Nucleolar number variation in Hordeum species,their haploids and interspecific hybrids

Nucleolar number variation has been investigated in root tip cells by cytologically determining the number of nucleoli per cell in autoploids and alloploids of Hordeum species, their haploids and interspecific hybrids. The nucleolus organisers in autoploid types of H. vulgare or H. bulbosum did not show any alterations irrespective of the ploidy level. The nucleolar number variation in these species results from a definite pattern of fusion and the maximum number of nucleoli per nucleus corresponds to the number of secondary constrictions. Nucleolus formation in alloploids and interspecific hybrids is impaired on some of the NORs, suggesting differential amphiplasty or nucleolar dominance. A comparison of nucleolar formation in the alloploid species (brachyantherum, arizonicum, procerum and parodii), their haploids, and the interspecific hybrids revealed different degrees of variation from the expected mean and maximum numbers of nucleoli. While the deviations in hybrids between alloploids (H. arizonicum and H. brachyantherum or H. procerum and H. brachyantherum) are marginal, nucleolar dominance is more pronounced in hybrids involving H. vulgare or H. bulbosum as one of the parents and is invariably associated with the disappearance of the secondary constriction(s) from the NOR(s) contributed by one of the parents, and the number of nucleoli is appropriately reduced.     

CYTOGENETIC ANALYSES OF AUTOPOLYPLOIDS: MODELS AND METHODS FOR TRIPLOIDS TO OCTOPLOIDS

Methods are presented for determining the frequencies and numbers of various meiotic configurations expected in autopolyploids. This allows one to test polyploids of unknown origin for agreement with expected meiotic configurations. Rejection of the autoploid hypothesis may indicate the presence of Ph-like genes or some type of alloploid. The models consider mean chiasma frequencies of 2, 3, and 4 per bivalent for triploids and tetraploids and 2 per bivalent for pentaploids, hexaploids, heptaploids, and octoploids. Literature data for a known autotriploid, autotetraploids, allotetraploids, and allopentaploids were tested against expectations of the models. There was generally good agreement between number of observed autoploid meiotic configuration and those expected in the models.     

Transfer of Ph I genes promoting homoeologous pairing from Triticum speltoides to common wheat

Diploid-like chromosome pairing in polyploid wheat is controlled by several Ph (pairing homoeologous) genes with major and minor effects. Homoeologous pairing occurs in either the absence of these genes or their inhibition by genes from other species (Ph ^I genes). We transferred Ph ^I genes from Triticum speltoides (syn Aegilops speltoides) to T. aestivum, and on the basis of further analysis it appears that two duplicate and independent Ph ^I genes were transferred. Since Ph ^I genes are epistatic to the Ph genes of wheat, homoeologous pairing between the wheat and alien chromosomes occurs in the F₁ hybrids. Using the Ph ^I gene stock, we could demonstrate homoeologous pairing between the wheat and Haynaldia villosa chromosomes. Since homoeologous pairing occurs in F₁ hybrids and no cytogenetic manipulation is needed, the Ph ^I gene stock may be a versatile tool for effecting rapid and efficient alien genetic transfers to wheat.Contribution no. 93-435-J from the Kansas Agricultural Experiment Station, Kansas State University, Manhattan, KS 66506-5502, USA     

Evidence for multiple, autoploid origins of agamospermous populations in Eupatorium sessilifolium (Asteraceae)

Comparative analyses were made of agamospermous populations of Eupatorium sessilifolium, which have previously been documented to be polyploid, to determine whether they are alloploid or autoploid in origin and to assess the possibility that they have arisen more than once. There was no variability in ITS sequences among seven agamospermous and eight sexual diploid populations of E. sessilifolium, which is consistent with morphological observations in suggesting that the agamospermous populations were autoploids. The ITS sequence characteristic of E. sessilifolium differs from all other North American species by a minimum of 15 changes, and heterogeneity or polymorphism would be expected if the agamospermous populations were alloploids. Analysis of the chloroplast-based trnC-psbM spacer region showed variability among both sexual diploid and agamospermous populations of E. sessilifolium, which suggested that the agamospermous populations stem from multiple origins. Analysis of ISSR data revealed considerable intraspecific variability within E. sessilifolium, and the distribution of variability, with agamospermous populations showing variability from one another, added further evidence for multiple origins of agamospermous populations. The results in conjunction with distributional evidence that the sexual diploid populations of E. sessilifolium are geographically restricted and uncommon suggest that monitoring of populations might be warranted to evaluate whether measures are needed to enhance their continued survival.     

C-banding in 6x-Triticale xSecale cereale L. hybrid cytogenetics

Summary The meiotic behaviour of F₁ hybrids of hexaploid Triticale that differed in their genotypic or chromosomic constitution, and diploid rye, was investigated. Meiotic analysis were done by Feulgen and C-banding staining methods. A differential desynaptic effect in the hybrids was detected and explained in terms of genetic differences in pairing regulators. The high homoeologous pairing (A-B wheat chromosomes and wheat-rye chromosomes) observed in the hybrids can be explained in terms of an inhibition of the effect of a single dose of thePh allele of the 5B chromosome produced by two doses of the 5R chromosome. The higher homoeologous pairing detected in the hybrid 188 x Canaleja could be the overall result of the balance between thePh diploidizing system (1 dose), the pairing promoter of the 5R chromosome (2 doses) and that of the 3D chromosome (1 dose coming from the parental line Triticale with the substitution 3R by 3D).     

CHROMOSOME PAIRING IN NATURAL FIRST AND ADVANCED GENERATION LIATRIS HYBRIDS

Chromosome pairing was analyzed in natural F₁ and advanced generation hybrids of crosses between Liatris aspera and L. spicata, and L. aspera and L. cylindracea. The status of the hybrids was determined on the basis of morphological and chromatographic criteria. The F₁ hybrids of L. aspera × L. spicata contained pairing irregularities in 30% of their PMC's as compared to 10% in the hybrid segregates. Notably, translocation figures appeared in 7% of the F₁ PMC's as compared to 0.6% in the segregate PMC's. The F₁ hybrids of L. aspera × L. cylindracea contained irregularities in 19% of their PMC's as compared to 7% in the segregates. Translocation figures occurred in 6% of the F₁ PMC's, but only in 0.7% of the segregate PMC's. The incidence of pairing and level of structural heterozygosity was far lower in the hybrid segregates than was anticipated on the basis of F₁ pairing relationships. Thus it would appear that there is selection for structural homozygosity in the hybrid segregates.     

New hybrids between Agropyron and wheat

Summary Intergeneric hybrids of Triticum aestivum (2n=42,AABBDD) with Agropyron ciliare (2n= 28,SSYY), A. trachycaulum (2n=28,SSHH), A. yezoense (2n=28,SSYY) and A. scirpeum (2n=28) are reported for the first time. F₁ hybrids of T. aestivum were also produced with A. intermedium (2n=42,E₁E₁E₂E₂Z₁Z₁) and A. junceum (2n=14,JuJu). All wheat-Agropyron hybrids were obtained by embryo rescue technique. Cultivars and reciprocal crosses differed for seed set, seed development and F₁ plant production. The F₁ hybrids were sterile. Attempts to obtain amphiploids were unsuccessful. However, backcross derivatives were obtained with wheat as the recurrent parent.The level of chromosome pairing in A. trachycaulum x wheat, A. yezoense x wheat and wheat x A. junceum hybrids provided no evidence of homologous or homoeologous pairing. Mean pairing frequencies in A. ciliare x wheat, wheat x A. scirpeum and wheat x A. intermedium hybrids indicated homoeologous or autosyndetic pairing. Ph gene was more effective in regulating homoeologous pairing in A. yezoense x wheat hybrids than in A. ciliare x wheat hybrid. Chromosome pairing data of BC₁ derivatives indicated that either some of the wheat chromosomes were eliminated or Agropyron chromosomes caused reduced pairing of wheat homologues.Contribution No. 82-653-J, Department of Plant Pathology, Kansas State Agricultural Experiment Station, Manhattan, Kan, USA     

Recent hybrid speciation in Cardamine (Brassicaceae) – conversion of nuclear ribosomal ITS sequences in statu nascendi

 The very recent allopolyploid speciation of Cardamine insueta and Cardamine schulzii is well documented. We used this system for a further understanding of the evolution of the internal transcribed spacers (ITS) of nuclear ribosomal DNA in recently formed hybrids. The ITS sequencing of the two parent species and the alloploid offspring suggests a synopsis of the types of ITS evolution, reported so far in the literature. We detected homogenization to one parental ITS type with a very strong bias to the maternal sequence. Nevertheless, maintainance of both parental ITS sequences in the alloploids was also recorded. Our data suggest: (1) rapid genomic change in newly formed alloploids, and (2) a multiple origin of C. insueta and C. schulzii. Received: 24 August 1998 / Accepted: 8 September 1998     

HETEROSIS AND OUTBREEDING DEPRESSION IN INTERPOPULATION CROSSES SPANNING A WIDE RANGE OF DIVERGENCE

The intertidal copepod Tigriopus californicus was used as a model organism to look at effects of crossing distance on fitness and to investigate the genetic mechanisms responsible. Crosses were conducted between 12 pairs of populations spanning a broad range of both geographic distance (5 m to 2007 km) and genetic distance (0.2% to 22.3% sequence divergence for a 606-bp segment of the mitochondrial COI gene). For each pair of populations, three fitness components (hatching number, survivorship number, and metamorphosis number) were measured in up to 16 cohorts including parentals, reciprocal F₁, F₂, F₃, and first-generation backcross hybrids. Comparisons of each set of cohorts allowed estimation of within- and between-locus gene interaction. Relative to parentals, F₁ hybrids showed a trend toward increased fitness, with no correspondence with population divergence, and a decrease in variance, which in some cases correlated with population divergence. In sharp contrast, F₂ hybrids had a decrease in fitness and an increase in variance that both corresponded to population divergence. Genetic interpretation of these patterns suggests that both the beneficial effects of dominance and the detrimental effects of breaking up coadaptation are magnified by increasing evolutionary distance between populations. Because there is no recombination in T. californicus females, effects of recombination can be assessed by comparing F₁ hybrid males and females backcrossed to parentals. Both recombinant and nonrecombinant backcross hybrids showed a decline in fitness correlated with population divergence, indicating that segregation among chromosomes contributes to the breakup of coadaptation. Although there was no difference in mean fitness between the two backcross types, recombinational backcrosses showed greater variance for fitness than nonrecombinational backcrosses, suggesting that the breakup of parental gene ombinations within chromosomes has both beneficial and detrimental effects.     

Linear and non-linear regression analysis of genotype X environment interactions in pearl millet

Summary Regression analysis was computed on the grain yield of 15 single cross F₁ hybrids of pearl millet (Pennisetum typhoides (Burm.) S. & H.) evaluated in 20 environments at 19 sites in India to assess the nature of genotype X environment interactions. Linear, quadratic, cubic, twoand three-intersecting straight line models were examined for fit. The interactions of six hybrids viz. MH 110, MH 113, MH 114, MH 115, MH 120 and MBH 110 were explained by the linear regression model. The response of the remaining nine hybrids was largely non-linear. The two and three-intersecting straight line models fit better than the quadratic and cubic models and explained non-linearity of response. The two-intersecting straight line models fit for 6 hybrids MH 106, MH 107, MH 112, MH 116, MH 117 and BJ 104. The response of MH 109 was best explained by a three-intersecting straight line model, but there still existed a significant remainder variation. The truncation of environmental range by assuming moving division points was more efficient than the fixed division points for the segmental regression models. The stability of hybrid varieties on the best fitting model has been discussed.     

Electrophoretic relationships and phylogeny of Nordic polyploids inDraba (Brassicaceae)

104 populations of 15 Nordic species (2x–16x) of the taxonomically complex genusDraba were investigated using enzyme electrophoresis. The polyploids were genetic alloploids showing high levels of fixed heterozygosity and electrophoretic variation; the diploids were homozygous and genetically depauperate. Thus, the data suggest that alloploidy in arctic-alpineDraba serves as an escape from genetic depauperation caused by inbreeding at the diploid level. Although some populations probably have local alloploid origins, electrophoretic data indicate that several polyploids have migrated repeatedly into the Nordic area.Draba crassifolia (2n = 40) is probably octoploid based on x = 5. A hypothesis on the evolutionary history of the polyploids based on x = 8 is presented. Diploids contributing to numerous polyploid genomes and multiple origins of polyploids have seriously blurred taxonomic relationships. Relationships inferred from genetic data do not always correspond to those based on morphology; two morphologically very similar polyploids,D. alpina andD. oxycarpa, were, for example, genetically distant and probably represent independent lineages.     

HYBRID FITNESS IN THE LOUISIANA IRISES: ANALYSIS OF PARENTAL AND F1 PERFORMANCE

The assumption of hybrid inferiority is central to the two models most widely applied to the prediction of hybrid zone evolution. Both the tension zone and mosaic models assume that natural selection acts against hybrids regardless of the environment in which they occur. To test this assumption, we investigated components of fitness in Iris fulva, I. hexagona and their reciprocal F₁ hybrids under greenhouse conditions. The four cross types were compared on the basis of seed germination, vegetative and clonal growth, and sexual reproduction. In all cases, the hybrids performed as well as, or significantly better than, both of their parents. These results suggest that F₁ hybrids between I. fulva and I. hexagona are at least as fit as their parents. The results of this study are therefore inconsistent with the assumptions of both the tension zone and mosaic models of hybrid zone evolution.     

CHROMOSOMES AND CROSSING BEHAVIOR OF SOME SPECIES OF SANSEVIERIA

Menzel , Margaret Y. (Florida State U., Tallahassee), and James B. Pate . Chromosomes and crossing behavior of some species of Sansevieria. Amer. Jour. Bot. 47(3) : 230—238. Illus. 1960.–Approximately 20 species (28 clones) studied were diploids, tetraploids or hexaploids of the basic numbers x = 20; about 40% of the taxa were polyploid. All species had similar karyotypes, except for chromosome number. Five of 12 combinations of diploid species gave fertile F₁ hybrids; 4 studied cytologically showed 20 bivalents at metaphase I. Two triploid interspecific hybrids showed high trivalent frequencies. In contrast, multivalent formation in polyploid species was variable but rather low. Morphological relationships appeared reticulate among and between diploids and polyploids and did not coincide with barriers to crossing or to hybrid fertility. The following tentative hypothesis concerning relationships in the genus is proposed: Sansevieria is monophyletic and speciation has proceeded through genetic variation and hybridization at the diploid level and by allopolyploidy (of the segmental type) ; a low level of chromosome differentiation has accompanied speciation such that complete pairing occurs in diploid hybrids, but considerable preferential pairing occurs in allopolyploids. The occurrence of both polyploid and hybrid vigor, the fertility of hybrids between species differing greatly in morphology and physiology, and the high potential for vegetative propagation make the genus a favorable subject for breeding based on interspecific hybridization.     

Intergeneric hybridization between Hordeum and Hordelymus (Poaceae)

Intergeneric hybridization between the monotypic Eurasian genus Hordelymus and species of Hordeum was performed. Hybrids were obtained in nine combinations and the meiotic pairing was analyzed in four combinations (6 families). Hordelymus europaeus is probably an alloploid with two different genomes. The very low pairing in the hybrids with H. jubatum, H. brachyantherum , and H. pusillum indicates that these species do not have any genomes in common with Hordelymus. The hybrids with H. depressum have an intermediate pairing which suggests that there is a certain homoeology in one genome.     

ON THE METHODS OF EXPERIMENTAL TAXONOMY

By studying seedling progenies from individual plants it may be decided whether the material under investigation is allogamous, autogamous or apomictic. Chromosome counts disclose whether the material is cytologically stable or variable. If variation occurs it may be a question of polyploidy or aneuploidy. Aneuploidy may either be an occasional consequence of polyploidy or it may represent dysploidy and the formation of new basic numbers. Chromosome counts combined with measurements of chromosome size may reveal cases of pseudopolyploidy of the kind first observed in Luzula. If chromosome morphology is different between two taxa this indicates reproductive isolation. Different species usually differ with regard to their chromosome structure. Studies of meiosis are desirable in polyploids in order to distinguish between auto- and alloploidy. Cases of genic control of meiosis in polyploids of Phleum and Triticum demonstrate, however, that partial or complete homology between genomes does not always result in multivalent formation. Apparent alloploids may in reality be more or less autoploid. Incompatibility is not a reliable criterion of species differentiation. Strong or absolute barriers between diploids and autotetraploids may have purely quantitative causes. Hybrid sterility is not always a reliable measure of the degree of genetic difference between the parents, but may be caused by heterozygosity for chromosomal rearrangements. Cytological and experimental methods are, nevertheless, indispensable tools in plant taxonomy.     

DNA divergence as a criterion of a sweet pepper (Capsicum annuum L.) selection for heterosis

The identification of perspective parental lines for the creation of high-yield hybrids is the most labor-consuming stage of selection, because it needs extensive trials of combining ability. Based on evaluation of the genetic divergence of the parental lines, the prediction accuracy of F₁ hybrids performance was investigated. The value of the divergence was calculated using biometric and molecular methods, such as inter simple sequence repeats (ISSR) and random amplified polymorphic DNA (RAPD). Based on estimates of divergence, 10 lines were chosen for cyclic cross (scheme I) and testcross (scheme II). In most crosses, the F₁ hybrids were significantly superior to the parents in the main economically valuable traits. The level of heterosis was higher among hybrids of scheme I. Analysis of the relationship between parental divergence and F₁ performance showed that the hybrid productivity of scheme I was predetermined by ISSR divergence in 86%, and productivity was caused by RAPD divergence in 69%, whereas the F₁ yield of scheme II was not related to the value of genetic distances. Since the values of DNA divergence were closely associated both with midparent level and F₁ performance, we assumed that part of the polymorphic DNA fragments of the parental lines of scheme I is related to heterotic loci (HTL), which may be considered potential key markers for the heterotic selection of the sweet pepper.     

AUTOTETRAPLOIDY IN HAPLOPAPPUS SPINULOSUS HYBRIDS: EVIDENCE FROM NATURAL AND SYNTHETIC TETRAPLOIDS

Haplopappus spinulosus (Asteraceae) is a herbaceous, perennial weed common throughout the western Great Plains of North America and includes both diploid and tetraploid populations. A number of populations in southeastern Colorado were analyzed cytogenetically and morphologically for two reasons. First, initial observations during a routine population survey showed they were morphologically intermediate between the diploid subspecies glaberrimus and spinulosus, suggesting they might have arisen via hybridization. Second, cytological examination revealed that they were tetraploid. Because there was indication of hybrid origin, it was of interest to determine whether the populations were behaving as autopolyploids or segmental allopolyploids. The distinction between these two polyploid types is not easily made since both are likely to form multivalents at meiosis, but equations derived from a model proposed by Jackson and Hauber (1982) have made it possible to determine statistically whether a tetraploid individual is behaving meiotically as an autotetraploid. Meiotic configuration frequencies at diakinesis were determined for each tetraploid plant sampled, and observed frequencies were compared to those expected for an autotetraploid having the same maximum number of chiasmata per bivalent, chiasma frequency and chromosome number. In general, the meiotic behavior of the tetraploids was no different from that expected for autoploids. The initial hypothesis that the populations were derived from hybridization was tested by a detailed cytogenetic and morphological study of the presumed parental subspecies, F, hybrids, and natural putative hybrids. The evidence supports the hypothesis that the natural autotetraploids arose from the hybridization of ssp. glaberrimus and ssp. spinulosus.     

CHROMOSOME NUMBERS IN PLATYSCHKUHRIA RYDBERG (COMPOSITAE) AND THEIR SYSTEMATIC SIGNIFICANCE

Diploid (2n = 12_II), tetraploid, pentaploid, and hexaploid level chromosome numbers are reported for Platyschkuhria Rydb. (Compositae). First reports of accessory chromosomes in the genus are also presented. Known polyploidy in Platyschkuhria is thus far restricted to P. integrifolia (Gray) Rydb. var. desertorum (M. E. Jones) Ellison from south-central Utah. Cytogenetic analysis, using methods and models of Jackson and Hauber (1982), suggests alloploid origins for the tetraploid levels. Diploid and polyploid populations correlate strongly with distinctively different edaphic situations, yet no consistent morphological features were identified that could justify taxonomic recognition of the polyploids.     

High divergence of reproductive tract proteins and their association with postzygotic reproductive isolation in Drosophila melanogaster and Drosophila virilis group species

The possible association between gonadal protein divergence and postzygotic reproductive isolation was investigated among species of the Drosophila melanogaster and D. virilis groups. Protein divergence was scored by high-resolution two-dimensional electrophoresis (2DE). Close to 500 protein spots from gonadal tissues (testis and ovary) and nongonadal tissues (malpighian tubules and brain) were analyzed and protein divergence was calculated based on presence vs absence. Both testis and ovary proteins showed higher divergence than nongonadal proteins, and also a highly significant positive correlation with postzygotic reproductive isolation but a weaker correlation with prezygotic reproductive isolation. Particularly, a positive and significant correlation was found between proteins expressed in the testis and postzygotic reproductive isolation among closely related species such as the within-phylad species in the D. virilis group. The high levels of male-reproductive-tract protein divergence between species might be associated with F₁ hybrid male sterility among closely related species. If so, a lower level of ovary protein divergence should be expected on the basis that F₁ female hybrids are fully fertile. However, this is not necessarily true if relatively few genes are responsible for the reproductive isolation observed between closely related species, as recent studies seem to suggest. We suggest that the faster rate of evolution of gonadal proteins in comparison to nongonadal proteins and the association of that rate with postzygotic reproductive isolation may be the result of episodic and/or sexual selection on male and female molecular traits. Correspondence to: A. Civetta