Jacqueline M. Nugent

Physical and gene mapping studies reveal that chloroplast DNA from geranium (Pelargonium hortorum) has sustained a number of extensive duplications and inversions, resulting in a genome arrangement radically unlike that of other plants. At 217 kilobases in size, the circular chromosome is about 50% larger than the typical land plant chloroplast genome and is by far the largest described to date, to our knowledge. Most of this extra size can be accounted for by a 76-kilobase inverted duplication, three times larger than the normal chloroplast DNA inverted repeat. This tripling has occurred primarily by spreading of the inverted repeat into regions that are single copy in all other chloroplast genomes. Consequently, 10 protein genes that are present only once in all other land plants are duplicated in geranium. At least six inversions, occurring in both the inverted repeat and large single-copy region, must be postulated to account for all of the gene order differences that distinguish the geranium genome from other chloroplast genomes. We report the existence in geranium of two families of short dispersed repeats and hypothesize that recombination between repeats may be the major cause of inversions in geranium chloroplast DNA.

ABSTRACT Plant development depends on the activity of meristems which continually reiterate a common plan. Permutations around this plan can give rise to a wide range of morphologies. To understand the mechanisms underlying this variation, the effects of parallel mutations in key developmental genes are being studied in different species. In Antirrhinum, three of these key genes are: (1) floricaula (flo) a gene required for the production of flowers (2) centrora- dialis (cen), a gene controlling flower position (3) cycloidea (eye), a gene controlling flower symmetry. Several plant species, exhibiting a range of inflorescence types and floral symmetries are being analysed in detail. Comparative genetic and molecular analysis shows that inflorescence architecture depends on two underlying parameters: a basic inflorescence branching pattern and the positioning of flowers. The flo and cen genes play a key role in the positioning of flowers, and variation in the site and timing of expression of these genes, may account for many of the different inflorescence types. The evolution of inflorescence structure may also have influenced the evolution of floral asymmetry, as illustrated by the cen mutation which changes both inflorescence type and the symmetry of some flowers. Conflicting theories about the origins of irregular flowers and how they have coevolved with inflorescence architecture can be directly assessed by examining the role of eye- and cen-like genes in species displaying various floral symmetries and inflorescence types.

The respiratory gene cox2, normally present in the mitochondrion, was previously shown to have been functionally transferred to the nucleus during flowering plant evolution, possibly during the diversification of legumes. To search for novel intermediate stages in the process of intracellular gene transfer and to assess the evolutionary timing and frequency of cox2 transfer, activation, and inactivation, we examined nuclear and mitochondrial (mt) cox2 presence and expression in over 25 legume genera and mt cox2 presence in 392 genera. Transfer and activation of cox2 appear to have occurred during recent legume evolution, more recently than previously inferred. Many intermediate stages of the gene transfer process are represented by cox2 genes in the studied legumes. Nine legumes contain intact copies of both nuclear and mt cox2, although transcripts could not be detected for some of these genes. Both cox2 genes are transcribed in seven legumes that are phylogenetically interspersed with species displaying only nuclear or mt cox2 expression. Inactivation of cox2 in each genome has taken place multiple times and in a variety of ways, including loss of detectable transcripts or transcript editing and partial to complete gene loss. Phylogenetic evidence shows about the same number (3-5) of separate inactivations of nuclear and mt cox2, suggesting that there is no selective advantage for a mt vs. nuclear location of cox2 in plants. The current distribution of cox2 presence and expression between the nucleus and mitochondrion in the studied legumes is probably the result of chance mutations silencing either cox2 gene.