The variation and evolution of selected species of solanum section basarthrum

A study of greenhouse-grown and field populations ofSolarium caripense Humboldt & Bonpland ex Dunal,S. tabanoense Correll, andS. trachycarpum Bitter & Sodiro, all diploid species (n = 12) of high altitudes of southern Central America and northern South America, revealed great morphological variation. Polygonographs utilizing seven characters (number of leaflets, leaf index, length of pubescence, number of flowers, anther length, corolla index, and pollen diameter) showed a wide range of variation and led to recognition of six morphologically distinct groups. Hybridizations with greenhouse populations showed that five of the morphological groups are reproductively isolated as well. A complex pattern of genetic variation involving various degrees and combinations of low crossing success, low seed set, lowered F₁ pollen fertility, and nonreciprocal crossability was found. Examinations of meiotic figures from hybrids revealed no gross chromosomal structural differences. Evidence indicates that genetic differences, including gene-cytoplasm interactions, are significant isolating barriers. A key to the species studied plus appropriate taxonomic notes are provided.Solanum heiseri is described as new, andS. trachycarpum is placed inS. sect.Basarthrum seriesCaripensia.     

HYBRIDIZATION BETWEEN ANNUAL SPECIES OF PHLOX: POPULATION STRUCTURE

Phlox cuspidata (n = 7) and P. drummondii subsp. drummondii (n = 7) are closely related annuals which are indigenous to eastern and central Texas. The species typically occupy different ecological niches but may form contiguous or confluent populations in disturbed habitats and hybridize therein. On the basis of correlative interpretations of exomorphic, chromatographic and fertility information, hybridizing populations can be segregated into three distinct classes: (1) highly fertile plants with the morphological and phenolic attributes of P. drummondii; (2) highly fertile plants with the morphological and chemical attributes of P. cuspidata; (3) sterile plants with manifestly intermediate morphology and complementary chromatographic patterns. These data strongly suggest that hybridizing populations of P. drummondii and P. cuspidata are tritypic, being composed of “pure” or essentially “pure” parental species and a group of plants which has all of the attributes characteristic of an F₁ hybrid.     

Natural intersectional hybridization in perityle (Compositae)

A small number of supposed F₁ hybrids was discovered in an isolated sympatric population ofP. parryi (n = 17) andP. rupestris var.albiflora (n = 17) in the Chisos Mts. of Texas. Although the parental species are morphologically quite distinct, being assigned to separate taxonomic sections ofPerityle, their hybrids displayed relatively high fertility. Origin of the presumed natural hybrids was documented by the production of artificial hybrids betweenP. parryi andP. rupestris var.albiflora. Fertility data based on pollen stainability were accumulated for the available natural hybrids, and for artificial hybrids through F₃ and backcross generations. A brief discussion of biosystematic implications is included.     

Chromosome numbers in callirhoe (Malvaceae)

Forty-four chromosome counts from the North American genusCallirhoe are reported, including first-recorded counts ofC. alcaeoides, C. digitata, C. bushii, C. triangulata, andC. involucrata var.tenuissima. Information concerning chromosome numbers from present and previous reports is summarized and evaluated. Observations are presented on meiosis for the genus in general, for male sterile individuals of gynodioecious species, and for selected F₁ hybrids. Pollen fertility data are recorded for the latter. Chromosomally three species groups are recognized: an exinvolucellate, diploid species group withn=14; an involucellate, tetraploid and octoploid species pair,n=28 and 56; and an involucellate, diploid and tetraploid species group withn=15 and 30. Intrageneric relationships and those within the tribe Malveae subtribe Malvinae are discussed.     

ARTIFICIAL HYBRIDIZATION OF RUBBER-BEARING GUAYULE WITH COLD-TOLERANT PARTHENIUM LIGULATUM

ABSTRACT Caespitose and cold-tolerant plants of Parthenium ligulatum (Jones) Barneby (Asteraceae) from a native population in the Uinta Basin, Utah, were uprooted, potted, and transferred to a greenhouse in California. Approximately two years after transfer, the plants flowered and subsequently were crossed to diploid guayule (Parthenium argentatum Gray), the rubber-bearing species, native to the state of Durango, Mexico. Only female guayule × male P. ligulatum crosses produced F₁ hybrids. Only crosses involving guayule as female parent and F₁ plants as male parents produced backcross (BC,) plants. Hybrid plants were variable with respect to their growth habit, inflorescence, and leaf shape. Both parents and F₁ hybrids had 2n = 36 chromosomes. Unlike the parents, however, meiosis was irregular in the hybrids which showed a range of 0–5 and an average of 2.1 univalents at metaphase I. Hybrids averaged 0.87 laggards at anaphase I and 0.83 micronuclei at the tetrad stage. The crossability of guayule and P. ligulatum, the high degree of chromosome pairing of the F₁ hybrids, and the production of BC₁ plants indicate that the two species are related in spite of their distinct morphological and ecological differences. This study suggests that the cold-tolerance trait of P. ligulatum may be transferred to guayule through interspecific hybridization followed by backcrossing. The development of cold-tolerant guayule cultivars is expected to expand the areas of guayule production beyond that of the Chihuahuan desert and similar climates.     

Detecting hybridization in mixed populations ofRhinanthus minor andRhinanthus angustifolius

Rhinanthus minor andRhinanthus angustifolius are known to hybridize in mixed populations in nature. These hybridization events can have important evolutionary consequences. The development and use of species-specific RAPD and ISSR markers allowed the detection of hybrid individuals not always distinguishable with morphological characters. Two mixed populations of different ages were studied. In a young mixed 2-year-old population, both individuals of the two parental species and F₁ hybrids were found using genetic analysis, showing that hybridization occurred rapidly. Flower morphology of F₁ hybrids was too variable to distinguish all these hybrids from the parental species. This morphological variability of F₁ hybrids was also confirmed in artificial crosses in the greenhouse. In an old and no longer mixed 30-year-old population, onlyR. angustifolius plants and a few genetically introgressed individuals close toR. angustifolius were present. Genetic markers showed traces of past hybridization and introgression. Unidirectional introgression ofR. minor intoR. angustifolius with the complete disappearance ofR. minor from this population was observed.     

Autogamy and hybridization as evolutionary mechanisms in Panicum subgenus Dicanthelium (Gramineae)

A predominantly autogamous breeding system is described forP. occidentale Scribn.,P. pacificum Hitchc. & Chase, andP. thermale Boland. Five synthetic hybrids and three generations of progeny are also described for crosses among these three species. All parents haven = 9 chromosomes and normal meiosis. Hybrids exhibit various irregularities during meiosis, but nine bivalents are usually formed. One population ofP. occidentale and one ofP. thermale were found to have chromosomes with interchanges larger than those of other populations. The F₂ and F₃ generations had markedly increased fertility, as determined by pollen stainability. It is proposed that autogamy and hybridization are a common means of maintaining and stabilizing genetic variability in these grasses and that these means account for much of the taxonomic difficulty posed by the subgenus. The possibility of a low rate of introgression between populations is also proposed.     

Production of Polyploids and Unreduced Gametes in Lilium auratum × L. henryi Hybrid

Intergenomic F₁ hybrids between L. auratum x L. henryi and their BC₁ progeny were investigated through genomic in situ hybridization technique (GISH) to determine their potential value in lily breeding. We confirmed that F₁ intergenomic hybrids possessed a set of chromosomes (x=12) from both parents and that flowers of the F₁ auratum × henryi hybrid showed an intermediate morphological phenotype. Pollen size, viability and germination ability were measured through microscopic observations. F₁ intergenomic hybrids produced a relevant frequency of 2n-gametes, which were successfully used to perform crosses with Oriental hybrids, resulting in the triploid Oriental Auratum Henryi (OAuH) hybrid. Twenty BC₁ plants were generated by crossing between four different Oriental hybrid cultivars and F₁ AuH hybrids using an in vitro embryo rescue technique, after which the genome constitution and chromosome composition were analyzed by GISH. All plants were triploid, showing 12 from female parents (diploid Oriental hybrid) and 24 from male parents (diploid F₁ AuH hybrid). Overall, 16 out of 20 BC₁ progeny possessed recombinant chromosomes with 1-5 crossover sites per plant. Cytological analysis of 20 BC₁ plants by GISH verified that the occurrence of 2n pollen formation in all F₁ AuH hybrids was derived from the FDR (first division restitution) mechanism, in which the genome composition of all BC₁ plants possess 12 Oriental + 12 L. auratum + 12 L. henryi chromosomes. Allotriploids derived from the AuH hybrid were used as female for crossing with the diploid Oriental hybrid cultivar ''Sorbonne'' and considerable numbers of plants (0-6.5 plants per ovary) were only obtained when female OAuH (BC₁) triploids were used. Taken together, the results of this study indicate that production and analysis of F₁ AuH hybrids and their progeny through sexual polyploidization can be useful for efficient creation of important horticultural traits.     

CO(2) Exchange, Cytogenetics, and Leaf Anatomy of Hybrids between Photosynthetically Distinct Flaveria Species

Hybrids between the C₄-like species, Flaveria brownii, A. M. Powell and the C₃-C₄ intermediate species Flaveria linearis Lag., Flaveria floridana Johnston, and Flaveria oppositifolia (DC.) Rydb. exhibited bivalent chromosome pairing during meiosis and stainability of pollen was high, ranging from 51 to 95%. An F₂ population produced from an F. brownii × F. linearis F₁ hybrid, exhibited bivalent chromosome pairing and high pollen stainability indicating a high degree of fertility in the hybrid. Oxygen inhibition of apparent photosynthesis averaged 6.8% for F. brownii and 22.2% for the C₃-C₄ species (in two experiments), and F₁ hybrids exhibited inhibitions which were intermediate to their parents. Values of carbon dioxide compensation concentration determined at low irradiance were 4.0, 34.0, and 6.5 microliters per liter for F. brownii, F. linearis and their F₁ hybrid, respectively. The mean value at low irradiance for 33 F₁ plants was 6.8 microliters per liter, and individual values ranged only from 3.7 to 11.7 microliters per liter. Anatomical characteristics for the F₁ hybrid leaves were intermediate to those of the parents, and there was considerable variation among F₂ plants derived from F. brownii × F. linearis. In the F₂ population δ¹³C values ranged from −27‰ to −20‰. The expression of more C₄-like characteristics by the F₁ hybrids in this study and their apparent high fertility make them promising specimens for producing segregating populations for use in C₄ inheritance studies.     

VARIATION IN CHROMATOGRAPHIC PATTERNS IN THE TRAGOPOGON DUBIUS-PRATENSIS-PORRIFOLIUS COMPLEX (COMPOSITAE)

Natural populations of the diploid species, Tragopogon dubius, T. pratensis and T. porrifolius, the F₁ hybrids of these species, and the two amphidiploids, T. mirus and T. miscellus, were obtained from the same localities used for previous morphological and cytological studies of the evolutionary relationships of this complex. Inter- and intra-populational comparisons were made utilizing 2-dimensional chromatograms of these taxa. Members of the complex shared a group of 18 flavonoid-type pattern components, no one of which was completely species specific. Therefore, species patterns were expressed for a population rather than an individual with pattern components being expressed as a frequency. Patterns for F₁ hybrids and amphidiploids, expressed in the same manner, were relatable to the parental species. Diploid species and F₁ hybrid patterns were the most variable in areas of greatest sympatry, indicating considerable genetic interchange between species. The distribution of specific components in certain populations was interpreted as chemical introgression. Evidence was presented for separate origins of T. mirus in two localities. The results confirm previous interpretations of the evolutionary relationships in this complex, and when compared to those obtained for Baptisia, they implicate the different breeding systems as important factors in establishing the kind of variation observed.     

The variation and evolution of selected species of Solanum section Basarthrum (Solanaceae). II

Five taxa ofSolanum sect.Basarthrum were studied in an effort to clarify their taxonomic position and to determine the effective evolutionary mechanisms. Methods included an analysis of chromosome number and behavior, artificial hybridizations and a study of herbarium material. The data suggest thatS. canense andS. suaveolens are closely related and that 5.suaveolens may have been the progenitor ofS. canense. The only successful interspecific cross involving one of these two species was vigorous but highly sterile. Evidence from both morphology and crossing studies indicates a close relationship betweenS. basendopogon andS. caripense. Hybrids between these two species with relatively high fertility through the F₃ generation were secured. The status ofS. basendopogon f.obtusum remains a problem since there is but one collection of the typical form. The placement ofS. sanctae-marthae in sect.Basarthrum is considered problematic. Virtually none of 170 interspecific crosses with this species were successful. Seed size and the presence and size of a seed wing are proposed as useful morphological characters in sect.Basarthrum. There is apparently a correlation between short styles and self-compatibility. All species are diploid (n = 12) and no chromosomal or meiotic aberrations were noted in the species or hybrids. Most of the more than 1,000 interspecific crosses failed. Most of the hybrid fruits bore no seeds or seeds which did not germinate. The primary barriers separating species are considered to be strong prefertilization isolating mechanisms and ecogeographic factors.     

Morphological,chemical, and cytogenetic characters of F1 hybrids betweenParthenium argentatum (Guayule) andP. fruticosum var.fruticosum (Asteraceae) and their potential in rubber improvement

Interspecific hybrids betweenParthenium argentatum, the guayule rubber plant, andP. fruticosum var. fruticosum were evaluated for their potential rubber content and quality. Fifteen-mo-old field-grown plants ofP. fruticosum var.fruticosum measured four times more in height and spread than those ofP. argentatum, but contained less than 0.05% rubber of low mol wt.Parthenium argentatum showed 2% rubber content, with a mol wt of about one million. Resin contents varied little among parents or hybrids. The same age F₁ hybrids were intermediate in height and spread and had low rubber content, but showed presence of high mol wt rubber like the guayule parent. This indicates that high mol wt rubber is expressed over the low mol wt rubber in F₁ hybrids. Despite low rubber content but favorable biomass production, F₁ hybrids revealed irregular meiotic chromosome behavior and low pollen and seed germination. These results suggest that interspecific F₁ hybrids may be used in backcross programs to increase biomass and rubber content in guayule.     

A systematic study of the genus hulsea (Asteraceae)

A systematic treatment based on genetic, biochemical, and morphological studies is presented for the 11 recognized taxa ofHulsea. A uniform diploid complement of 38 chromosomes was found in the 37 populations examined, including all species and representing first reports forH. brevifolia, H. californica, andH. mexicana. Thirteen flavonoid compounds, based on the aglycones quercetin, apigenin, cyanidin, and the 4′-O-methyl ether of luteolin, were detected inHulsea. Patterns of the flavonoid compounds are discussed with respect to the systematic treatment. The results of 595 synthetic crosses indicate thatHulsea species are self-incompatible, relatively cross-compatible, and that their F₁ hybrids are relatively fertile. Heterozygosity for one or two translocations is reported for 11 interspecific and intersubspecific hybrid combinations.     

Coreopsis nuecensis (Compositae) and a related new species from southern Texas

An aneuploid chromosome series ofn = 6, 7, 8, 9, and 10 inCoreopsis nuecensis Heller was analyzed for morphological, distributional, and genetic correlations with the chromosome number classes. The results show that two sets of basic chromosome numbers occur within what has been treated asC. nuecensis: n = 6, 7 andn = 9, 10. Then = 7 class frequently carries a pair of B chromosomes, forming then = 8 class. The base chromosome numbers are correlated with some minor but consistent morphological differences, with distributional differences, and with strong sterility barriers in the F₁ hybrids. It is proposed that then = 9, 10 segment be recognized as a new species,Coreopsis nuecensoides.     

Reproductive isolation and hybridization in sympatric populations of three Dactylorhiza species (Orchidaceae) with different ploidy levels

Background and Aims

The potential for gene exchange between species with different ploidy levels has long been recognized, but only a few studies have tested this hypothesis in situ and most of them focused on not more than two co-occurring species. In this study, we examined hybridization patterns in two sites containing three species of the genus Dactylorhiza (diploid D. incarnata and D. fuchsii and their allotetraploid derivative D. praetermissa).

Methods

To compare the strength of reproductive barriers between diploid species, and between diploid and tetraploid species, crossing experiments were combined with morphometric and molecular analyses using amplified fragment length polymorphism markers, whereas flow cytometric analyses were used to verify the hybrid origin of putative hybrids.

Key Results

In both sites, extensive hybridization was observed, indicating that gene flow between species is possible within the investigated populations. Bayesian assignment analyses indicated that the majority of hybrids were F₁ hybrids, but in some cases triple hybrids (hybrids with three species as parents) were observed, suggesting secondary gene flow. Crossing experiments showed that only crosses between pure species yielded a high percentage of viable seeds. When hybrids were involved as either pollen-receptor or pollen-donor, almost no viable seeds were formed, indicating strong post-zygotic reproductive isolation and high sterility.

Conclusions

Strong post-mating reproductive barriers prevent local breakdown of species boundaries in Dactylorhiza despite frequent hybridization between parental species. However, the presence of triple hybrids indicates that in some cases hybridization may extend the F₁ generation.     

Chromosome evolution in the Gibasis linearis group (Commelinaceae)

Karyotypes, DNA amounts, and meiotic behaviour were examined in population samples of two closely related species, Gibasis venustula and G. heterophylla, and their F₁ hybrids. All samples were diploid (2n=12). DNA amount was similar in G. heterophylla, G. venustula ssp. robusta, and some populations of G. venustula ssp. venustula but in 13 samples of the latter, it showed as much as a 60% difference despite karyotypic uniformity. Although DNA levels were related to the altitude of the habitat, there was some variation within two morphological and ecological races of subspecies venustula. Analysis of hybrid meiosis and pollen fertility demonstrated the cytogenetical discreteness of populations, races, and subspecies. Populations and races were concluded still to be evolving following ecological isolation. The significance of quantitative and qualitative genome changes in evolution and their bearing on the classification of these species is discussed.     

Chromosome numbers and taxonomic notes for Mexican coreopsis,sections electra and pseudoagarista (compositae: heliantheae)

Notes on morphology and chromosome numbers are given for several species of MexicanCoreopsis, most of which were poorly known prior to recent collections.Coreopsis parvifolia (sectionElectra) is a large, shrubby, octoploid (2n=112) species apparently restricted to a small area in Puebla. It andC. cuneifolia (a diploid with 2n=28) appear to be closely related but differ by a number of morphological features in addition to ploidy level. A first chromosome report forC. pringlei (sectionPseudoagarista) shows this rare species to be diploid (2n=26), a fact which is in agreement with the base number of the section, i.e.,x=13. Additional collections of the very rareC. rudis andC. mcvaughii show them to be similar yet distinct species. Chromosome determinations forC. petrophila from Nayarit and Durango agree with previous counts for Jalisco populations of the species, i.e., 2n=26. Considerable morphological variation exists within this species but no subspecific entities are recognized.     

Development and cultivation of F2 hybrid between Undariopsis peterseniana and Undaria pinnatifida for abalone feed and commercial mariculture in Korea

The kelp Undariopsis peterseniana (Kjellman) Miyabe et Okamura is warm water tolerant and consequently there is currently considerable interest in developing commercial cultivation techniques for this species in Korea. Undaria is an important species for both the abalone industry and for commercial seaweed mariculture. In an attempt to extend the culture period of Undaria we bred and cultured hybrid kelp that is a cross between free-living gametophytes of U. peterseniana and Undariopsis pinnatifida. The morphological characteristics of the F₁ hybrid sporophytes were intermediate between those of the parent plants having shallow pinnated blades and forming both sorus and sporophyll. A F₂ generation was produced by isolating zoospores from sorus and sporophyll separately from a F₁ hybrid thallus. Using free-living gametophyte seeding and standard on-growing techniques, F₀ (female U. pinnatifida and male U. peterseniana), F₁, and F₂ gametophytes were cultured from December 2011 to May 2013. The morphological differences between the F₁ and F₂ generations were assessed. The F₂ hybrids were found to have longer pinnate blades and narrower midribs than the F₁ hybrid and only formed sporophylls. Growth and morphology of F₂ hybrids originating from the sporophyll or sorus of the F₁ hybrids were not morphologically different from each other. Both of the F₂ hybrids exhibited late maturation, with the early stages of sporophylls appearing in April. This late maturation of F₂ hybrids is beneficial in the development of hybrid strains that extend the period of availability of Undaria for abalone feed and cultivation in Korea.     

Variation among tripsacum populations of Mexico and Guatemala

Seven species of the genusTripsacum, one with a well marked subspecies, are indigenous to Mexico and Guatemala. Two diploids,T. zopilotense andT. maizar, are restricted to western and southwestern Mexico. The typical diploid form of a third species,T. latifolium, occurs in south central and eastern Guatemala; atypical tetraploid forms of this species are found in western Mexico. Four tetraploids,T. dactyloides, T. dactyloides ssp.hispidum, T. lanceolatum, andT. pilosum, have ranges extending from northwestern Mexico southeastward into Guatemala. The seventh species is the highly sterile tetraploid,T. laxum, of doubtful occurrence in the wild. It is readily propagated vegetatively and has been widely distributed in Latin America and the West Indies as a forage plant. Field studies of the range of variation in 80 Mexican and Guatemalan populations included an evaluation of 16 definitive morphological characteristics from an average of approximately six individuals of each population. Seventy-three of the 80 populations were allopatric, and of these, 50 were classed as typical, 13 as atypical, and ten as intermediates. Thirty-nine of the 50 populations classed as typical were tetraploid, and 11 were diploid. Of the 23 atypical and intermediate populations, 17 were tetraploid and six were diploid. Fifty-eight tetraploid populations, which included typical, atypical, and intermediate forms ofT. dactyloides, T. dactyloides ssp.hispidum, T. lanceolatum, T. pilosum, andT. latifolium, comprised an inclusive intergrading series having different combinations of characteristics distinguishing the two very dissimilar diploids,T. mizar andT. zopilotense. This series was interpreted as support for the hypothesis of Randolph & Hernández (1950), that the tetraploid populations originated as alloploid derivatives of these or closely related diploid species. The occurrence in some of the tetraploid populations of characteristics not seen in either of the putative parental species was noted, and the possibility that other taxa also might have been involved in their origin is under investigation. In addition to the 73 allopatric populations included in this survey there were seven sympatric and mixed associations of diploid and tetraploid taxa. Among these were hybrids with various combinations of the characteristics differentiating taxa of the associated populations. These hybrids effectively obscured differences between taxa, thus creating a syngameon complex from which genetic recombinants were observed to have spread into recently disturbed habitats of neighboring areas. From such direct evidence, and the existing interrelationships among allopatric and sympatric populations, it was concluded that the origin of the Mexican and Guatemalan tetraploidTripsacum species and various atypical and intermediate variants, has involved, and is continuing to involve, alloploid recombinants of diploid species and syngameon complexes, of which those encountered in this survey are examples.     

Unidirectional hybridization and reproductive barriers between two heterostylous primrose species in north-west Yunnan,China

Background and Aims

Heteromorphy in flowers has a profound effect on breeding patterns within a species, but little is known about how it affects reproductive barriers between species. The heterostylous genus Primula is very diverse in the Himalaya region, but hybrids there have been little researched. This study examines in detail a natural hybrid zone between P. beesiana and P. bulleyana.

Methods

Chloroplast sequencing, AFLP (amplified fragment length polymorphism) markers and morphological comparisons were employed to characterize putative hybrids in the field, using synthetic F₁s from hand pollination as controls. Pollinator visits to parent species and hybrids were observed in the field. Hand pollinations were conducted to compare pollen tube growth, seed production and seed viability for crosses involving different morphs, species and directions of crossing.

Key Results

Molecular data revealed all hybrid derivatives examined to be backcrosses of first or later generations towards P. bulleyana: all had the chloroplast DNA (cpDNA) of this species. Some individuals had morphological traits suggesting they were hybrids, but they were genetically similar to P. bulleyana; they might have been advanced generation backcrosses. Viable F₁s could not be produced with P. bulleyana pollen on P. beesiana females, irrespective of the flower morphs used. Within-morph crosses for each species had very low (<10 %) seed viability, whereas crosses between pin P. bulleyana (female) and pin P. beesiana had a higher seed viability of 30 %. Thus genetic incompatibility mechanisms back up mechanical barriers to within-morph crosses in each species, but are not the same between the two species. The two species share their main pollinators, and pollinators were observed to fly between P. bulleyana and hybrids, suggesting that pollinator behaviour may not be an important isolating factor.

Conclusions

Hybridization is strongly asymmetric, with P. bulleyana the only possible mother and all detected hybrids being backcrosses in this direction. Partial ecological isolation and inhibition of heterospecific pollen, and possibly complete barriers to F₁ formation on P. beesiana, may be enough to make F₁ formation very rare in these species. Therefore, with no F₁ detected, this hybrid zone may have a finite life span as successive generations become more similar to P. bulleyana.