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The flowers of Clarkia gracilis subsp. sonomensis have large petals each with a large, central, red-purple spot while the flowers of subsp. gracilis are small and unspotted. Other pigmentation (anthocyanin) patterns also vary within and between these subspecies. We carried out a genetic analysis of differences in floral patterns and petal size. A novel basal petal spot appeared in the F2. The analysis indicated that the novel petal spot was specified by an allele in subsp. gracilis at a locus governing spot position. This allele is not normally expressed in subsp. gracilis because of the action of a modifier gene at a second locus. The study also indicated single factor inheritance for presence versus absence of pigmentation on the hypanthium, stamens, and the lower portion of the petals. Multifactorial inheritance was observed for differences in petal length and width. Most of the possible recombinant floral patterns were recovered in the F2 and F3. This system can be used to study developmental regulation of floral traits as well as ecological relationships between floral pattern and pollination system. The recovery of a normally unexpressed allele for basal petal spot points to the difficulty of extrapolating from phenotypic analysis to an understanding of morphological evolution.

Allotetraploid plant species combine the genomes of related diploid species, but little is known about whether homologous genes from the diploid genomes are expressed, how they interact, or whether they evolve differently when in a common tetraploid nucleus. Polyploidy may lead to gene silencing, but few molecular characterizations of silenced genes encoding enzymes in polyploids and related diploids have been reported. Here we describe the PgiC genes in the tetraploid Clarkia gracilis and related diploid species, which are native from California to southern Washington. PgiC encodes the cytosolic isozyme of phosphoglucose isomerase (PGIC; EC 5.3.1.9). The gene was duplicated in the basal stock of Clarkia and now both genes, PgiC1 and PgiC2, are active in about half of the diploid species, whereas only PgiC1 is active in the others. Clarkia gracilis was found to have three PgiC genes: two PgiC1s and a PgiC2. Reverse-transcriptase-polymerase chain reaction (RT-PCR) experiments, starting with mRNAs prepared from seedling leaves of C. gracilis, showed that the three genes are expressed. Analysis of their sequences showed they are evolving at similar rates to their homologues and that they have the same intron-exon structure. The presence of an expressed PgiC2 in C. gracilis was unexpected because all related diploid species, including one identified as a parent, have only active PgiC1s. The donor of the PgiC2 is now presumed extinct, but parsimony analysis identified its phylogenetic position. None of the PgiC genes that were active when C. gracilis arose were silenced. A possible example of gene conversion involving a 300-nuclectide region of one PgiC1 and PgiC2 was identified, but it probably occurred in the diploid parental species rather than in C. gracilis. PgiC2 is the first known example of an active locus in a tetraploid plant species that is no longer expressed in its diploid relatives.

Clarkia (Onagraceae) is a genus of 42 annual species, mostly native to California, that has served as a model for many studies of plant evolutionary biology, particularly morphological, cytological, and genetic divergence; reproductive isolation; and speciation. Section Sympherica is the largest section with eight diploid and one allotetraploid species. Species in the section have provided important evidence about the evolution of reproductive isolation (C. lingulata derived from C. biloba) and large morphological change (C. dudleyana thought to be sister to the morphologically distinct C. heterandra, recently transferred into Clarkia from the monotypic Heterogaura). Clarkia epilobioides, another diploid species in the section, was previously shown to be one parent of the allotetraploid C. delicata, the other parent being C. unguiculata from sect. Phaeostoma. Lewis and Lewis (1955) interpreted the parentage of C. delicata and other evidence of intersectional hybridization to mean that the diploid sections of the genus, though highly diverse, were closely related and should be maintained in the single genus Clarkia. Here we assess phylogenetic relationships among the species of sect. Sympherica and related species by analyzing the nucleotide sequences of PgiC1 and PgiC2, a pair of paralogous genes that encode the cytosolic isozyme of phosphoglucose isomerase (EC 5.3.1.9). The major results were the following: (1) C. unguiculata and both genomes of C. delicata are within a well-defined "Sympherica" clade; thus, C. delicata should not be considered an intersectional hybrid; (2) C. heterandra belongs in the clade and is closely related to C. unguiculata; and (3) on the evidence of PgiC1, C. dudleyana is not in the clade and is not closely related to C. heterandra.