar chromosome of the A genome. Very rarely, it paired with another pair of the A genome. Thus, it was concluded that the alien C-genome chromosome of the addition line is primarily homoeologous to the nucleolar chromosome and secondarily to another chromosome of the A genome. Three of the four derived B. campestris trisomic plants were identified as B campestris nucleolar trisomics. Trisomy in the fourth plant involved another chromosome. The cytological mechanism underlying the origin of trisomics in the addition line and chromosome homoeology relationships between B. campestris and B. alboglabra are envisaged.     

Chromosome evolution by robertsonian translocation in Gibasis (Commelinaceae)

The plant species Gibasis schiedeana (Kunth) D. R. Hunt sens. lat. contains two cytotypes viz. a self-sterile diploid with 2n=10 (x=5) and a selffertile cytological autotetraploid with 2n=16 (x=4). Single chromosome sets of these plants consist of 2 metacentrics +3 acrocentrics, and 3 metacentrics +1 acrocentric chromosomes respectively suggesting a Robertsonian relationship between them. Their artificial F₁ hybrids show the pairing of acrocentrics with metacentric arms confirming the supposed nature of the chromosome affinities. Both breeding systems and ploidy levels show that the direction of the change has been from x=5 to x=4 by a translocation of the Robertsonian type.     

Chromosome elimination in trisomics of Coix aquatica Roxb.

Summary Somatic chromosome elimination was identified and its patterns studied in a trisomic (2n=11) with marker genes in Coix aquatica Roxb. In a cross between a recessive trisomic with green base and white style (ccc ii ss) and a dominant disomic having purple base and purple style (CC II SS), all the F₁ seedling progeny were purple based because of the presence of C, I and S. For C. to be expressed in seedling base, either I should be absent or S should be present with I. In style colour, however, irrespective of the presence of I and S, C produces purple phenotype.In one trisomic (Ccc ii ss) plant (designated as 4–15) of the F₁ progeny, a part of the seedling base was green. All the tillers coming up from the green side of the main tiller also had green base, and those arising on the purple side were purple based. Similarly, the pistillate spikelets on the green side of the main culm and on the tillers with green base were white styled, and the male spikes showed 10 chromosomes. Female spikelets on the purple side of the main tiller and on the tillers with purple base were mostly purple styled and the male spikes had 11 chromosomes. In some of the purple based tillers, however, there were both 11 and 10 chromosomes in different regions or different inflorescence clusters on the tiller. In these tillers, where the chromosome number was 11, style colour was purple, and white style occurred when there were 10 chromosomes. In one tiller, the style colour was purple but the chromosome number was 10.The recessive phenotype of the style in the trisomic conceivably resulted from an elimination of the extra chromosome carrying the dominant allele C. On the basis of the morphological features of the extra chromosome, such as length, centromere position and distribution pattern of the hetero and eupycnotic regions, it was identified as chromosome No. 2 in the complement. It was therefore possible to place with certainty the gene c on this chromosome. Sometimes, however, the extra chromosome carrying c also was eliminated giving 10 chromosomes and purple style.In the other trisomic plants of the F₁ progeny, one plant showed 11 chromosomes but in a tiller there were only 10 chromosomes and white styles. In two other plants, although the chromosome number was 11 throughout, white style was present in a single cluster of inflorescences in one plant, and in one pistillate spikelet in the other. In the latter two cases, white style was believed to have arisen as a result of a mutation from C to c or somatic crossing over, giving the genotype ccc in the affected regions. In a single plant, chromosome elimination was observed in only one cell.Apparently the 10-chromosome sectors arose from the 11-chromosome condition by selective elimination of the extra chromosome during mitosis in the primordium giving rise to these sectors. In the affected plants, elimination did not obviously occur at the same stage but at different times in their ontogeny. Instability is probably governed by one or a few major genes, associated with a number of modifiers, exhibiting incomplete penetrance and variable expression. Chromosome elimination did not apparently follow any particular pattern but was erratic. Probably some intracellular environment is necessary to trigger the mechanism governing the elimination into action. The unstable system, occurring in combination with other favourable features like the functional nature of the aneuploid gametes, sexual reproduction, monoecious condition favouring cross pollination and tolerance of extra chromosomes by the sporophyte, could be an important factor in the cytogenetic evolution of the species.     

Variable karyotype with B-chromosomes in Bellevalia saviczii (Liliaceae)

Bellevalia saviczii (Liliaceae), a hexaploid (2n = 6x = 24), was collected from mountain slopes in the Shiraz valley, southern Iran. The basic karyotype consists of one long metacentric chromosome, a slightly shorter acrocentric, and two shorter submetacentrics. In 32% of the plants, one and sometimes two of the acrocentrics (IIa) appeared with a pericentric inversion that changed it into a metacentric (IIm). This metacentric IIm was present in a Hardy--Weinberg ratio throughout the collection range of more than 300 km. A number of aneuploids were found, pentasomics [2n – 1 = 23 (4.9%)], heptasomics [2n + 1 = 25 (2.5%)], and one octasomic [2n + 2 = 26 (0.3%)]. Re-assorted karyotypes with 2n = 24, but with odd numbers of two of the chromosomes, were also present (6.6%). B-chromosomes were found in root meristems and in pedicels in a number (19.1%) of the plants that grew around a 30 km marsh. Their numbers ranged from 1B to 8Bs per plant peaking in the 2B mode. The B-chromosomes (two polymorphs with terminal and near-terminal sticking points) were telocentric with a few metacentric iso-Bs. No B-carrying plants were found in the drier regions away from the marsh. The B-frequency distribution for aneuploids and pericentric inversions was much different than for the standard euploid. Dr Mark Gettner died on November 2nd, 2002. He is fondly remembered by his family, his many students and friends for his kindness and generosity, and for his dedication to his scientific work. Dr Broun is the son-in-law of Dr Gettner and he will be the corresponding author for this article (pb22@york.ac.uk).     

加拿大引进的二倍体燕麦种质的核型鉴定

采用常规压片法对砂燕麦、西班牙燕麦和短燕麦3个二倍体燕麦种进行了核型研究。结果表明:砂燕麦染色体核型公式为2n=2x=14=10m+4sm(2SAT),具近中部和中部着丝点染色体,第4对染色体组的短臂上有1对随体,核不对称系数为68.17%;西班牙燕麦染色体核型公式为2n=2x=14=10m+4sm(2SAT),具近中部和中部着丝点染色体,第7对染色体短臂上有1对随体,核不对称系数为59.31%;短燕麦染色体核型公式为2n=2x=14=6m+4sm+4st(2SAT),具近端部、近中部和中部着丝点染色体,第6对染色体组的短臂上有1对随体,核不对称系数为63.91%。虽然3个燕麦种的核型均为2A,但它们的染色体形态有明显不同,比较认为砂燕麦相对进化,短燕麦次之,西班牙燕麦较原始。本研究对燕麦种质资源的核型分析及进化地位研究具有参考价值。     

Tertiary trisomics in the garden pea as a model of B chromosome evolution in plants

It is hypothesized that, in plants, genetically empty B chromosomes may originate from the extra chromosome (E) of tertiary trisomics if (i) the region of basic chromosomes homologous to the E (H-region) harbors a sporophytic lethal covered by the wild-type allele in E, and (ii) crossing-over between E and the H-region is suppressed. Under these conditions, most loss-of-function mutations occurring in the H-region are deleterious for haploid gametophytes, whereas those occurring in E are neutral or advantageous for hyperploid (n+1) gametophytes. As a result, natural selection at the gametophyte level can lead to the degeneration of E, leaving the H-region intact. Using Hammarlund translocation T(3-6)a, we synthesized two trisomic lines of the garden pea (Pisum sativum L.), where E was composed of the short arms of chromosomes 3 and 6 and the H-region carried recessive markers. In the trisomic line TRIS, we found few crossovers between E and the H-region. In the trisomic line TRUST, obtained after a change of basic chromosome constitution, recombination in this region was completely suppressed. After induction in the H-region of TRUST of a recessive sporophytic mutation rmv, two 15-chromosome lines of stable trisomics were established. One of them passed 11 generations, having produced more than 6000 individuals, all of them trisomic, and E remained present as a single element with no pairing partners. No tetrasomics were detected in these lines. If such trisomics occurred in nature, their extra chromosomes are likely to become a B chromosome.     

广东四个墨兰品种的核型研究

研究了广东四个墨兰品种金嘴墨兰、银边墨兰、企剑黑墨和企剑白墨的染色体数目和核型。结果表明,企剑黑墨和企剑白墨的染色体数目为2n=40,为二倍体,核型公式分别为：2n=2x=40=30m＋10sm和2n=2x=40=2M＋36m＋2st;金嘴墨兰和银边睾兰的染色体数目为2n=41,为非整倍体。4个墨兰品种的染色体结构主要由中部着丝粒染色体组成,除银边墨兰为1B型外,其它均为2B型。     

The Effect of Substitution of Chromosome 5A with an Extra Chromosome 5B in the Progeny of a Somaclonal Common Wheat Line Monosomic for 5A

Cytogenetic analysis was employed in studying the cause of generation of fertile awned forms in the progeny of plants, which were selected from a speltoid somaclonal wheat line monosomic for chromosome 5A (2n = 41 = 20II + I), had speltoid spikes, and were reproduced by self pollination. On cytogenetic and genetic evidence, chromosome 5A was eliminated and the copy number of chromosome 5B increased in the plants examined. The appearance of an extra chromosome 5B is probably caused by nondisjunction of bivalent 5B in the presence of a telocentric originating from the long arm of chromosome 5A. A difference in meiotic segregation was observed for univalent chromosomes 5A and 5B.     

Occurrence of macro B chromosomes in Astyanax scabripinnis paranae (Pisces,Characiformes, Characidae)

B chromosomes occur in several Neotropical fish species. Cytogenetic analysis of 27 specimens (15 females and 12 males) of Astyanax scabripinnis paranae from the Araquá river (a small headwater tributary of the Tietê river) shows that this population has 2n=50 chromosomes (4M+30 SM+4ST+12A), two chromosome pairs with NORs and conspicuous C-band positive blocks in the terminal position of the long arm of four chromosome pairs. In this population, eight females presented 2n=51 chromosomes and the extra chromosome was a large metacentric similar in size and morphology to the first chromosome pair in the karotype. This accessory chromosome is entirely heterochromatic in C-banded metaphases and shows a late replication pattern evidenced by BrdU incorporation. There was no significant correlation between the presence of B chromosomes and increased NOR activity at the P>0.05 level. Some aspects related to these B chromosomes are discussed.     

Determination of n+1 Gamete Transmission Rate of Trisomics and Location of Gene Controlling 2n Gamete Formation in Chinese Cabbage (Brassica rapa)

A set of trisomics of Chinese cabbage was used for determining the n+1 gamete transmission rate and locating the gene controlling 2n gamete formation on the corresponding chromosome. The results showed that the transmission rates of extra chromosomes in different trisomica varied from 0% to 15.38% by male gametes and from 0% to 17.39% by female gametes. Of the nine F2 populations derived from the hybridizations between each triaomic and Bp058 (2n gamete material), only Tri-4×Bp058 showed that the segregation ratio of plants without 2n gamete formation to plants with 2n gamete formation was 10.38:1, which fitted the expected segregation ratio of the trisomics (AAa) based on the 7.37% of n+1 gamete transmission through female and 5.88% through male. In other populations the segregation ratios varied from 2.48:1 to 3.72:1, which fitted the expected 3:1 segregation ratio of the bisomice (Aa). These results suggested that the gene controlling 2n gamete formation in Chinese cabbage Bp058 was located on chromosome 4. Further trisomic analysis based on the chromosome segregation and the incomplete stochastic chromatid segregation indicated that the gene locus was tightly linked to the centromere.     

The Biology of Allium monanthum (Liliaceae) I. Polyploid Complex and Variations in Karyotype

Abstract Plants of Allium monanthum Maxim., whose gender expression are usually dioecious, but rarely hermaphrodite or gynomonoecious, proved to constitute a polyploid complex, consisting of diploid, triploid, and tetraploid individuals. The basic chromosome complement of this species consists of seven metacentric or submetacentric chromosomes and one acrocentric, the latter possessing a satellite on the short arm. Thus, the karyotype formula is expressed as 7V+11 (x=8). The diploid plants (2n = 16) were confined to central Honshu, Japan. Typical female plants possessed the standard karyotype, whereas male plants were heterozygous for two kinds of translocations. The 3x plants (2n=24) are somewhat widely distributed in the areas from the Kanto to Hokuriku district in Honshu. All female triploid plants possessed the standard karyotype. The geographical distribution of 4x plants (2n=32) which express mostly a female phenotype occurred nearly throughout the whole areas investigated; they are geographically isolated from the 2x plants. A majority of 4x plants had the standard karyotype. The remaining tetraploids were of the aberrant type, 4x/51, which has five acrocentric chromosomes, and two aneuploids 4x+1 and 4x-1. Both 3x and 4x forms seem to be of autopolyploid origin. Three kinds of aberrant nucleolar chromosomes with an extra satellite or an inseried secondary constriction were found in the heterozygotes for translocations of 2x plants and also in some plants of the 3x form. These aberrant plants usually form their own homogeneous populations, but were somewhat scattered throughout the range in their distribution. Thus, these individuals are considered to have perpetuated these types of chromosome aberrations which originated in the remote past.     

3个彩色马蹄莲引进品种的核型分析

利用普通压片法对3个引进彩色马蹄莲(Zantedeschia hybrid)品种的染色体数与核型进行了分析。结果表明:所试验品种染色体数均为2n=32。染色体形态比较一致,多是由中部(m)以及近中部(sm)着丝粒染色体组成。其中,‘Allure’为2n=2x=32=14m(2SAT)+2sm,‘Cupdio’的核型公式为2n=2x=32=14m+2sm,Odessa的核型公式为2x=32=1M+15m(1SAT)。3个品种核型不对称系数分别为56.72%,56.25%和56.38%,核型分类显示其均为1A型。