Gynodioecy in <i>Plantago lanceolata</i>:VI. Functions of cytokinins in growth, development, and reproduction of two sex types

Theoretical models predict that male sterile plants of gynodioecious species should show at least some compensation for their disadvantage of not reproducing as males through female component of fitness. In this study, growth, development, and reproduction of a hermaphrodite and a male sterile family of Plantago lanceolata L. were compared under controlled conditions. The male sterile plants produced more and longer spikes and had relatively longer styles. The male sterile plants achieved their final biomass sooner, by an earlier formation of side rosettes, and flowered earlier. The hypothesis was tested as to whether cytokinins in the plants are involved as a pleiotropic factor in either or both sex expression and the various plant characteristics associated with the male sterile phenotype. The roots of the male sterile plants had higher concentrations of putative zeatin riboside than the roots of the hermaphroditic plants, as quantified by an enzyme-linked immunoassy after separation of cytokinins by high performance liquid chromatography. Spraying the plants with benzyladenine did not affect internal cytokinin concentrations or sex expression. Benzyladenine spray increased the growth rate of the main rosette and stimulated floral initiation. Our results indicate that cytokinins are possibly involved in determining the morphological differences between sex types in this species

Genetics of Male-Sterility in Gynodioecious Plantago-Coronopus .2. Nuclear Genetic-Variation

Inheritance of male sterility was studied in the gynodioecious species Plantago coronopus using five plants and their descendants from an area of similar to 50 m(2) from each of four locations. In each location, crosses between these five plants yielded the entire array of possible sex phenotypes. Both nuclear and cytoplasmic genes were involved. Emphasis is placed on the nuclear (restorer) genetics of two cytoplasmic types. For both types, multiple interacting nuclear genes were demonstrated. These genes carried either dominant or recessive restorer alleles. The exact number of genes involved could not be determined, because different genetic models could be proposed for each location and no common genetic solution could be given. At least five genes, three with dominant and two with recessive restorer allele action, were involved with both cytoplasmic types. Segregation patterns of partially male sterile plants suggested that they are due to incomplete dominance at restorer loci. Restorer genes interact in different ways. In most instances models with independent restorer gene action were sufficient to explain the crossing results. However, for one case we propose a model with epistatic restorer gene action. There was a consistent difference in the segregation of male sterility between plants from the two cytoplasmic types. Hermaphrodites of cytoplasmic type 2 hardly segregated male steriles, in contrast to plants with cytoplasmic type 1. [KEYWORDS: Cytoplasmic male-sterility; origanum-vulgare l; nucleocytoplasmic polymorphism; lanceolata l; evolution

Effects of seed size, inbreeding and maternal sex on offspring fitness in gynodioecious Plantago coronopus

Male steriles (MS) must have a fitness advantage relative to hermaphrodites (H) if they are to be maintained in gynodioecious species. We report experiments in which we disentangle the relative contributions of seed size, inbreeding and maternal sex to the fitness advantage of male steriles in Plantago coronopus L. 2 Seed size effects were observed throughout growth experiments in the glasshouse and were reflected in all biomass measurements. In the field, seed size effects resulted in a fourfold increase in standardized seed production per initial buried seed after 2 years between small (mean weight = 0.13 mg) and large (0.20 mg) seeds. 3 Inbreeding depression, calculated from seed to seed was = 0.37 after one generation of selfing and = 0.93 after the second generation of selfing. Regression of log(1 ) on inbreeding level suggested synergistic epistasis in fitness. 4 Even after taking into account the effects of seed size and inbreeding level, the offspring of a male sterile mother had a 16% advantage over a hermaphrodite, but this disappeared when the progeny sex ratio (the ratio of MS : H individuals among the offspring) was taken into account. 5 In the field, offspring of large seeds had both a higher overall incidence of flowering, and a higher probability of flowering in their first year, thus generating an extra cohort of individuals. The high inbreeding depression in fitness after two generations of selfing was also due to a very low incidence of flowering among the S2 individuals. Flowering probability therefore appears to be a critical trait in this system. 6 In the field, the contributions of seed size variation (15%) and inbreeding (9%) combine with 48% higher seed production to give a total fitness advantage of 70% of male steriles relative to hermaphrodites. This is probably sufficient for maintenance of gynodioecy under nuclear-cytoplasmic inheritance of male sterility. 7 Both inbreeding effects (as a consequence of the sexual system) and pleiotropic effects (of the genes coding for male sterility) play a role in the maintenance of gynodioecy in this species, with an apparently greater role for the latter. [KEYWORDS: compensation ; flowering ; germination ; growth analysis ; inbreeding depression ; male sterility ; maternal sex ; seed size ; sex ratio ; spatial variation]