BIOSYSTEMATIC STUDIES IN THE BOUTELOUA CURTIPENDULA COMPLEX. III. POLLEN SIZE AS RELATED TO CHROMOSOME NUMBERS

Pollen size statistics are presented for 10 closely related species of Bouteloua and relationships between pollen size and chromosome numbers are presented for 13 populations of 5 species and 3 varieties. With 1 exception, all populations of all taxa conformed to a general pattern of pollen size dependent upon chromosome number. Chromosome numbers varied from 2n = 20 to 2n = ca. 103, with several independent aneuploid series. Statistical analyses were made of pollen size as related to chromosome number in the 3 varieties of B. curtipendula. These data showed that tetraploids (2n = 40) of var. tenuis had significantly greater pollen size and coefficient of variation than diploids (2n = 20) of the same variety. Similarly, aneuploids of var. curtipendula with 2n = 45 to 2n = 64 chromosomes had significantly larger and more variable pollen than tetraploids (2n = 40) of the same variety. Highly significant positive regression coefficients were obtained from analyses of chromosome numbers and mean pollen size, and chromosome numbers and coefficient of variation, for var. curtipendula. Regression coefficients for var. caespitosa populations with chromosome numbers over the hexaploid (2n = 60) level were not significant.     

BIOSYSTEMATIC STUDIES IN THE BOUTELOUA CURTIPENDULA COMPLEX. V. MEGASPOROGENESIS AND EMBRYO SAC DEVELOPMENT

Megasporogenesis and embryo sac development in the sexually reproducing taxa Bouteloua warnockii (2n = 22), B. media (2n = 20), B. uniflora Vasey var. uniflora (2n = 20), B. uniflora var. coahuilensis Gould and Kapadia (2n = 20), and B. curtipendula var. curlipendula (2n = 40) all were found to be of the Adoxa type, in which all 4 megaspores persist and divide once to form an 8-nucleate embryo sac. On the other hand, evidence indicated that plants of B. curtipendula var. caespitosa with high ancuploid chromosome numbers reproduce by pseudogamous fertilization of an aposporous embryo sac. In this taxon the megaspore mother cell did not go beyond the first anaphase of meiosis and the functional embryo sac developed from a nucellar cell. Although the 8-nucleate embryo sac was typical, a 3-nucleate embryo sac was observed to develop in some cases.     

ANEUSOMATY IN ANEUPLOID POPULATIONS OF CLAYTONIA VIRGINICA

Lewis Walter H. (Stephen F. Austin State Coll., Nacogdoches, Texas.) Aneusomaty in aneuploid populations of Claytonia virginica. Amer. Jour. Bot. 49(9): 918–928. Illus. 1962.—From 2 central east Texas populations of Claytonia virginica, 15 chromosome numbers, 2n = 14, 15, 16, 18, 25, 26, 27, 28, 29 30, 31, 32, 33, 36, and 58, were found among a sample of 181 plants. The most frequently encountered numbers were 2n = 14, 28, and 29. Among an additional 14 plants the pollen mother cells in the same bud differed from one another in chromosome number, as well as the pollen and premeiotic cells from the same plant. The chromosomes of the most unstable plant varied from 2n = 14–36. Numerous meiotic abnormalities, including inversions, dicentrics, bridges, fragments, non-disjunctions, univalents, and multivalents, were observed for the aneusomatic and trisomic plants. It is suggested that the origin of the aneusomatics is related to the numerical disparity of the gametic chromosomes composing them. Since the species is perennial in habit, thereby allowing the unstable plants to produce gametes with varying chromosome numbers year after year, it is further proposed that the wide range of aneuploid known for C. virginica resulted, at least in part from the presence of aneusomatic individuals.     

Cytological studies on meiotic configurations of intraspecific aneuploids ofCarex blepharicarpa (Cyperaceae) in Japan

Chromosome configurations at meiotic metaphase I ofCarex blepharicarpa were determined for 245 individuals collected from 11 localities in the Chugoku District of Japan. Nine intraspecific aneuploids, 2n=26–33 and 41, were found. The most common diploid number was 29, and found in 73 individuals. No clear geographical pattern was suggested by a distribution of these aneuploids. A consecutive series of chromosome numbers from 2n=26 to 32 was found in two populations from Okayama Prefecture. At meiotic metaphase I, univalents and multivalents were found, and one to three trivalents were observed in each aneuploid. Heteromorphic chain trivalents comprising large, medium and small chromosomes were prevalent. Homomorphic trivalents, which are thought to be originated from chromosome duplications caused by unreduced gametes, were found in only one individual with 2n=41. The high frequency of heteromorphic trivalents in this species indicates that most aneuploidy probably results from fission and/or fusion of chromosomes.     

Trisomics and aneuploids of ryegrass

Summary Primary trisomics (2 n + 1 = 15), double trisomics (2 n + 1 + 1 = 16) and aneuploids with 24 to 30 chromosomes, as well as a diploid and tetraploids, were found in the progeny of a hypertriploid (2 n = 22) plant of perennial ryegrass, Lolium perenne L. Trisomics and double trisomics differed in their mean chromosome association, chiasma number and spike morphology. A few aneuploids and tetraploids had reciprocal translocations. The diploid, primary trisomics and tetraploids were more fertile than the double trisomics and aneuploids. Most trisomics and aneuploids were probably produced through female transmission. One double trisomic had a high univalent number, a low chiasma number and loose chromosome coiling. Both the extra chromosomes carried secondary constrictions. The gene for desynapsis might be located on one of these chromosomes.     

CYTOGENETICS OF RUBUS. II. CYTOLOGICAL STUDIES OF THE VARIETIES ‘YOUNG,’ ‘BOYSEN,’ AND RELATED FORMS

Thompson , Maxine M. (U. California, Davis.) Cytogenetics of Rubus. II. Cytological studies of the varieties ‘Young,’ ‘Boysen’ and related forms. Amer. Jour. Bot. 48(8): 667–673. Illus. 1961.—Chromosome numbers are given for the trailing blackberry varieties, ‘Young’ (2n = 49), ‘Boysen’ (2n = 49), ‘Nectar’ (2n = 49) and for related forms which include the parents of ‘Young,’ ‘Phenomenal’ (2n = 42) and ‘Mayes’ (2n = 56), and 3 cytologically resynthesized ‘Young’ plants (2n = 49) as a basis for interpreting the postulated origin of ‘Young.’ Cytological evidence substantiated the conclusion that ‘Young’ is a hybrid between ‘Phenomenal’ and ‘Mayes.’ Contributions to the understanding of genomic relationships in Rubus are offered from detailed analyses of meiosis in ‘Phenomenal,’ ‘Mayes,’ ‘Young,’ and ‘Boysen.’ ‘Phenomenal’ and ‘Mayes’ both had a very regular meiosis. ‘Young,’ as well as ‘Boysen,’ showed a greater degree of chromosome association than either parent of ‘Young.’ Meiotic behavior in ‘Boysen’ presented a close parallel to that of ‘Young’ which, correlated with morphological similarities and the same 2n chromosome number, suggests a similar origin. The mode of reproduction in ‘Young’ and ‘Boysen’ was found to be sexual on the basis of morphological variation in the open-pollinated (selfed) progeny, the varying aneuploid somatic chromosome numbers in these progeny (2n = 32–54) and aneuploid chromosome numbers in hybrids having either variety as one parent. The productiveness of ‘Young’ and ‘Boysen’ in commercial plantings and their successful utilization in breeding programs indicate a high fertility regardless of their having an odd multiple of the basic number. It is concluded that the production of balanced euploid gametes is not necessarily a criterion of fertility, at least not at this high level of ploidy.     

Occurrence of aneuploid progenies from an asynaptic amphidiploid ofScilla scilloides (lindley) druce II. Mechanism of production of the various aneuploid progenies

The mechanism of production of the various aneuploid progenies was clarified in the asynaptic amphidiploid plants (2n=34+4f+2F, AABB) ofScilla scilloides. Its asynaptic nature and chromosomal stickiness lead to the unequal segregation at anaphase I (AI) in PMC's. The observed values in 18 segregation patterns, 17:17 to 0: 34, were different from the expected values estimated from random segregation of chromosomes. Nevertheless, the preferential transmission of special chromosomes among genomes A (x=8=a₁−a₈) and B (x=9=b₁−b₉) had not occurred. As the result of unequal segregation, the pollen grains with various chromosome numbers were observed. Almost all of the 200 pollen grains contained chromosome numbers more than 17 (range 8 to 34). The observed values of each chromosome number were roughly similar to the expected values of containing the complete set of genome A or B in the random distribution without preferential segregation of chromosomes at AI. The difference between the index of polien mitosis and the pollen fertility was significant in the Wilcoxon matched-pairs signed rank test and suggested the selection for some genomically unbalanced pollen grains during maturation. Consequently, viable pollen grains with various chromosome constitutions are a few (mean pollen fertility of 5.8%) but might produce many aneuploids by self- and cross-pollination.     

CHROMOSOME NUMBERS IN BROMELIACEAE

Eighty-three chromosome counts are reported for 72 taxa of the Bromeliaceae. Fifty-eight of these counts are the first known chromosome number reports for their respective taxa. A model of chromosomal evolution in the Bromeliaceae (n = 25) is presented. The model is parsimonious and consistent with existing data on meiotic chromosome numbers within the family and in the closely related Velloziaceae (n = 9). Two hypothesized paleodiploids (n = 8 and n = 9) hybridized to form a tetraploid that in turn hybridized with the n = 8 lineage. The resultant n = 25 is the extant base number for the family. Two alternative hypotheses could explain the unique extant base number (n = 17) for Cryptanthus: 1) Cryptanthus represents the paleotetraploid level, i.e., prior to the second round of hybridization, or 2) the lower number represents the result of a more recent series of aneuploid reductions from n = 25. Given the existence of intergeneric hybrids involving Cryptanthus, aneuploid reduction is the more likely interpretation.     

CHROMOSOME NUMBERS IN POPULATIONS OF CLAYTONIA VIRGINICA FROM THE NEW YORK METROPOLITAN AREA

Buds of Claytonia virginica were examined from 9 locations in the metropolitan New York area. A large amount of meiotic irregularity was characteristic of the populations studied. Chromosome counts on pollen mother cells were higher than any reported previously for the taxon, which contains an extensive aneuploid series including at least 40 different numbers ranging from 2n = 12 to ca. 190. In one New Jersey population, plants having approximately 90 chromosomes were associated with those having only 16. All the observations indicated that Claytonia virginica is an unusual species with an ability to tolerate fluctuations in chromosome material without apparent effects on fertility and morphology.     

SEMISTERILITY IN THE INTERSPECIFIC HYBRID MELILOTUS POLONICA × M. ALBA

Jaranowski , J. K. (Coll. of Agriculture, ul. Wojska Polskiego 71c, Poznan, Poland.) Semisterility in the interspecific hybrid Melilotus polonica × M. alba. Amer. Jour. Bot. 48(1): 28–35. Illus. 1961.—Interspecific hybrids between Melilotus polonica (n = 8) and M. alba (n = 8) are readily secured. The F₁ hybrids are intermediate between the parents and partially sterile with a mean percentage of 58.8 (ranging from 46.8 to 72.6) defective pollen grains. Six bivalents and a chain or ring of 4 chromosomes occur at diakinesis and metaphase-I of microsporogenesis. A crossshaped configuration characteristic of a reciprocal translocation is present at pachytene, indicating that one of the parents is homozygous for an interchange of relatively large section between two of the members of the chromosome complex. Chromosome bridges, lagging chromosomes, movement of the univalents to the same pole and precocious division of the univalents lead to aberrant chromosome distribution during the course of meiosis. Reduction in self-fertility indicates a corresponding aberrant distribution of chromosomes during megasporogenesis. Pollen sterility in the F₂ generation ranged from 24.8% to 72.5% with a mean value of 54.6%. Two plants in the F₂ generation which had relatively low pollen sterility proved to be aneuploids (2n + 1). Meiotic irregularities in the F₂ plants were comparable to those exhibited by the F₁ plants.     

Variable karyotype with B-chromosomes in Bellevalia saviczii (Liliaceae)

Bellevalia saviczii (Liliaceae), a hexaploid (2n = 6x = 24), was collected from mountain slopes in the Shiraz valley, southern Iran. The basic karyotype consists of one long metacentric chromosome, a slightly shorter acrocentric, and two shorter submetacentrics. In 32% of the plants, one and sometimes two of the acrocentrics (IIa) appeared with a pericentric inversion that changed it into a metacentric (IIm). This metacentric IIm was present in a Hardy--Weinberg ratio throughout the collection range of more than 300 km. A number of aneuploids were found, pentasomics [2n – 1 = 23 (4.9%)], heptasomics [2n + 1 = 25 (2.5%)], and one octasomic [2n + 2 = 26 (0.3%)]. Re-assorted karyotypes with 2n = 24, but with odd numbers of two of the chromosomes, were also present (6.6%). B-chromosomes were found in root meristems and in pedicels in a number (19.1%) of the plants that grew around a 30 km marsh. Their numbers ranged from 1B to 8Bs per plant peaking in the 2B mode. The B-chromosomes (two polymorphs with terminal and near-terminal sticking points) were telocentric with a few metacentric iso-Bs. No B-carrying plants were found in the drier regions away from the marsh. The B-frequency distribution for aneuploids and pericentric inversions was much different than for the standard euploid. Dr Mark Gettner died on November 2nd, 2002. He is fondly remembered by his family, his many students and friends for his kindness and generosity, and for his dedication to his scientific work. Dr Broun is the son-in-law of Dr Gettner and he will be the corresponding author for this article (pb22@york.ac.uk).     

Below-ground carbon and nitrogen accumulation in perennial grasses: A comparison of caespitose and rhizomatous growth forms

An experiment was conducted to compare below-ground soil organic carbon and total nitrogen accumulation between caespitose and rhizomatous perennial grasses in long-term (<25 yrs) grazed and ungrazed sites in semi-arid and mesic communities in the North American Great Plains. Development of greater nutrient pools beneath than between clones occurred at minimal clone basal areas (<60 cm²) for both caespitose species. Caespitose grasses accumulated substantially greater pools of carbon (20–200 fold) and nitrogen (50–500 fold) in soils to a depth of 10 cm beneath clones than rhizomatous grasses accumulated in rhizomes in both communities. Carbon and nitrogen pools in soils beneath caespitose clones exceeded combined (soil + rhizome) pools for rhizomatous grasses for a majority of the clone basal areas (>90 cm²) in the mesic community. In contrast, both pool sizes were smaller beneath the caespitose grass at all clone basal areas than the combined pools for the rhizomatous grass in the semi-arid community. The occurrence of larger soil nutrient pools beneath the rhizomatous species in the semi-arid community was largely a consequence of niche separation for microsites characterized by soils with higher nutrient concentrations, rather than plant-induced increases in nutrient concentrations. Although nutrient islands do not occur beneath rhizomatous grasses, their distribution in the semi-arid community was restricted to microsites characterized by soils with higher SOC and N concentrations. A greater efficiency of nutrient accumulation per unit rhizome mass and the maintenance of rhizome nutrient pools of similar magnitude to those of the rhizomatous grass in the mesic community may also contribute to the distribution of rhizomatous grasses in semi-arid communities. The existence of nutrient islands beneath a wide range of clone sizes in both mesic and semi-arid communities provides circumstantial evidence to suggest that nutrient islands beneath caespitose grasses may contribute to clone fitness in this growth form.     

Aneuploidization under segmental allotetraploidy in rice and its phenotypic manifestation

Key message

We report a repertoire of diverse aneuploids harbored by a newly synthesized segmental allotetraploid rice population with fully sequenced sub-genomes and demonstrate their retention features and phenotypic consequences.

Abstract

Aneuploidy, defined as unequal numbers of different chromosomes, is a large-effect genetic variant and may produce diverse cellular and organismal phenotypes. Polyploids are more permissive to chromosomal content imbalance than their diploid and haploid counterparts, and therefore, may enable more in-depth investigation of the phenotypic consequences of aneuploidy. Based on whole-genome resequencing, we identify that ca. 40% of the 312 selfed individual plants sampled from an early generation rice segmental allotetraploid population are constitutive aneuploids harboring 55 distinct aneuploid karyotypes. We document that gain of a chromosome is more prevalent than loss of a chromosome, and the 12 rice chromosomes have distinct tendencies to be in an aneuploid state. These properties of aneuploidy are constrained by multiple factors including the number of genes residing on the chromosome and predicted functional connectivity with other chromosomes. Two broad categories of aneuploidy-associated phenotypes are recognized: those shared by different aneuploids, and those associated with aneuploidy of a specific chromosome. A repertoire of diverse aneuploids in the context of a segmental allotetraploid rice genome with fully sequenced sub-genomes provides a tractable resource to explore the roles of aneuploidy in nascent polyploid genome evolution and helps to decipher the mechanisms conferring karyotypic stabilization on the path to polyploid speciation and towards artificial construction of novel polyploid crops.

     

INTERSPECIFIC HYBRIDIZATION IN HAPLOPAPPUS AND ITS BEARING ON CHROMOSOME EVOLUTION IN THE BLEPHARODON SECTION

Jackson , R. C. (U. Kansas, Lawrence.) Interspecific hybridization in Haplopappus and its bearing on chromosome evolution in the Blepharodon section. Amer. Jour. Bot. 49 (2) : 119–132. Illus. 1962.—Cytological analyses of interspecific hybrids between H. gracilis (n = 2) and H. spinulosus ssp. australis (n = 8) indicate that ssp. australis is a segmental allotetraploid, derived from past hybridization between 2 taxa with chromosome numbers of n = 4. Analysis of hybrids between H. gracilis (n = 2) and H. ravenii (n = 4), a previously undescribed species, has shown that the chromosome segments of these 2 species are almost completely homologous. Differential contraction is suggested as the explanation for the disappearance in late pachytene of presumed non-homologous segments which were evident in some cells at early pachytene. The pairing relationship of gracilis and ravenii chromosomes at pachytene and later prophase I stages of meiosis indicates that gracilis has evolved from ravenii by an aneuploid reduction process similar to that described for Crepis. The close morphological relationship of the 2 species adds further support to this proposition. Data from the cytological analysis of both interspecific hybrids indicate that x = 4 is the basic chromosome number for the Blepharodon section of Haplopappus.     

Chromosome variation in wheat plants regenerated from cultured immature embryos

Summary A cytological study has been made of plants regenerated from cultured immature embryos of four wheat cultivars (Triticum aestivum, 2n = 6x = 42). In total, 29% of the 192 plants examined were aneuploid with a range in chromosome numbers of 38–45. Evidence of chromosome structural changes was also found. This variation occurred in regenerants of all four cultivars, but there were large differences in the proportions of aneuploids arising from individual cultures which meant that no significant differences could be demonstrated between cultivars. Chromosome abnormalities were present in plants regenerated both from embryogenic cultures and from cultures in which the origin of shoots could not be distinctly defined.     

Karyotypic changes in potato plants regenerated from protoplasts

Over two hundred plants were regenerated from shoot-culture derived proto-plasts of potato (Solanum tuberosum L. cv. Majestic). Some had grossly aberrant phenotypes but the majority were similar to, or indistinguishable from normal control Majestic. Cytological examination showed that on average, 57% of the regenerants had the normal chromosome number (2n=4x=48). The remainder were aneuploids and fell into two classes in approximately equal numbers. The first class was limited at about the euploid level (ie, 2n=44–49). The second class contained plants with higher chromosome numbers ranging from 2n=73 to the octaploid level (2n=8x=96). The overall results represent an improvement over our earlier studies on chromosome variation in protoplast-derived potato plants. In addition, three cases of structural chromosome variation were observed.     

Cytogeography and meiotic chromosome configurations of six intraspecific aneuploids ofCarex conica Boott (Cyperaceae) in Japan

Chromosome numbers were determined for 340 plants ofCarex conica from 83 populations in Japan. Six aneuploids, 2n=32, 33, 34, 36, 37 and 38, were found. Plants with even diploid chromosome numbers 2n=32, 34, and 36 were the most common and had different geographical distributions. Individuals with 2n=32 were from islands in the Seto Inland Sea and nearby coastal areas of the Chugoku District of Honshu; those with 2n=34 were from the Kanto, Chubu and Kinki Districts of Honshu; those with 2n=36 were from the mountainous areas of Chugoku, Shikoku and Kyushu Districts. Canonical discriminant analysis of 17 morphological characters demonstrated that the plants with 2n=32 were clearly distinct from those with 2n=34 or 36. All four aneuploids with even chromosome numbers showed normal bivalent pairing at meiotic metaphase I and probably represent cytogenetically stable cytodemes. Plants with 2n=33 had one heteromorphic trivalent and 15 bivalents, indicating a structural mutation. At mitotic metaphase I, one chromosome was markedly larger than the others, suggesting that the 2n=33 plants arose from 2n=34 plants by fusion of two chromsomes. The plant with 2n=37 was intermediate in morphology betweenCarex conica (2n=36) andC. morrowii (2n=38) and probably originated as an interspecific hybrid between these species.     

CHROMOSOMAL RELATIONSHIPS AND TAXONOMIC CONSIDERATIONS IN THE GENUS CASSIA

Irwin , H. S. and B. L. Turner . (U. Texas, Austin.) Chromosomal relationships and taxonomic considerations in the genus Cassia. Amer. Jour. Bot. 47(4): 309-318. 1960.—All chromosome numbers known for the genus Cassia are arranged in a table which follows the infrageneric organization of Bentham (1871). Included are new counts for 88 taxa, of which 68 are first reports. Evidence is represented which suggests that the base number for the genus is 7, the other numbers having been derived by aneuploid loss from various euploid levels. The basic number 8, of general occurrence in the Subsection Leiocalyx, Section Chameacrista, is presumed to have been derived from the ancestral generic base by aneuploid gain of 1 chromosome. The results of the present study do not support elevation of the section Chamaecrista to generic rank. In general, the basic soundness of Bentham's revision is upheld.     

The fitness costs and benefits of trisomy of each Candida albicans chromosome

Candida albicans is a prevalent human fungal pathogen. Rapid genomic change, due to aneuploidy, is a common mechanism that facilitates survival from multiple types of stresses including the few classes of available antifungal drugs. The stress survival of aneuploids occurs despite the fitness costs attributed to most aneuploids growing under idealized lab conditions. Systematic study of the aneuploid state in C. albicans has been hindered by the lack of a comprehensive collection of aneuploid strains. Here, we describe a collection of diploid C. albicans aneuploid strains, each carrying one extra copy of each chromosome, all from the same genetic background. We tested the fitness of this collection under several physiological conditions including shifts in pH, low glucose, oxidative stress, temperature, high osmolarity, membrane stress, and cell wall stress. We found that most aneuploids, under most conditions, were less fit than their euploid parent, yet there were specific conditions under which specific aneuploid isolates provided a fitness benefit relative to the euploid parent strain. Importantly, this fitness benefit was attributable to the change in the copy number of specific chromosomes. Thus, C. albicans can tolerate aneuploidy of each chromosome and some aneuploids confer improved growth under conditions that the yeast encounters in its host niches.     

Frequency,origin, and survival of aneuploids in tetraploid ryegrass

Aneuploids with 26 to 30 chromosomes occurred in progenies of induced tetraploids of Lolium perenne in a frequency of 6 to 23% in C₁ families, 12% in C₂ and 6% in advanced generations. The most frequent types had 27 and 29 chromosomes. Most were apparently recovered through female transmission, which varied for aneuploid gametes with different chromosome numbers and which probably depended on the chromosome involved. In general, aneuploids showed a reduction in fertility. Even with the same chromosome number, fertility varied perhaps as a result of differences between the chromosomes involved. Aneuploids could not be distinguished from eu-tetraploids on a morphological basis. Tetraploids late and early in respect to flowering did not differ in aneuploid frequency. No reversion to diploidy was observed in successive generations.