Interaction between rye B-chromosomes and wheat genetic systems controlling homoeologous pairing

The interactive effect on homoeologous pairing of rye B-chromosomes with the absence of both pairing suppressor (3A, 3D, 5B) and promotor (3B, 5A, 5D) chromosomes of common wheat (Triticum aestivum L.) is analyzed by comparison of pairing at Metaphase I of 27-, 27+2B, 28- and 28+2B-chromosome plants. These plants were obtained from crosses between the respective wheat monosomics (2n=41) and rye plants (Secale cereale L.) carrying or not carrying two B-chromosomes (2n=14 or 14+2Bs). —The effect of rye B-chromosomes on pairing depends on the function of the wheat chromosome which is absent in the appropriate hybrids, i.e., rye B-chromosomes have a suppressor effect on pairing when the pairing suppressing wheat chromosomes 3A, 3D or 5B are absent, while they behave as promotors when the pairing promoting chromosomes 3B, 5A or 5D are absent.     

Genomic constitutions of four Chinese endemic Elymus species: E. brevipes, E. yangii, E. anthosachnoides, and E. altissimus (Triticeae, Poaceae).

The objectives of this study were to determine the genomic constitution and to explore the genomic variation within four Chinese endemic Elymus species, i.e., E. brevipes (Keng) L?ve (2n = 4x = 28) and E. yangii B.R. Lu (2n = 4x = 28), E. anthosachnoides (Keng) L?ve (2n = 4x = 28), and E. altissimus (Keng) L?ve (2n = 4x = 28). Intraspecific crosses between different populations of the four Elymus species, as well as interspecific hybridizations among the four target species, and with six analyzer species containing well-known genomes, i.e., E. caninus (L.) L. (2n = 4x = 28, SH), E. sibiricus L. (2n = 4x = 28, SH), E. semicostatus (Lees ex Steud.) Melderis (2n = 4x = 28, SY), E. parviglumis (Keng) L?ve (2n = 4x = 28, SY), E. tsukushiensis Honda (2n = 6x = 42, SHY), and E. himalayanus (Nevski) Tzvelev (2n = 6x = 42, SHY), were achieved through the aid of embryo rescue. Chromosome pairing behaviors were studied in the parental species and their hybrids. Numerical analysis on chromosome pairing was made on the interspecific hybrids. With one exception, each meiotic configuration at metaphase I in the hybrids involving the target taxa and the analyzer species containing the "SH" genomes fit a 2:1:1 model with x-values ranging between 0.91 and 1.00; chromosome pairing in the hybrids involving analyzer parents with the "SY" genomes match a 2:2 model, with x-values between 0.97 and 0.99. All pentaploid hybrids with a genomic formula "SSYYH," except for two crosses having unexpected low c-values, had pairing patterns fitting the 2:2:1 model with x-values varying between 0.96 and 1.00. It is concluded based on hybridization, fertility, and chromosome pairing data that (i) the four target Elymus species are strictly allotetraploid taxa, (ii) they are closely related species, all comprised of the "SY" genomes, (iii) minor genomic structural rearrangements have occurred within the four Elymus species, and (iv) meiotic pairing regulator(s) exists in some of the Elymus taxa studied.     

Synthesis and cytological characterization of trigeneric hybrids of durum wheat with and without Ph1.

Wild grasses in the tribe Triticeae, some in the primary or secondary gene pool of wheat, are excellent reservoirs of genes for superior agronomic traits, including resistance to various diseases. Thus, the diploid wheatgrasses Thinopyrum bessarabicum (Savul. and Rayss) A. Love (2n = 2x = 14; JJ genome) and Lophopyrum elongatum (Host) A. Love (2n = 2x = 14; EE genome) are important sources of genes for disease resistance, e.g., Fusarium head blight resistance that may be transferred to wheat. By crossing fertile amphidiploids (2n = 4x = 28; JJEE) developed from F1 hybrids of the 2 diploid species with appropriate genetic stocks of durum wheat, we synthesized trigeneric hybrids (2n = 4x = 28; ABJE) incorporating both the J and E genomes of the grass species with the durum genomes A and B. Trigeneric hybrids with and without the homoeologous-pairing suppressor gene, Ph1, were produced. In the absence of Ph1, the chances of genetic recombination between chromosomes of the 2 useful grass genomes (JE) and those of the durum genomes (AB) would be enhanced. Meiotic chromosome pairing was studied using both conventional staining and fluorescent genomic in situ hybridization (fl-GISH). As expected, the Ph1-intergeneric hybrids showed low chromosome pairing (23.86% of the complement), whereas the trigenerics with ph1b (49.49%) and those with their chromosome 5B replaced by 5D (49.09%) showed much higher pairing. The absence of Ph1 allowed pairing and, hence, genetic recombination between homoeologous chromosomes. Fl-GISH analysis afforded an excellent tool for studying the specificity of chromosome pairing: wheat with grass, wheat with wheat, or grass with grass. In the trigeneric hybrids that lacked chromosome 5B, and hence lacked the Ph1 gene, the wheat-grass pairing was elevated, i.e., 2.6 chiasmata per cell, a welcome feature from the breeding standpoint. Using Langdon 5D(5B) disomic substitution for making trigeneric hybrids should promote homoeologous pairing between durum and grass chromosomes and hence accelerate alien gene transfer into the durum genomes.     

Meiotic pairing in hybrids between tetraploid Triticale and related species: new elements concerning the chromosome constitution of tetraploid Triticale

Summary Two F5 strains of tetraploid triticale (2n= 4x=28), obtained from 6x triticaleX2 rye progenies, were crossed with diploid and tetraploid rye, some durum and bread wheats, and various 8x and 6x triticale lines. Meiosis in the different hybrid combinations was studied. The results showed that the haploid complement of these triticales consists of seven chromosomes from rye and seven chromosomes from wheat. High frequencies of PMCs showing trivalents were observed in hybrids involving the reference genotypes of wheat and triticale. These findings proved that several chromosomes from the wheat component have chromosome segments coming from two parental wheat chromosomes. The origin of these heterogeneous chromosomes probably lies in homoeologous pairing occurring at meiosis in the 6x triticaleX2x rye hybrids from which 4x triticale lines were isolated. A comparison among different hybrids combinations indicated that the involvement of D-genome chromosomes in homoeologous pairing is quite limited. In contrast, meiotic patterns in 4x triticale X 2x rye hybrids showed a quite high pairing frequency between some R chromosomes and their A and B homoeologues.     

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.     

小麦族鹅观草属三种植物的生物系统学研究

本文研究了禾本科小麦族鹅观草属的3个种:缘毛鹅观草(Roegneria pendulina Neuski 2n=4x=28),纤毛鹅观草(R. ciliaris (Trin.) Nevski 2n=4x=28)和鹅观草(R. kamoji Ohwi 2n=6x=42)及其种间杂种的形态变异和染色体配对行为。各杂种F_1的减数分裂染色体配对数较高,但杂种高度不育。在杂种减数分裂中还观察到一定频率的多价体形成。以上结果充分表明该3种植物享有两个共同的基本染色体组,在S和Y染色体组之间发生过染色体相互易位,缘毛鹅观草的染色体组可拟定为S~PY~P。     

Synthesis and cytological characterization of trigeneric hybrids involving durum wheat,Thinopyrum bessarabicum,and Lophopyrum elongatum

Summary In an attempt to transfer genes for salt tolerance and other desirable traits from the diploid wheatgrasses, Thinopyrum bessarabicum (2n=2x=14; JJ genome) and Lophopyrum elongatum (2n=2x=14; EE genome), into durum wheat cv Langdon (2n=4x=28; AABB genomes), trigeneric hybrids with the genomic constitution ABJE were synthesized and cytologically characterized. C-banding analysis of somatic chromosomes of the A, B, J, and E genomes in the same cellular environment revealed distinct banding patterns; each of the 28 chromosomes could be identified. They differed in the total amount of constitutive heterochromatin. Total surface area and C-banded area of each chromosome were calculated. The B genome was the largest in size, followed by the J, A, and E genomes, and its chromosomes were also the most heavily banded. Only 25.8% of the total chromosome complement in 10 ABJE hybrids showed association, with mean arm-pairing frequency (c) values from 0.123 to 0.180 and chiasma frequencies from 3.36 to 5.02 per cell. The overall mean pairing was 0.004 ring IV + 0.046 chain IV + 0.236 III + 0.21 ring II + 2.95 rod II + 20.771. This is total pairing between chromosomes of different genomes, possibly between A and B, A and J, A and E, B and J, B and E, and J and E, in the presence of apparently functional pairing regulator Ph1. Because chromosome pairing in the presence of Ph1 seldom occurs between A and B, or between J and E, it was inferred that pairing between the wheat chromosomes and alien chromosomes occurred. The trigeneric hybrids with two genomes of wheat and one each of Thinopyrum and Lophopyrum should be useful in the production of cytogenetic stocks to facilitate the transfer of alien genes into wheat.     

加拿大披碱草和圆柱披碱草BC1植株的细胞遗传学鉴定

以加拿大披碱草(2n=4x=28)和圆柱披碱草(2n=6x=42)为材料,对其BC1植株染色体数目、配对构型以及花粉育性和结实性等进行了鉴定。结果显示:BC1代85%以上细胞染色体数目为28条(2n=4x=28);BC1植株的花粉母细胞减数分裂中期Ⅰ染色体配对行为较规则,其平均染色体构型分别为0.04Ⅰ 13.98Ⅱ,且环状二价体多于棒状二价体;BC1植株的花粉可育率和自然结实率分别为84.83%和70.98%,说明BC1植株的育性已得到恢复,为其后代优良株系的选育奠定了基础。     

Prevalence of B chromosomes in Orthoptera is associated with shape and number of A chromosomes

We analyze the prevalence of B chromosomes in 1,601 species of orthopteran insects where chromosome number and shape are known. B chromosomes have been reported in 191 of these species. Bs are not uniformly distributed among orthopteran superfamilies, with evident hotspots in the Pyrgomorphoidea (32.3% of species carrying Bs), Grylloidea (14.9%), Acridoidea (14.6%) and Tetrigoidea (14.3%). As expected under the theory of centromeric drive, we found a correlation between B chromosome presence and A chromosome shape-Bs are more frequent in karyotypes with more acrocentric A chromosomes. We also found that Bs are less common in species with high chromosome numbers and appear to be most common at the modal chromosome number (2n = 24). Study effort, measured for each genus, was not associated with B prevalence, A chromosome shape or A chromosome number. Our results thus provide support for centromeric drive as an important and prevalent force in the karyotypic evolution of Orthoptera, just as it appears to be in mammals. We suggest that centromeric drive may provide a mechanistic explanation for White's principle of karyotypic orthoselection.     

Polyploidy and karyotype variation in Turkish Bellevalia (Hyacinthaceae)

There are 20 Bellevalia species in Turkey, half of which are endemic. Chromosome numbers are known for 15 species. A chromosome survey of 145 Bellevalia individuals showed that the karyotype is remarkably stable. All are based on x  = 4. The majority are diploid with 2 n  = 8, but there is also a polyploid series of 2 n  = 16, 24 and 32. Aneuploidy occurs only at the octoploid level. Eleven individuals had metacentric B chromosomes, one had acrocentric Bs and one had telocentric Bs. Bellevalia pycnantha and B. paradoxa are morphologically similar, with B. pycnantha reduced to a synonom of B. paradoxa .  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 87–98.     

赖草属5个种的核型与进化

报道了国产赖草属５个种的核型,即大赖草,２ｎ＝４ｘ＝２８＝２４ｍ（２ＳＡＴ）＋４ｓｍ（２ＳＡＴ）;粗穗赖草２ｎ＝４ｘ＝２８＝２２ｍ（２ＳＡＴ）＋４ｓｍ＋２ｓｔ（２ＳＡＴ）;若羌赖草,２ｎ＝４ｘ＝２８＝２０ｍ（４ＳＡＴ）＋６ｓｍ＋２ｓｔ（２ＳＡＴ）;羊草,２ｎ＝４ｘ＝２８＝２２ｍ（４ＳＡＴ）＋２ｓｍ＋４ｓｔ（４ＳＡＴ）;窄颖赖草,２ｎ＝４ｘ＝２８＝２２ｍ（２ＳＡＴ）＋４ｓｍ（２ＳＡＴ）＋２ｓｔ（２Ｓ     

东北蒿属艾组12种植物核型研究

报道东北蒿属艾组12种植物的染色体数及核型资料,结果如下：宽叶山蒿（Artemisia stolonifera (Maixm.)Komar.)2n=4x=36=28m 4sm(4SAT) 4st(4SAT);野艾蒿（A.lavandulaefolia DC)2n=2x=50=40m 4sm 6st(6SAT); 矮蒿（A.lancea Van.)2n=2x=16=10m 4sm 2st;蒙古蒿（A.mongolica (fisch.ex Bess.)Nakai]2n=2x=16=14m 2st;红足蒿（A.rubripes Nakai)2n=2x=16=14m 2st;歧茎蒿（A.igniaria Maixm.)2n=2x=34=28m 2sm 4st(4SAT);柳叶蒿（A.integrifolia Linn.)2n=4x=36=20m 12sm 4st;线叶蒿（A.subula 4st(4SAT);柳叶蒿（A.integrifolia Linn.)2n＝4x=36=20m 12sm 4st;线叶蒿（A.subulata Nakai)2n=2x=16=14m 2st;高岭蒿（A.brachyphylla Kitam.)2n=2x=18=14m 2sm 2st;林文稿[A.uiridissima(Komar.)Pamp.]2n=2x=18=14m 4sm;奄癌（A.keiskeana Miq.)2n=2x=18=16m 2sm;阴地蒿（A.sylvatica Maxim.)2n=2x=16=14m 2sm,核型对称性野艾蒿（A.lavandulaefolia)为2B型,其余均为2A型,依据核型资料对个别种的演化 分类进行了初步探讨。     

我国某些蔷薇属花卉的核型研究

本文报道了我国产的5种和14个品种的蔷薇属花卉的染色体数目和核型,结果如下:小花型为二倍体,2n=2x=14,少数为混倍体;中花型为三倍体,2n=3x=21;大花型为四倍体,2n=4x=28。大部分种的核型均由其中部和近中部着丝点染色体组成,少数种具近端着丝点染色体。它们可以区分为3种核型类型,即1A、2A和1B。     

7种橐吾属植物的核型

研究了7种橐吾属（Ligularia）植物的染色体和核形态。干崖子橐吾（L．kanaitzensis）的核型为2n＝2x＝58＝26m＋28sm 4st;窄头橐吾（L.stenocephala）的核型为2n=2x=58=26m 32sm;细茎橐吾（L.hookeri）的核型为2n=2x=58=30m 26sm 2st;宽戟橐吾（Llatihastata）的核型为2n=2x=58=28m 26sm(2sat) 4st;网脉橐吾（L.dictyoneura）的核型为2n=2x=58=26m 28sm 2st 2t;蹄橐吾（L.hodgsonii）的核型为2n=2x=58=28m 28sm 2t;棉毛橐吾（L.vellerea）的核型为2n=2x=58=22cm 34sm 2t。虽然这7个种的染色体数目相同,2n=58,核型主要是由m和sm染色体构成,但各类的染色体数目在种间有差异。核的对称性高,着丝点端值（T．C）为61．45％－64．96％。除窄头橐吾和鹿蹄橐吾的染色体数与前人报道的相同外,其它5个种的染色体数目和核型为首次报道。     

Chromosome pairing in triploid intergeneric hybrids ofFestuca pratensis withLolium multiflorum, revealed by GISH

Genomic in situ hybridisation (GISH) was used to reveal chromosome pairing in two partly fertile, triploid (2n = 3x = 21) hybrids obtained by crossing the diploid (2n = 2x = 14) Festuca pratensis Huds. (designated FpFp), used as a female parent, with the autotetraploid (2n = 4x = 28) Lolium multiflorum Lam. (designated LmLmLmLm), used as a male parent. The pattern of chromosome pairing calculated on the basis of the mean values of chromosome configurations identified in all 100 PMCs analysed, was: 0.71I Lm + 2.24I Fp + 2.18II Lm/Lm + 0.54II Lm/Fp + 4.18III Lm/Lm/Fp. A relatively high number of Lm/Lm bivalents and Fp univalents, and a low number of Lm/Fp bivalents and Lm univalents indicated that the pairing was preferential between L. multiflorum chromosomes. Other observations regarding chromosome pairing within the Lm/Lm/Fp trivalents also confirmed this preferential pairing in the analysed triploids, as the Fp chromosome was not randomly located in the chain- and frying-pan-shaped trivalents. The similarities and differences in chromosome pairing at metaphase I and the level of preferential pairing between Lolium chromosomes in the different triploid Lolium-Festuca hybrids are discussed.     

Analysis of meiosis in triticale (XTriticosecale Wittmack) X rye (Secale cereale L.) F1 hybrids at three ploidy levels

Summary Triticales (XTriticosecale Wittmack) at three ploidy levels (8x, 6x, 4x, x=7) were crossed with diploid rye (Secale cereale L.) to produce a solitary hypopentaploid hybrid (2n=32), and a number of tetraploid (2n=4x=28) and triploid (2n=3x=21) hybrids. The hybrids exhibited a morphology which was intermediate between the parents. The number of bivalents ranged from 1–7 (4.65 per cell) in hypopentaploid, from 2–12 (7.13 per cell) in tetraploid and from 4–9 (6.84 per cell) in triploid hybrids. In 4x and 3x hybrids, trivalents and quadrivalents were also observed at low frequencies (range 0–1; mean 0.01–0.03 per cell). Chiasmata frequency was highest in triploid hybrids (12.44 per cell), lowest in hypopentaploid (5.37 per cell) and intermediate in tetraploids (10.54 per cell). More than 7¹¹ were found in 39.7% pollen mother cells (PMC's) in the 4x hybrids and in 5.0% PMCs in 3x hybrids. It is concluded that an increase in the relative proportion of wheat chromosomes in the hybrids had a slight suppression effect on homologous as well as homoeologous pairing of rye chromosomes. Contrary to this, the relative increase in rye complement promoted homoeologous pairing between wheat chromosomes. In triploid hybrids, the chiasmata frequency as well as the c value were the highest, suggesting that in tetraploid hybrids rye chromosomes had a reduced pairing (low frequency of ring bivalents).     

Two new Tibetan species of Elymus (Poaceae: niticeae) and their genomic relationships

Two Elymus species, E. cacuminus (2n = 4x = 28) and E. retroflexus (2n = 2x = 28). native to western China are described as species new to science. In order to determine genomic constitutions of the two species, intergeneric and interspecific hybridizations were carried out between the two new species and Pseudoroegneria spicata (2n = 2x = 14, SS), and sixteen other Elymus species, fifteen tetraploids (2n = 4x = 28) and one hexaploid (2n = 6x = 42), containing the "SH", "SY", and "SYH" genomes. Chromosome pairing behaviour was studied in the two species and their hybrids. Generally low meiotic pairing was observed in the hybrids with S- and SH-genome species and high pairing in the hybrids with SY- and SYH-genome species. It is concluded from this study that (i) the two new species are strict allotetraploid (2n = 4x = 28) and self-pollinating taxa (inbreeders) in nature; (ii) they contain the "SY" genomes and; (iii) they have close genomic relationships with the SY-genome Elymus species from the same or geographically adjacent areas.     

B chromosome variation in Euphydryas colon (Lepidoptera: Nymphalidae)

Cytogenetic analysis of an Idaho population of the checkerspot butterfly, Euphydryas colon, has revealed considerable inter- and intra-individual variation in chromosome number which turns out to be a classic case of B chromosome variation. The basic chromosome complement of the species is n (, )=31. The A chromosomes were aligned equatorially at mitotic metaphase and metaphase II, and axially at metaphase I, indicating a restriction of centric activity at the first meiotic division. No failure of pairing between homologous A chromosomes was observed and, although a marked asynchrony of chromatid separation was found to be characteristic of mitotic telophase and telophase II, the frequency of macrospermatid formation was low. The B chromosomes were at least partly heterochromatic but exhibited some variation in both pycnosity and size. Mitotically stable B-containing individuals showed a preponderance of unpaired Bs at first metaphase and these divided at either first or second anaphase. The presence of Bs was associated with a heightened production of abnormal spermatids particularly in individuals with high numbers of B chromosomes. Among the 25 individuals sampled, 21 carried from 1–6 B chromosomes, and of these 14 were mitotically stable. In all 7 unstable individuals the mean number of B chromosomes per cell exceeded the modal number. Assuming that the modal number represents the zygotic number, these results suggest that a mechanism to boost the number of B chromosomes exists in males of E. colon.     

Synaptic patterns of rye B chromosomes. II. The effect of the standard B chromosomes on the pairing of the A set

In order to elucidate the possible effects of rye B chromosomes (Bs) on synapsis and metaphase-I associations of the A set, a comparative study between pachytene and metaphase-I-cells of rye plants carrying different numbers of Bs (0–8) has been carried out. The number of Bs was found to be positively correlated with the frequency of synaptic irregularities of the A set, i.e. multivalents and foldback pairing, and with the frequency of pachytene interlockings. It is proposed that interlockings are the origin of these irregularities because both appeared in close proximity in many nuclei. Examples of A-B pairing are described. The frequency of synaptic abnormalities seems to be unrelated to the mean of A chromosome-bound arms at metaphase I.     

Contributions to the cytòtaxonomy of the Commelinaceae

Representatives of the Tradescantia geniculata alliance (genus Gibasis Rafin.) in cultivation at Kew display considerable cytological and morphological diversity. Chromosome complements consist of either large or small chromosomeS. Plants with small chromosomes have either 2 n = 16 or 2 n = 32, and all form only bivalents at meiosis and have presumed basic numbers of x – 8 and x = 8 or 16 respectively. The 2 n = 32 accessions are considered to represent T.geniculata in its strict sense; the identity of the 2 n =16 plant is uncertain. Plants with large chromosomes have either 2 n = 16 or 2n = 20 and each forms quadrivalents to the extent that they can be considered as cytological autotetraploidS. Basic numbers are then x = 4 and x =5, the lowest yet recorded for the Commelinaceae. The identity of the x = 4 species is uncertain. The x = 5 plants are readily identified as T.karwinskyana.