Preface. Part I: Molecules and genomes in plant systematics. Chloroplast DNA and the study of plant phylogeny: present status and future prospects - M T Clegg and G Zurawski Use of chloroplast DNA rearrangements in reconstructing plant phylogeny - S R Downie and J D Palmer Mitochondrial DNA in plant systematics: applications and limitations - J D Palmer Ribosomal RNA as a phylogenetic tool in plant systematics - R K Hamby and E A Zimmer Evolution of the NOR and 5S DNA loci in the Triticeae - R Appels and B Baum Part II: Molecular approaches to plant evolution Intraspecific chloroplast DNA variation: systematic and phylogenetic implications - D E Soltis, P S Soltis and B G Milligan Molecular data and polyploid evolution in plants - P S Soltis, J J Doyle and D E Soltis Molecular systematics and crop evolution - J Deobley Part III: Model studies of phylogenetic relationships Contributions of molecular data to polyploid evolution in plants - P S Soltis, J J Doyle and D E Soltis Molecular systematics and crop evolution - J Deobley Contributions of molecular data to papilionoid legume systematics - J J Doyle, M Levin and A Bruneau Chloroplast DNA variation in the asteraceae: phylogenetic and evolutionary implications - R K Jansen, H J Michaels, R S Wallace, K-J Kim, S C Keeley, L E Watson and J D Palmer Chloroplast DNA restriction site variation and the evolution of the annual habit in North American Coreopsis (Asteraceae) - D J Crawford, J D Palmer and M Kobayashi Molecular systematics of onagraceae: examples from Clarkia and Fuschia - K J Systema and J E Smith Floral morphology and chromosome number in the subtribe oncidiinae (Orchidaceae): evolutionary insights from a phylogenetic analysis of the chloroplast DNA restriction site variation - M W Chase and J D Palmer Part IV: Theoretical perspectives The suitability of molecular and morphological evidence in reconstructing plant phylogeny -M J Donaghue and M J Sanderson Character-site weighting for restriction site data in phylogenetic reconstruction, with an example from chloroplast DNA - V A Albert, B D Mishler and M W Chase Polymorphism, hybridization and variable evolutionary rate in molecular phylogenies - K Ritland and J E Eckenwalder Index.

Molecular Systematics of Plants

The evolvement of duplicated gene loci in allopolyploid plants has become the subject of intensive studies. Most duplicated genes remain active in neoallopolyploids contributing either to a favourable effect of an extra gene dosage or to the build-up of positive inter-genomic interactions when genes or regulation factors on homoeologous chromosomes are divergent. However, in a small number of loci (about 10%), genes of only one genome are active, while the homoeoalleles on the other genome(s) are either eliminated or partially or completely suppressed by genetic or epigenetic means. For several traits, the retention of controlling genes is not random, favouring one genome over the other(s). Such genomic asymmetry is manifested in allopolyploid wheat by the control of various morphological and agronomical traits, in the production of rRNA and storage proteins, and in interaction with pathogens. It is suggested that the process of cytological diploidization leading to exclusive intra-genomic meiotic pairing and, consequently, to complete avoidance of inter-genomic recombination, has two contrasting effects. Firstly, it provides a means for the fixation of positive heterotic inter-genomic interactions and also maintains genomic asymmetry resulting from loss or silencing of genes. The possible mechanisms and evolutionary advantages of genomic asymmetry are discussed.

Genomic asymmetry in allopolyploid plants: wheat as a model

We examined the structure, intranuclear distribution and activity of ribosomal DNA (rDNA) in Nicotiana sylvestris (2n = 2x = 24) and N. tomentosiformis (2n = 2x = 24) and compared these with patterns in N. tabacum (tobacco, 2n = 4x = 48). We also examined a long-established N. tabacum culture, TBY-2. Nicotiana tabacum is an allotetraploid thought to be derived from ancestors of N. sylvestris (S-genome donor) and N. tomentosiformis (T-genome donor). Nicotiana sylvestris has three rDNA loci, one locus each on chromosomes 10, 11, and 12. In root-tip meristematic interphase cells, the site on chromosome 12 remains condensed and inactive, while the sites on chromosomes 10 and 11 show activity at the proximal end of the locus only. Nicotiana tomentosiformis has one major locus on chromosome 3 showing activity and a minor, inactive locus on chromosome 11. In N. tabacum cv. 095-55, there are four rDNA loci on T3, S10, S11/t and S12 (S11/t carries a small T-genome translocation). The locus on S12 remains condensed and inactive in root-tip meristematic cells while the others show activity, including decondensation at interphase and secondary constrictions at metaphase. Nicotiana tabacum DNA digested with methylcytosine-sensitive enzymes revealed a hybridisation pattern for rDNA that resembled that of N. tomentosiformis and not N. sylvestris. The data indicate that active, undermethylated genes are of the N. tomentosiformis type. Since S-genome chromosomes of N. tabacum show rDNA expression, the result indicates rDNA gene conversion of the active rDNA units on these chromosomes. Gene conversion in N. tabacum is consistent with the results of previous work. However, using primers specific for the S-genome rDNA intergenic sequences (IGS) in the polymerase chain reaction (PCR) show that rDNA gene conversion has not gone to completion in N. tabacum. Furthermore, using methylation-insensitive restriction enzymes we demonstrate that about 8% of the rDNA units remain of the N. sylvestris type (from ca. 75% based on the sum of the rDNA copy numbers in the parents). Since the active genes are likely to be of an N. tomentosiformis type, the N. sylvestris type units are presumably contained within inactive loci (i.e. on chromosome S12). Nicotiana sylvestris has approximately three times as much rDNA as the other two species, resulting in much condensed rDNA at interphase. This species also has three classes of IGS, indicating gene conversion has not homogenised repeat length in this species. The results suggest that methylation and/or DNA condensation has reduced or prevented gene conversion from occurring at inactive genes at rDNA loci. Alternatively, active undermethylated units may be vulnerable to gene conversion, perhaps because they are decondensed and located in close proximity within the nucleolus at interphase. In TBY-2, restriction enzymes showed hybridisation patterns that were similar to, but different from, those of N. tabacum. In addition, TBY-2 has elevated rDNA copy number and variable numbers of rDNA loci, all indicating rDNA evolution in culture.

Gene conversion of ribosomal DNA in Nicotiana tabacum is associated with undermethylated, decondensed and probably active gene units.

Regulation of cytosine methylation in ribosomal DNA and nucleolus organizer expression in wheat

Polyploidy is a significant force in plant evolution. Approximately 47% to 52% of all angiosperm species are polyploid (V. Grant, 1981). Estimates of the frequency of polyploidy in pteridophytes range from 43.5% for the ferns alone (Vida, 1976) to 95% for pteridophytes as a whole (V. Grant, 1981), suggesting ancient polyploidy in several lineages of pteridophytes. In byrophytes polyploidy is common in mosses, but rare in liverworts (V. Grant, 1981). In contrast to angiosperms, pteridophytes, and bryophytes, polyploidy in gymnosperms is very rare and sporadic. Polyploidy has not been detected in cycads or ginkgo, and only 1.5% of the species of Coniferales are polyploid (Khoshoo, 1959). In the Gnetales, tetraploidy is common in Ephedra and rare or nonexistent in Gnetum and Welwitschia (Delevoryas, 1980).

Molecular Data and Polyploid Evolution in Plants

The major nucleoli in the wheat variety Chinese Spring are formed at the nucleolus organisers (NORs) on chromosomes 1B and 6B. Minor nucleoli are formed from the NORs on chromosomes 5D and 1A. Nucleolar volume is poorly correlated with the number of ribosomal RNA genes in the NOR region. The nucleolus on chromosome 1B is twice the volume of that on chromosome 6B even though only half as many ribosomal RNA genes reside at the 1B locus compared with the number at the 6B locus. Nucleolus size is correlated with the size of the NOR constriction seen in metaphase chromosomes. When major NORs are deleted, the volumes of the remaining nucleoli increase to compensate for the deletions. Varying the dosage of many of the chromosomes in the total complement influences the volume of the minor nucleoli. Chromosomes 1D and 6B have been shown to carry genes regulating total nucleolar volume. Although total nucleolar volume differs between cells at different stages of development, the genetic variation influencing volume appears to have similar effects in root tip, premeiotic and young pollen cells.

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Guido Martini

Papers

The control of nucleolus volume in wheat, a genetic study at three developmental stages