THE HYBRID ORIGIN OF ASTER ASCENDENS (ASTERACEAE)

Aster ascendens Lindl. is a rhizomatous perennial of the Great Basin region of the western United States, with a chromosome base number of x = 13. Populations in the southern and western parts of the range are mainly diploid (n = 13) and those in the north and east are mainly tetraploid (n = 26). This species is postulated to be an amphiploid which has resulted from hybridization between the western Cordilleran A. occidentalis (T. & G.) Hook, and the Great Plains species A. falcatus Lindl. Aster ascendens is intermediate between these two species in most morphological traits, but resembles A. occidentalis in ray color and general growth habit, and A. falcatus in the shape of the outer phyllaries. The karyotype of A. ascendens also exhibits a combination of the features found in A. occidentalis and A. falcatus karyotypes. Of the specimens collected for this study, 13% were putative backcrosses of A. ascendens to one of its parent taxa or to a closely related species; this suggests that there is potentially continuing genetic interchange between the species groups that gave rise to A. ascendens.     

A CYTOTAXOXOMIC SURVEY OF MELAMPODIUM (COMPOSITAE-HELIANTHEAE)

Turner , B. L.. and R. M. King . (U. Texas, Austin.) A cytotaxonomic survey of Melampodium (Compositae-Heliantheae). Amer. Jour. Bot. 49(3): 233–26. Illus. 1962.—Chromosome counts are reported for individuals from 89 populations of Melampodium representing 26 species The genus is multibasic with x = 9, 10, 11, 12, 16 and 23. Chromosome numbers on a base of x = 10 characterize the section Melampodium while basic numbers of x = 23, 16, 12, 11 and 9 occur in the section Zarabellia. Melampodium camphoratum (n = 16) differs from all other species examined in having relatively small meiotic chromosomes. Only 6 of the 23 species are polyploid or have polyploid races. Melampodium leucanthum and M. cinereum have both diploid and tetraploid populations; the latter occur without any apparent morphological or geographical correlation and are probably autoploid in origin. A survey of the basic chromosome numbers known for other genera of the subtribe Melampodinae (12 of 22 genera) is presented. and it is suggested that x = 10 is the most probable basic number of the genus and subtribe.     

CYTOGEOGRAPHICAL STUDIES IN THE GENUS ARNICA (COMPOSITAE: SENECIONEAE). I

A total of 134 chromosome counts representing 21 taxa of the genus Arnica are presented. Counts are published here for the first time for A. lonchophylla (n = 38) and A. nevadensis (n = 38). Ten previously unreported counts representing 9 taxa are also presented. The basic chromosome number of the genus is x = 19. With respect to chromosome number, the genus exhibits maximum diversity in the Rocky Mountains of Colorado and Wyoming. Arnica angustifolia, A. cordifolia and A. mollis are recognized as mature polyploid complexes, containing several wide ranging polyploid races and only a few highly restricted or scattered diploid races. Within the genus in general, diploids tend to be restricted to unglaciated areas while polyploids are much wider ranging, particularly in unglaciated areas.     

A review of chromosome cytology in Hyacinthaceae subfamily Ornithogaloideae (Albuca, Dipcadi, Ornithogalum and Pseudogaltonia) in sub-Saharan Africa

The chromosome cytology of Hyacinthaceae subfamily Ornithogaloideae is reviewed within the framework of a recent molecular-based classification, with particular emphasis on its center of diversity in sub-Saharan Africa. We also provide new chromosome counts for sections that are unknown or poorly known cytologically. Albuca subgen. Namibiogalum (9 spp.) probably has an ancestral base number of x = 10 but subgen. Albuca (± 70 spp), subgen. Monarchos (9 spp.) and subgen. Osmyne (36 spp.) have x = 9. The pattern in subgen. Urophyllon (3 spp.) is remarkable: although x = 6 is likely, the species in the section exhibit a range of 2n = 12, 10, 8, 6 and 4 (exclusive of polyploidy). All karyotypes have three large chromosome pairs and a variable number of small chromosomes. Pseudogaltonia (2 spp.) has x = 9 and Dipcadi (26 spp.) possibly x = 9 in series Uropetalum and x = 6 in series Dipcadi, which exhibits a pattern of descending dysploidy leading to n = 3 in D. marlothii. In Ornithogalum (± 130 spp.) chromosome numbers are known for only 24 of the ± 84 sub-Saharan species, mostly from subgen. Aspasia and subgen. Ornithogalum sect. Linaspasia, both of which have x = 6, and from subgen. Galtonia, which has x = 8. In contrast, x = 7 is basic for the Eurasian sects. Honorius and Melophis, and x = 18 seems likely for sect. Cathissa. Sect. Ornithogalum, the cytology of which we does not examine in detail, may have x = 9. Polyploidy is apparently rare in the sub-Saharan African ornithogaloids, in marked contrast to the high frequency of polyploidy among Eurasian species. In Albuca just 3 or possibly 4 sub-Saharan species (9% or 13% of those counted) are exclusively polyploid and 5 more have diploid and polyploid races; and in sub-Saharan Ornithogalum, only the tropical O. gracillimum is exclusively polyploid, and the western southern African O. hispidum has diploid and polyploid races.     

CHROMOSOMAL STUDIES OF SUBGENUS TRIDENTATAE OF ARTEMISIA: EVIDENCE FOR AUTOPOLYPLOIDY

The sagebrushes (subgenus Tridentatae of Artemisia—new combination presented in the text) are western North America's most widespread and populous shrub group. Chromosome counts from 120 populations confirm the base chromosome number at x = 9 with numerous 2n = 2x = 18 diploids and 2n = 4x = 36 tetraploids. Few higher polyploids are known, and aneuploidy and supernumerary chromosomes are rare. All 11 Tridentatae species are now known cytologically. All but the narrowly endemic A. argillosa are known from at least three locations: A. arbuscula (2n = 18, 36), A. bigelovii (2n = 18, 36), A. cana (2n = 18, 36), A. longiloba (2n = 18, 36), A. nova (2n = 18, 36), A. pygmaea (2n = 18, 36), A. rigida (2n = 18, 36), A. rothrockii (2n = 18, 36, 54, ca. 72), A. tridentata (2n = 18, 36, 54), and A. tripartita (2n = 18, 36). The chromosome number of A. argillosa, reported here for the first time, is 2n = 36. Chromosome numbers of eight subspecies also have been determined. The subgenus is characterized by autopolyploidy as indicated by morphologically indistinguishable 2x and 4x plants, a few mixed ploidy populations, consistent formation of IVs in 4x PMCs, a relatively uniform 2x karyotype, and a 4x karyotype, which is approximately twice the 2x one. Karyotypic differences, if they exist at all, are on a populational level rather than a systematic taxonomic level. The Tridentatae have apparently rapidly differentiated in situ in North America under the stimulus of recurring aridic cycles since late Tertiary or early Quaternary. They likely derive from more primitive circumboreal stock originating from the Eurasian homeland of Artemisia. The differentiation of myriad forms of Tridentatae was seemingly achieved through genic rather than genomic means. Karyotypic analysis supports a position within Tridentatae of A. rigida, A. bigelovii, and A. pygmaea.     

Decreased DNA content of birch (Betula) chromosomes at high ploidy as determined by cytophotometry

Relative amounts of DNA were determined on telophase nuclei by Feulgen cytophotometry for euploid taxa of birch (Betula) with somatic chromosome numbers of 28, 42, 56, 70, and 84. A direct correlation was found between observed DNA absorbance and chromosome number except for plants of B. papyrifera with 84 somatic chromosomes. The DNA density value for nuclei of the 84-chromosome plants fitted a 12.25 ratio instead of the expected 13.0 ratio. The DNA density value for these plants was calculated to be approximately equivalent to plants which would possess 63 somatic chromosomes. The average DNA value per chromosome was 2.73 for the 84-chromosome plants in contrast to 3.50 per chromosome in each of the lower euploids. Nuclear diameters of the 84-chromosome plants were directly related to chromosome number and not to DNA density value. The genomic number of Betula was considered to be x=7, rather than x=14, since a DNA value equivalent to 63 chromosomes is a multiple of 7 and not 14. Diploid birch species (2n=2x=28), therefore, would actually be tetraploids (2n=4x=28). The reduction in DNA content may be an adaptation for the establishment of higher ploidy in birches.     

THE CHROMOSOMES OF SPLNACIA OLERACEA

Ellis , J. R., and Jules Janick . (Purdue U., Lafayette, Ind.) The chromosomes of Spinaeia oleracea. Amer. Jour. Bot. 47(3) : 210—214. Illus. 1960.—The somatic chromosomes of S. oleracea are described and each has been associated with one of the 6 morphological trisomics derived from triploid-diploid crosses. Of these 6 primary trisomies, reflex had been shown by genetic studies to be trisomic for the chromosomes carrying the sex-determining factors. This chromosome is the longest of the somatic complement and has a sub-median centromere. No obvious heteromorphism of this chromosome pair was observed in staminate plants. Heteromorphism involving this chromosome pair has been reported recently in 2 varieties of cultivated spinach by Zoschke (1956) and Dressier (1958) and was earlier reported by Araratjan (1939) for the wild species, S. tetandra. However, their accounts differ markedly from each other and with the present results in respect to the morphology of this chromosome pair. This study suggests the existence of races which differ with respect to the morphology of the chromosome pair containing the X Y factors.     

CHROMOSOMES AND SYSTEMATICS OF ASPLENIUM SECT. HYMENASPLENIUM (ASPLENIACEAE)

Gametic and somatic chromosome numbers of eight Asian species belonging to Asplenium sect. Hymenasplenium were determined. Seven species were observed to have chromosome numbers based on x = 39, and one on x = 38. These chromosome numbers are exceptional in Asplenium which is well known to have chromosomes of n = 36 or multiples thereof. Decisions on the taxonomic status of the species of section Hymenasplenium were facilitated by cytological observations. Systematic recognition of this section is also supported by the peculiarity in the chromosome numbers, and evidence for the addition to the section of several other species is presented.     

A taxonomical re-evaluation of the Valais chromosome race of the common shrew<Emphasis Type="Italic">Sorex araneus</Emphasis> (Insectivora: Soricidae)

The common shrewSorex araneus Linnaeus, 1758 is subject to intense chromosomal polymorphism. About 65 chromosome races are presently known. One of these chromosome races (the Valais race) is karyologically, morphologically, biochemically, and genetically clearly distinct from all other chromosome races of the species. Recent studies of hybrid zones between the Valais race and other chromosome races in the Swiss and French Alps add further strong evidence for the specific taxonomic status of the Valais race. Chromosomes and diagnostic protein markers reveal sharp frequency clines and strong heterozygote deficits. In one hybrid zone, the maintenance of the strong genetic differentiation of the hybridizing taxa was confirmed by a study with autosomal microsatellites indicating minimal gene flow. A microsatellite marker on the Y-chromosome showed complete absence of male mediated gene flow suggesting hybrid male sterility. To clarify the taxonomic status of this taxon, additional analyses were conducted. A morphometric analysis of the mandible indicated the Valais race is morphologically as distinct from neighbouring chromosome races ofS. araneus as from other relatedSorex species. In a phylogeny based on complete mitochondrial DNA cytochromeb gene sequences, the Valais race clearly appears as the sister taxon to all other races ofS. araneus. Therefore, the chromosome race Valais ofS. araneus herein is elevated to specific status and the nameSorex antinorii Bonaparte, 1840 is applied.     

Chromosome evolution in the Laurales based on analyses of original and published data

We present a summary of currently available chromosome information for all seven families in the order Laurales on the basis of original and previously published data and discuss the evolution of chromosomes in this order. Based on a total of 53 genera for which chromosome data were available, basic chromosome numbers appear consistent within families: x = 11 (Calycanthaceae); x = 22 (Atherospermataceae and Siparunaceae); x = 19 (Monimiaceae); and x = 12 and 15 (Lauraceae). The Hernandiaceae have diverse numbers: x = 15 (Gyrocarpoideae) and x = 18 and 20 (Hernandioideae). Karyotype analyses showed that Hennecartia, Kibaropsis, and Matthaea (all Monimiaceae) contained two or three sets of four distinct chromosomes in 38 somatic chromosomes, suggesting that 2n = 38 was derived by aneuploid reduction from 2n = 40, a tetraploid of x = 10. In light of the overall framework of phylogenetic relationships in the Laurales, we show that x = 11 is an archaic base number in the order and is retained in the Calycanthaceae, which are sister to the remainder of the order. Polyploidization appears to have occurred from x = 11 to x = 22 in a common clade of the Siparunaceae, Atherospermataceae, and Gomortegaceae (although 2n = 42 in the Gomortegaceae), and aneuploid reduction from x = 11 to x = 10 occurred in a common clade of the Hernandiaceae, Lauraceae, and Monimiaceae. To understand chromosome evolution in the Lauraceae, however, more studies are needed of genera and species of Cryptocaryeae.     

Primulina melanofilamenta sp. nov. (Gesneriaceae) from Guangxi,China

A new species of Gesneriaceae, Primulina melanofilamenta Ying Liu & F. Wen from Guangxi, China, is illustrated and described. It is morphologically distinct from the most similar species P. eburnea and P. atroglandulosa by the color of the corolla, dark purplish black filaments, anthers backed with a beard of white hairs, the length of staminodes and the hairs of styles. The somatic chromosome number of P. melanofilamenta is also reported as 2n = 36.     

Karyotypic variation in the Australian Gekko Phyllodactylus marmoratus (Gray) (Gekkonidae: Reptilia)

The gekko Phyllodactylus marmoratus has at least three distinct chromosome races; 2n=36, 2n=36 ZZ/ZW and 2n=34. Specimens from these races are morphologically distinguishable, have a degree of habitat specialization and occur in a defined distribution. The 2n=36 race found in Eastern Australia is the presumed primordial type. The 2n=34 race occurs in Western Australia and is regarded as a fusion derivative. The 2n=36 ZZ/ZW race, which is only found on the Murray River system in Eastern Australia has a heteromorphic sex chromosome system present in the female. Giemsa banding suggests that this heteromorphism is the result of a pericentric inversion.     

CHROMOSOME RACES AND STRUCTURAL HETEROZYGOSITY IN CALYCADENIA CILIOSA GREENE (ASTERACEAE)

A biosystematic study of Calycadenia ciliosa resulted in the recognition of five homoploid (n = 6) chromosome races. The cytogenetic evidence indicates that each of these races is differentiated from its nearest relative by a single reciprocal chromosome translocation, although at one point in the evolutionary history a pericentric inversion may have been a concurrent cytological event. The data also show that the chromosome phylogeny includes two instances of redundant translocation. Mean pollen stainabilities of interracial hybrids range from 16–80%. In a survey of four natural populations 30–60% of the individuals sampled were found to be structurally heterozygous for reciprocal chromosome translocations. Pericentric inversion heterozygosity was also detected in one population. Another population contained morphologically indistinguishable individuals separated by as little as 120 m that were differentiated by a minimum of four chromosome translocations. These observations were compared with similar instances in other species in an effort to determine their significance in the process of plant evolution.     

THE HORDEUM VIOLACEUM COMPLEX OF IRAN

Fifty-seven Iranian collections of Hordeum violaceum Boiss. & Huet, a perennial forage grass, contained diploid (2n = 14), tetraploid (2n = 28), and hexaploid (2n = 42) chromosome races. All collections came from moderate to high elevations in the Alborz and Zagros mountains and adjacent plateau areas of Iran. Each chromosome race had a discrete distribution, and the hexaploids were the most widespread. The diploids were cytologically regular, except for a chromosome interchange that occurred in about half of the plants. The tetraploids and hexaploids behaved cytologically as autopolyploids. The hexaploids were taller, coarser and later-flowering than the diploids and tetraploids, and they had fewer but thicker culms and larger seeds. The tetraploids were the leafiest and most productive, making them the most desirable from an agronomic standpoint. All races were more or less self-sterile, a characteristic that sets H. violaceum apart from most other Hordeum species. The taxonomic status of H. violaceum and its closest relatives, H. turkestanicum Nevski and H. brevisubulatum Link, is uncertain because of close morphological similarities and the occurrence of chromosome races in each taxon.     

Production and characterization of arboreous and fertile <Emphasis Type="Italic">Solanum melongena</Emphasis> + <Emphasis Type="Italic">Solanum marginatum </Emphasis>somatic hybrid plants

In crop plants the shift from being annuals to perennials may allow future agricultural systems requiring less energy inputs. The practicability of this was tested for Solanum melongena. Leaf protoplasts of S. melongena (2n = 2x = 24) and one of the related arborescent species Solanum marginatum (2n = 2x = 24) were electrofused and fertile somatic hybrids with arborescent habit regenerated. The magnetic cell sorter (MACS) technique was used for the selection of heterokaryons. The hybrid nature of 18 regenerated plants was assessed on the banding patterns generated by inter-simple sequence repeat PCR. When taken to maturity in the greenhouse, hybrids grew more vigorously compared to the parental species. Their morphological traits were intermediate between those of S. melongena and S. marginatum. Hybrids flowered and produced an average of 85% stainable viable pollen and fertile fruits. The somatic hybrids were maintained in the greenhouse for more than 3 years and continued to produce flowers developing into two types of fruits with plentiful seeds. Fruits were either striated green containing non-germinable seeds or yellow with fully germinable seeds. Their S₁ progenies showed common features with S₀ hybrids, including fertility and arborescent habit. Cytologically, somatic hybrids exhibited the expected chromosome number of 2n = 4x = 48, while chromosome pairing during microsporogenesis was associated with a low frequency of intergenomic pairing. It is concluded that an arborescent perennial species has been obtained by somatic hybridization. The usefulness of this species per se or in eggplant breeding will depend not only on the transmission of the arborescent habit to cultivated eggplant varieties, but also on the variability that should be created from backcrossing the S. melongena + S. marginatum hybrids to S. melongena.     

Widespread Chromosome Variation in the Endangered Grassland Forb Rutidosis leptorrhynchoides F. Muell. (Asteraceae: Gnaphalieae)

Rutidosis leptorrhynchoides is an endangered plant endemic tosoutheastern Australia. Chromosome analysis of 19 of the 24known populations of the species has identified 17 differentchromosome variants or cytotypes. The most common cytotypesare a diploid and a tetraploid based on x = 11, and triploidand hexaploid plants with this basic number were also observed.Diploids, triploids and tetraploids based on a second basicnumber ofx = 13 were also seen. Plants with 2 n = 24 were shownto be hybrids between diploids with the two different basicnumbers. Meiotic chromosome pairing analysis of the plants with2n = 24 showed a maximum of two trivalents indicating the presenceof extra copies of one pair of large and one pair of small chromosomesin the 2 n = 26 plants. In addition, a number of different aneuploidsof the 2 n = 22 and 2 n = 44 races were found and many of thesealso showed structural chromosomal variation. The distributionof the two main chromosome races is disjunct with the tetraploidsconfined to southern Victoria. To avoid dysgenic effects, futurere-establishment efforts for this species should avoid mixingseed from different chromosome races. Copyright 2001 Annalsof Botany Company Aneuploidy, conservation genetics, karyotypes, meiosis, polyploidy     

A cytological study ofPelargonium sect.Eumorpha (Geraniaceae)

The chromosome numbers of seven species ofPelargonium sect.Eumorpha have been determined from material of known wild origin, and karyotypic comparisons have been made. Within the section there is variation in basic chromosome number (x = 4, 8, 9, 11), variation in chromosome size, and two species have polyploid races. The three species with chromosome numbers based on x = 11 have the smallest chromosomes (1.0–1.5 µm); chromosomes are larger (1.0–3.0 µm) in the other species.P. elongatum has the lowest chromosome number in the genus (2n = 8).P. alchemilloides is exceptional in that it has four cytotypes, 2n = 16, 18, 34 and 36, and the form with 2n = 36 has large chromosomes (2.0–5.0 µm). Evidence from a synthesized hybrid suggests thatP. alchemilloides with 2n = 16 may be of polyploid origin. The three species based on x = 11 appear to be more closely related to species from other sections ofPelargonium that have the same basic chromosome number and small chromosome size, rather than to other species of sect.Eumorpha.     

The cytology of Brachycome lineariloba

A comparison of the karyotypes of races D (2n=8), E (2n=10), B (2n=12) and C (2n=16) of B. lineariloba suggests that these races have in common a basic set of four chromosome pairs, and that the higher chromosome number races are related to race D by successive chromosome addition. — A study of meiosis in B × C and A₁ × B hybrids supports this contention and elucidates the homologies of the additional chromosomes. — Meiotic pairing in hybrids between A and C is very complex. At present it can only be stated that there are extensive interchromosomal homologies between the two races. — Two phyletic schemes of the relationships of the races are considered. The second, which is favoured, involves successive chromosome addition, with the quasidiploid race E (2n = 10) giving rise to race B by diploidisation of the univalent chromosomes. This scheme is supported by features of univalent behaviour in the various races and their hybrids. — The ecogeographic distribution pattern of the races shows replacement of D by E by B by C as the species extends into more arid and more harsh environments. This replacement is also associated with increasing vigour. — It seems most likely that the addition chromosomes are derived from a race A (2n=4) source since they are added always by twos, and each addition increases both vigour and drought tolerance. Race A is the most vigorous and one of the most drought tolerant of the five races.It is suggested that the evolution of the races can be related to the increasing aridity of the Late Pleistocene and Recent geological epochs.     

Exceedingly high chromosome number in Strasburgeriaceae, a monotypic family endemic to New Caledonia

We present the first report on the chromosome number of Strasburgeria robusta, which is confined to montane forests of New Caledonia and is the only known species in Strasburgeriaceae. The species has 2n = 500, which is an exceedingly high chromosome number in angiosperms. Within Crossosomatales, molecular evidence has indicated that S. robusta is sister to Ixerba brexioides, which is endemic to New Zealand and is the sole species in Ixerbaceae. Comparisons to the chromosome number of I. brexioides (2n = 50) support a close affinity between the two species because they share the base number x = 25. It is generally accepted that an increase in ploidy is associated with the origin of novel adaptations. A high level of polyploidy (20x with x = 25) may have allowed S. robusta to survive on a fragment of Gondwana by adapting to its ultrabasic substrate.     

SOMATIC EMBRYOGENESIS AND PLANT REGENERATION IN TISSUE CULTURES OF PANICUM MAXIMUM JACQ.

Callus tissue cultures were initiated from immature embryos, mature embryos and young inflorescences of Guinea grass (Panicum maximum Jacq.) on Murashige and Skoog's (MS) medium supplemented with 2.5–10 mg/l 2,4-dichlorophenoxyacetic acid (2,4-D). Calluses were transferred onto the same nutrient medium with 0.2 mg/l 2,4-D, or without 2,4-D. In callus cultures derived from immature embryos and young inflorescence segments, plantlets were produced via somatic embryogenesis after 3–5 wk. Young plants were successfully transplanted to pots and grown in the greenhouse. Plant development in callus obtained from mature embryos took place through the organization of shoot meristems. Regenerated plants were shown to have the normal tetraploid chromosome number of 2n = 4x = 32.