Morphology of the pachytene chromosomes ofPennisetum purpureum schumach

The morphology of the pachytene chromosomes ofPennisetum purpureum was studied. On the basis of relative lengths and arm ratios it was possible to identify them individually. They are numbered 1 to 14 in the order of their decreasing lengths and diagnostic characters are given for each chromosome. The first and the fourteenth chromosomes, the longest and the shortest of the complement respectively, are the nucleolus organizing chromosome. In all the fourteen chromosomes the centromeres are flanked by deep staining regions. This feature is common to the two species.P. typhoides andP. purpureum. Seven out of the fourteen chromosomes have terminal knobs in their long arms. This is a feature in which it differs from the other seven chromosome species.Pennisetum typhoides. These knobs, however, are not present uniformly in all populations; material obtained from Ghana did not show these knobs.     

Haploid meiosis and its bearing on the phylogeny of pearl millet,Pennisetum typhoides stapf et hubb.

Meiotic chromosome associations in a spontaneously originated haploid plant of pearl millet have been studied and their phyletic significance discussed. Chromosome pairing could be observed at pachytene and diplotene. Out of a total of 285 PMC's studied at diakinesis-metaphase 1, 43 showed one bivalent and 7 had two bivalents per cell. Both rod- and ring-bivalents were observed. Apart from synapsis accompanied by chiasma formation, close associations of univalent chromosomes were observed. Out of 150 cells without true bivalents, 41 showed 1 s-s association and five, 2 s-s pairs per cell. On the basis of the realization of a maximum of two bivalents per cell, as also of a maximum of 2 s-s pairs, it has been inferred that the chromosome complement ofP. typhoides (n=7) has evolved from a basic set ofn=5 chromosomes. Other available evidence supporting this inference is also discussed.     

Interspecific hybridization inCoix 1. Morphological and cytological studies of the hybrids of a new form ofCoix with 2n=32×Coix aquatica Roxb

A newly recorded form ofCoix with a haploid chromosome number (n=16) that is not an entire multiple of the presumed base numberx=5 andCoix aquatica (n=5) hybridize spontaneously. The hybrid plants display heterosis with regard to plant height, leaf and fruit size and anthocyanin pigmentation. All plants studied showed a total complement of 21 chromosomes. The karyotype has been classified into three size groups: long, medium and small. The chromosomes of the two parents show little homology and on the average only two bivalents are produced. Meiosis is abnormal. Only 4% viable pollen is produced and the plants are completely seedless. The production of sterile hybrids is taken to indicate a remote relationship between the genomes of the two species.Part of a Doctoral thesis approved by the University of Agra.     

Cytogenetics of double cross hybrids between Pennisetum americanum — P. purpureum amphiploids and P. americanum x Pennisetum squamulatum interspecific hybrids

Summary Pearl millet, Pennisetum americanum L. Leeke-napiergrass, Pennisetum purpureum Schum. amphiploids (2n=42) were crossed with pearl millet X Pennisetum squamulatum Fresen. interspecific hybrids (2n=41) to study the potential of germplasm transfer from wild Pennisetum species to pearl millet. These two interspecific hybrids were highly cross-compatible and more than two thousand trispecific progenies were produced from 17 double crosses. All doublecross hybrids were perennial and showed a wide range of morphological variations intermediate to both parents in vegetative and inflorescence characteristics. Some crosses resulted in sublethal progenies. Chromosomes paired mainly as bivalents (¯x15.88) or remained as univalents. At metaphase I, trivalents, quadrivalents, an occasional hexavalent and a high frequency of bivalents indicated some homeology among the genomes of the three species. Delayed separation of bivalents, unequal segregation of multivalents, lagging chromosomes, and chromatin bridges were observed at anaphase I. Although approximately 93% of the double-cross hybrids were male-sterile, pollen stainability in male-fertile plants ranged up to 94%. Seed set ranged from 0 to 37 seed per inflorescence in 71 plants under open-pollinated conditions. Apomictic embryo sac development was observed in double-cross progenies when crosses involved a pearl millet x P. squamulatum apomictic hybrid as pollen parent. These new double-cross hybrids may serve as bridging hybrids to transfer genes controlling apomixis and other plant characteristics from the wild Pennisetum species to pearl millet.     

Cytological studies on diploid and polyploid taxa of the genus Pennisetum Rich

The cytology of thirteen taxa and two hybrids in the genus Pennisetum indicated the distribution of the taxa among the four basic chromosome numbers 5, 7, 8 and 9. The diploid nature of P. ramosum and P. typhoides and the genomic allotetraploid status of P. purpureum was further confirmed. P. massaicum (2n=32), P. orientale (2n=36) and P. subangustum (2n=36) suggested probable autotetraploid nature and the three hexaploids (2n=54) investigated (P. polystachyon, P. longistylis and P. squamulatum) revealed allohexaploid constitution. The natural triploid, P. ruppellii (2n=27) and pentaploid P. villosum (2n=45) were found to be apomicts and they were allotriploid and allopentaploid respectively. The tetraploid-hexaploid complex of P. pedicellatum showed them to be cytotypes only. The cytogenetical behaviour of the hybrid, P. typhoides x P. purpureum with 2n=21 and the trispecies hybrid (P. typhoides x P. purpureum) x P. squamulatum with 2n=48 brought out the homology within the genomes of x=7 and x=9 and also between the genomes with x=7 and x=9. The significance of the inter- and intragenomic chromosome pairing had been brought out from the interspecific hybrids and the natural allotriploid and allopentaploid species having one genome in the haploid condition.The morphological sections of this genus did not correspond with the cytological groups. A high degree of evolutionary specialization was evident in species of the section Gymnothrix. A complete series of polyploids and high degree of heterogeneity from the morphological point of view was brought out in species with x=9.     

Genetic Control of Chromosome Pairing in Hexaploid Oats

THE common oat, Arena sativa, is a hexaploid (2n=6x=42) whose chromosomes form exclusively bivalents during meiosis. The bivalents are formed by the pairing of exact homologues resulting in disomic inheritance. Gene duplications and triplications were revealed by early inheritance studies¹ and homoeologous relationships of the three genomes were later supported by the good tolerance of deficiency for single and for pairs of chromosomes².     

Cytology and systematics in JapaneseArisaema (Araceae)

Chromosome numbers are determined from 37 populations attributed to 22 taxa of JapaneseArisaema. Of them, chromosome numbers ofA. limbatum var.conspicuum (2n=26),A. minus (2n=26),A. nambae (2n=28) andA. seppikoense (2n=26) are determined for the first time. New chromosome numbers, 2n=26, are reported forA. aequinoctiale, A. limbatum, A. stenophyllum, A. undulatifolium andA. yoshinagae. Three modes of basic chromosome numbers,x=14,x=13 andx=12, occur in JapaneseArisaema. Precise karyotypic comparisons of 20 taxa reveal that taxa withx=14 andx=13 share 26 major chromosome arms and have an obvious chromosomal relationship. One of two submeta-centric chromosomes inx=13 corresponds to two telo-centric chromosomes inx=14. InA. ternatipartitum with 2n=6x=72, ten out of 12 basic chromosomes are the most similar in size and arm ratio with larger ten chromosomes ofA. ringens among JapaneseArisaema examined. A basic chromosome number ofx=14 is the commonest in the genusArisaema and the remaining basic chromosome numbers,x=13 andx=12, seem to be derived through dysploidal reduction by translocating large segments of major arm of telo-centric chromosome onto other minor arm of telo-centric followed by loss of the remainings including a centromere, and by loss of two telo-centrics fromx=14, respectively. Some systematic problems of JapaneseArisaema are discussed based on new cytological data.Arisaema hatizyoense, A. minus andA. nambae are accepted as independent species.     

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     

Chromosome numerical mosaicism in some hybrids of theSolanum nigrum complex

The colchicine amphiploid (2n=72) of the hybridSolanum nodiflorum (2n=24)×S. villosum (2n=48) was crossed toS. memphiticum (2n=72) and toS. nigrum (2n=72). In the resulting F₁ hybrids chromosome numerical mosaicism was observed in the anthers of some plants. It was found that chromosome mosaicism did not affect chromosome pairing in normal cells. Disharmonious gene combination seems to be responsible for premeiotic somatic instability which results in chromosome mosaicism observed during meiosis in pollen mother cells. Gene or genes for fertility are transmitted independent of those for chromosome stability. It is suggested thatS. nigrum andS. memphiticum possess in common a gene complex, which when combined withS. nodiflorum andS. villosum genomes will induce somatic instability.     

Genesis of bivalent pairing in hexaploid clump Verbena

Both 6x Verbena aubletia (n=15) and 2x V. tenuisecta (n=5) form bivalents during meiosis, however, their 4x F₁ hybrid (V. aubletia × V. tenuisecta) shows almost complete homoeologous pairing involving on average 19.74 out of its 20 chromosomes. In 10% cells there are 4IV+2II indicating that essentially there may be 4 homoeologous sets of 5 chromosomes each in the F₁ hybrid. Evidently, V. aubletia is segmental allo-hexaploid involving 3 homoeologous genomes (A₁A₁ A₂A₂ A₃A₃). Whether its cytologically diploid behaviour is the result of a multivalent suppressor system or due to an acute property of preferential pairing, cannot be answered with certainty. In either case intergenomal homoeologies are totally suppressed resulting in bivalent pairing, meiotic isolation of the 3 genomes and institution of normal fertility.     

An ordered arrangement of bivalents at first meiotic metaphase of wheat

The spatial relationships between different types of marked pairs of bivalents or trivalents were studied at first meiotic metaphase of tetraploid wheat (2n=4x=28; genome AABB) by tallying the number of intervening bivalents between the marked pair. The study was carried out only in pollen mother cells with circularly arranged metaphases. Telocentric bivalents representing the two different arms of one chromosome were found very close to each other. Nonrelated trivalents of the same genome were significantly closer to each other than nonrelated trivalents of different genomes. Homoeologous trivalents tended to lie relatively close to one another. This pattern of chromosomal arrangement agrees well with previous findings in hexaploid wheat, both in mitotic and meiotic cells. It is concluded that in polyploid wheat the chromosomes of each genome lie relatively close to each other and are spatially separated from those of the other genomes.     

A SYSTEMATIC STUDY OF PENNISETUM SECT. PENNISETUM (GRAMINEAE)

Pennisetum sect. Pennisetum includes two reproductively isolated species. Pennisetum purpureum Schumach. is a tetraploid (2n = 28) perennial species which occurs throughout the wet tropics of the world. Pennisetum americanum (L.) Leeke is a diploid (2n = 14) annual species, native to the semi-arid tropics of Africa and India, and contains three morphologically diverse subspecies. Subspecies americanum includes the wide array of cultivated pearl millets. Subspecies monodii from the Sahel of West Africa is identified as the wild progenitor of pearl millet. Subspecies stenostachyum is morphologically intermediate between subsp. americanum and monodii and includes the mimetic weeds often associated with the cultivation of pearl millet.     

Intergeneric hybridization and C-banding patterns inHordelymus (Triticeae,Poaceae)

Crosses ofHordelymus europaeus (2n = 4x = 28) with four genera in theTriticeae were attempted. Adult hybrids were obtained in combinations withHordeum bogdanii (2x),Hordeum depressum (4x), andSecale cereale (2x). The meiotic pairing was very low in the hybrids withH. bogdanii andSecale cereale (0.12 and 0.30 chiasmata/cell, respectively), whereas high pairing (9.90 chiasmata/cell) was found in hybrids withH. depressum due to autosyndetic pairing ofH. depressum chromosomes. The chromosome complement ofHordelymus europaeus comprised 16 metacentrics, 8 submetacentrics, and 4 SAT-chromosomes. The Giemsa C-banding patterns of the chromosomes were characterized by small to minute bands at no preferential positions. It is hypothesized thatHordelymus europaeus may genomically be closest related toTaeniatherum andPsathyrostachys spp.     

Cytogenetic and genomic relationships ofThinopyrum elongatum with twoPsathyrostachys species and withLeymus secalinus (Poaceae)

Intergeneric hybridizations were made betweenT. elongatum, and twoPsathyrostachys and fiveLeymus species. The seed set obtained onT. elongatum ×Leymus hybrids ranged from 5.65% to 20.00%, depending onLeymus species. The seed set obtained onT. elongatum ×Psathyrostachys hybrids ranged from 16.07% to 19.70%. Meiotic pairing at metaphase-I in JN diploid hybrids ofT. elongatum ×Psathyrostachys species revealed a very low level homology between the basic J and N genomes, and further demonstrated that the two genomes are quite diverged. Chromosome pairing in theT. elongatum ×Leymus secalinus hybrid averaged 15.19 univalents + 2.62 rod bivalents + 0.26 ring bivalents + 0.02 trivalents, suggesting that the partial J^e chromosomes ofT. elongatum has homology withLeymus secalinus genomes.L. secalinus might have 3–4 chromosomes originating from J^e genome.     

Polyploidy and Evolution in Wild and CultivatedDahliaSpecies

Observations on the chromosomes of nine species ofDahliaCav.(Asteraceae, Heliantheae—Coreopsidinae) show that somehave 2n=32, others 2n=64, with a third group having both chromosomenumbers in the same taxon. Karyotype investigations showed thatthe chromosomes can be divided into groups of 14 metacentricsplus two submetacentrics per set of 16 chromosomes.In situhybridizationusing an rRNA gene probe indicated that the 2n=32 species haveeight hybridization sites whilst the 2n=64 species have 16 sites.Silver nitrate staining of these regions showed that not allof these nucleolar organizers are active. Meiotic analysis atmetaphase I and pachytene, by synaptonemal complex spreading,shows that the 2n=32 species have exclusive bivalent formationwhereas the 2n=64 species have small numbers of univalents plusquadrivalents in addition to bivalents. This study proposesthatDahliaspecies with 2n=32 are allotetraploids whereas thosespecies and chromosome races with 2n=64 are their autopolyploidderivatives. We suggest that a bivalent-promoting mechanismin the 2n=32 species may account for their meiotic behaviouras their component genomes appear so similar, and that thismechanism is also responsible for the low number of quadrivalentsin the 2n=64 taxa.Copyright 1998 Annals of Botany Comapny Chromosome pairing,Dahlia, in situhybridization, karyotype analysis, polyploidy, synaptonemal complex analysis     

Wheat Neocentromeres Found in F1 Triticale × Tritordeum Hybrids (AABBRH<Superscript>ch</Superscript>) After 5-Azacytidine Treatment

The DNA hypomethylation effect of 5-azacytine (5-AC; a cytosine analog) is widely known. This agent has been used for rRNA gene expression studies of Triticeae amphiploids and hybrids regarding rye rRNA genes suppression caused by the wheat nucleolar dominance phenomenon. However, this situation is reverted by 5-AC treatment which activates rye rRNA gene expression as it has been intensively observed in triticale. For nucleolar dominance studies, we produced F1 multigeneric hybrids (AABBRH^ch; 2n = 6x = 42) from crosses between the triticale cultivar ‘Corgo’ (AABBRR; 2n = 6x = 42) and the tritordeum cultivars HT9 and HT31 (AABBH^chH^ch; 2n = 6x = 42). The hybrid seeds were germinated in a low concentration of 5-AC (treatment) and in distilled water (nontreated control plants). Silver nitrate staining performed in one 5-AC-treated F1 hybrid revealed a reduced number of interphase cells with seven nucleoli, metaphases with eight Ag-NORs, and neocentromeres in the long arm of three wheat chromosomes. Nontreated hybrids presented six Ag-NORs per mitotic metaphase cell and a maximum of six nucleoli per interphase because of the 1R Ag-NOR suppression. No neocentromere was found in the control F1 hybrid plants. Both treated and nontreated seedlings were subsequently evaluated by fluorescent in situ hybridization performed with genomic and repetitive DNA probes to identify H^ch and rye genomes, to confirm Ag-NORs location, and to detect inactive rDNA loci. DAPI counterstaining was also helpful for the detection of neocentromeres in the long arm of three wheat chromosomes. This study allowed us to suggest that 5-AC treatment specifically induced wheat neocentromeres in the F1 multigeneric triticale × tritordeum hybrids.     

Chromosome studies in species and hybrids ofPetrorhagia sect.Kohlrauschia (Caryophyllaceae)

Cytogenetic investigations have been made in the fourPetrorhagia species and hybrids of the sectionKohlrauschia. The three diploid species show close similarities in chromosome number, size and morphology, with the exception ofP. velutina, where one pair of metacentric chromosomes is represented by a pair of telocentrics. Meiotic studies in hybrids indicate close genomic homology between the diploid species and also between the two floral forms ofP. prolifera. The tetraploidP. nanteuilii behaves as an allotetraploid forming only bivalents at meiosis and results suggest thatP. velutina andP. prolifera are the diploid progenitors of this species. Since meiosis in diploid and triploid hybrids results in extensive intergenomic pairing it is concluded that the natural tetraploid has a bivalent promoting mechanism that prevents pairing between the genomes of its diploid progenitors.     

Karyomorphology and relationships of Celtidaceae and Ulmaceae (Urticales)

Based on karyomorphological features, six (examined in this study) of nine genera of Celtidaceae are divided into three groups: 1)Celtis, Parasponia, Pteroceltis andTrema; 2)Aphananthe; 3)Gironniera, and six genera of Ulmaceae into two: 1)Holoptelea andPhyllostylon; 2)Hemiptelea, Planera, Ulmus andZelkova. The first four genera share the simple chromocenter type at the resting stage andx-10, with all chromosomes with submedian or median centromeres (frequency of chromosomes with subterminal or terminal centromeres 0%, although uncertain inTrema).Aphananthe hasx=13, but resembles the above four genera in other features.Gironniera is distinct from all other Celtidaceae in having the diffuse-complex chromocenter type andx=14, features which occur in Ulmaceae. InGironniera the frequency of chromosomes with subterminal or terminal centromeres is 43%, a proportion similar to those inHoloptelea (36%) andPhyllostylon (58%) of Ulmaceae. All six genera of Ulmaceae havex=14, yetHemiptelea, Planera, Ulmus andZelkova are distinct fromHoloptelea andPhyllostylon (with the simple chromocenter type) in having the diffuse-complex chromocenter type and in predominantly possessing chromosomes with subterminal or terminal centromeres (93%). Evidence from karyomorphology, as well as from other sources, suggests 1) thatAphananthe (x=13) is most distantly related to all genera withx=10 within Celtidaceae, 2) thatGironniera may have a key role for understanding evolutionary relationships between Celtidaceae and Ulmaceae, and 3) thatHoloptelea andPhyllostylon represent derivatives of a line that diverged early from a common ancestor or all Ulmaceae. On the basis of comparisons with other Urticales and the putative outgroups of the order, it is also suggested that the chromosome morphology of Ulmaceae represents the more derived state in Urticales.     

New occurrence of microchromosomes B in Moenkhausia sanctaefilomenae (Pisces, Characidae) from the Paraná River of Brazil: analysis of the synaptonemal complex

The Moenkhausia sanctaefilomenae specimens showed a karyotype consisting of 2n = 50 chromosomes with 12 metacentrics, 36 submetacentrics and two subtelocentrics. In addition to the basic karyotype, all the males specimens have cells ranging from zero to two B microchromosomes in mitotic metaphases. These chromosomes were not observed in the female specimens. C-band analysis showed a distribution pattern of characteristic heterochromatin with interstitial and centromeric blocks. However, the B chromosomes were faintly stained with C-banding and were not fluorescent with CMA₃ staining. The meiotic studies showed the formation of bivalents in metaphase I and in pachytene under an optical microscope. Through synaptonemal complex analysis with an electron microscope, the pachytene showed 25 bivalents completely paired and a small bivalent corresponding to the B chromosomes. In the same preparation, one of the B chromosomes was observed in a univalent form. On the basis of pairing behavior and morphology it is assumed that B chromosomes of M. sanctaefilomenae show homology between them and their evolutionary aspects are discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

Complete characterization of wheat–alien metaphase I pairing in interspecific hybrids between durum wheat (Triticum turgidum L.) and jointed goatgrass (Aegilops cylindrica Host)

The pattern of homoeologous metaphase I (MI) pairing has been fully characterized in durum wheat × Aegilops cylindrica hybrids (2n = 4x = 28, ABC^cD^c) by an in situ hybridization procedure that has permitted individual discrimination of every wheat and wild constituent genome. One of the three hybrid genotypes examined carried the ph1c mutation. In all cases, MI associations between chromosomes of both species represented around two-third of total. Main results from the analysis are as follows (a) the A genome chromosomes are involved in wheat–wild MI pairing more frequently than the B genome partners, irrespective of the alien genome considered; (b) both durum wheat genomes pair preferentially with the D^c genome of jointed goatgrass. These findings are discussed in relation to the potential of genetic transference between wheat crops and this weedy relative. It can also be highlighted that inactivation of Ph1 provoked a relatively higher promotion of MI associations involving B genome.