REPRODUCTIVE ISOLATION BARRIERS TO GENE EXCHANGE BETWEEN SOLANUM CHACOENSE AND S. COMMERSONII (SOLANACEAE)

In contrast to the general intercompatibility of diploid species of tuber-bearing potatoes of South America, Solanum chacoense Bitter and S. commersonii Dunal have well-established internal barriers to gene exchange. Pollen tubes of S. commersonii are inhibited in growth down the style of S. chacoense. While S. chacoense pollen tubes are not inhibited in S. commersonii stylar tissue, seed development is abnormal, resulting in production of large numbers of seeds that are not filled out. Even the apparently normal F₁ seed germinated poorly; approximately 0.1 germinable seeds were set per cross as compared with over 30 seeds per cross following intraspecific crosses. Lowland S. chacoense, including populations sympatric with S. commersonii, were not more effectively isolated from S. commersonii than were highland S. chacoense populations, over 500 km from the nearest S. commersonii. The F₁ hybrids produced were vigorous and highly fertile. Crosses between unrelated S. commersonii-chacoense F₁ hybrids produced only 1.3 germinable seeds per cross. This reduction resulted from low numbers of berries set, few seeds per berry, and low seed germination. The F₂ progeny were vigorous, but while many were both male and female fertile there were indications of an increase in sterility over that of the parents and F₁'s. The data were interpreted as indicating that the isolation barriers result from polygenic differences between the parent species and are composed principally of genes that function in several of the processes of seed setting. The accumulation of genes that keep these species isolated was interpreted as not having developed as a result of selection for such isolation in the region of overlap (Wallace effect), but rather as a byproduct of divergence that occurred during their allopatric evolution.     

FUNCTIONAL DIOECY AND ANDROMONOECY IN SOLANUM

Field and laboratory studies of 19 diclinous species endemic to Australia help to clarify the nature and evolution of andromonoecy, androdioecy, and dioecy in the genus Solanum. Ten species are andromonoecious; typically these species bear inflorescences with a single, large basal hermaphroditic flower and 12–60 distal, smaller staminate flowers. We suggest that the andromonoecious condition was derived from hermaphroditic-flowered ancestors in part by hemisterilization of flowers but largely by addition of staminate flowers. The resultant larger inflorescences are hypothesized to serve both to attract and to entrain pollinators, yielding more or higher-quality seed set in hermaphroditic flowers and/or greater dispersion of pollen from staminate flowers. We suggest that andromonoecy may also serve to reduce selling. Nine other species are morphologically androdioecious but functionally dioecious. In these species, staminate flowers, like those of the andromonoecious species, bear anthers with copious tricolporate pollen and a highly reduced gynoecium. The morphologically hermaphroditic flowers are functionally pistillate and borne singly in inflorescences, and they bear anthers with inaperturate pollen. The inaperturate pollen, although viable, never germinates and is hypothesized to be retained in pistillate flowers as a reward to pollinators in the nectarless Solanum flowers. All other species of Solanum studied with pollen dimorphism in which one pollen morph is inaperturate are also best treated as functionally dioecious. We conclude that there is no evidence for androdioecy in Solanum. A review of other families suggests that there is little support for this unusual breeding system in any other angiosperm group either. Preliminary analyses suggest that andromonoecy and dioecy are polyphyletic in Solanum. Furthermore, dioecy is as likely to have arisen from hermaphroditic as from andromonoecious ancestors.     

CYTOLOGY AND REPRODUCTIVE CHARACTERISTICS IN PENNISETUM SETACEUM

A green ecotype of fountaingrass, Pennisetum setaceum (Forsk.) Chiov., was found to be triploid (3x = 27) with 9 II + 9 I at metaphase I. Meiotic behavior of the bivalents was normal, but the univalents lagged and divided precociously at anaphase I. The univalent halves lagged at anaphase II and were distributed at random. Micronuclei were common, and stainable pollen was approximately 40%. The green type produced four progeny with 54 chromosomes. A purple strain of P. setaceum was found to be hexaploid (6x = 54) with varying chromosomal associations. Meiotic behavior was highly irregular and stainable pollen was almost zero. Megasporogenesis was initiated in both types, but the megaspores degenerated and nucellar and integument cells developed into four-nucleate, aposporous embryo sacs. Following self-pollination, the green type set approximately 10% seed and the purple type set 0.05% seed. Purple fountaingrass set as high as 18% seed following the application of pollen from Pennisetum ciliare (L.) Link (4x = 36), exemplifying a rare form of pseudogamy between two species. Improvement of P. setaceum through a standard breeding program appears unlikely.     

PHENOTYPIC VARIATION IN TRISOMICS OF CLARKIA UNGUICULATA

Vasek , F. C. (U. California, Riverside.) Phenotypic variation in trisomics of Clarkia unguiculata. Amer. Jour. Bot. 50(4): 308–314. 1963.—Progenies of 3n × 2n crosses included, in addition to diploids, plants trisomic for 1, 2, 3, 4, 5, 6 or 7 chromosomes. Means and variances were calculated for 15 phenotypic traits, including 3 width/length ratios, in one set of progenies, and for 10 of the traits, including 2 of the ratios, in another set of progenies. In 25 trait comparisons, including 15 different traits, the means for each chromosome number class were heterogeneous in 11 comparisons, which included 8 different traits. Single trisomics differed significantly from diploids in 5 comparisons (4 different traits). Despite these significant differences the variation followed no particular pattern except that sepal length increased and pollen fertility decreased with chromosome number, and trisomics, as a group, sometimes differed from diploids with regard to the width/length ratio of leaves or petals. The variances were heterogeneous in 5 comparisons (4 different traits). Sepal length and pollen fertility were the only traits for which single trisomics were more variable than diploids and for which the entire population was more variable than diploids. In addition, in 4 progenies of self-pollinated trisomics, diploids and trisomics (which within each progeny were trisomic for the same Chromosome) differed significantly in mean value in only 3 traits (out of 60 trait comparisons). Variances were significantly different in 6 comparisons (4 different traits) but, surprisingly, diploids were more variable than trisomics in 3 of these 6 comparisons. Except for sepal length, pollen fertility and some width/length ratios, a wide variety and number of extra chromosomes rarely had a significant effect on the mean or variability of various phenotypic traits, and single specific extra chromosomes had very little effect except sometimes on pollen fertility or an occasional ratio. A large amount of variation, probably caused by the environment and the general genetic background, may obscure possible specific trisomic phenotypes. Morphological identification of specific trisomics is considered impractical in this species.     

TRIVALENT FORMATION IN MULTIPLE TRISOMICS OF CLARKIA UNGUICULATA

Vasek, F. C. (U. California, Riverside.) Trivalent formation in multiple trisomics of Clarkia unguiculata. Amer. Jour. Bot. 50(3): 244–247. 1963.—A series of multiple trisomics, ranging from 2n + 1 to 2m + 7, was scored for univalents and trivalents at first metaphase. The mean number of trivalents per cell per extra chromosome was 0.43, 0.46, 0.46, 0.56, 0.54, 0.55 and 0.59 for plants with 1, 2, 3, 4, 5, 6 and 7 extra chromosomes respectively. These trivalent frequencies are interpreted to fall into 2 classes, indicating that the cytological differences between triploids and single trisomies may be based on a threshold effect. The results are compared with available data from other genera, and it is suggested that the observed increase in trivalent formation per extra chromosome may be associated with an increase in chiasma frequency.     

LABILE SEX EXPRESSION IN ANDROMONOECIOUS SOLANUM HIRTUM: FLORAL DEVELOPMENT AND SEX DETERMINATION

Sex expression (the proportions of hermaphrodite and staminate flowers produced) of the andromonoecious species Solatium hirtum is labile, and this lability of whole plant sex expression is due to labile sex expression of individual floral buds. In this paper I examine the developmental processes that underlie the differences in floral sex expression of hermaphrodite and staminate flowers of Solarium hirtum, focusing particularly on the processes responsible for the observed lability of floral sex expression. Differences in bud growth rate and relative growth of floral organs in these buds are evident at about the time of megasporocyte meiosis (11–12 days before anthesis). However, gynoecial sterility in staminate buds does not occur until just 6–7 days before anthesis. At this time, abnormalities in ovule development occur in staminate buds: the ovules begin to appear necrotic, the integumentary tapetum collapses, and the megagametophytes of many ovules cease normal development. These observations are consistent with the predictions of labile floral development.     

CYTOLOGY AND MORPHOGENESIS IN THE RHIZOMORPH OF ARMILLARIA MELLEA

Aided by the techniques of thin sectioning and electron microscopy, the apical region of the rhizomorph of Armillaria mellea has been examined. This region is composed of concentric zones of morphologically distinct tissues derived from a subapical meristematic zone designated the apical center. Meristematic activity is of two types: (1) primary, localized in the apical center, in which new hyphal elements are formed from apical initials, and (2) secondary, localized in the lateral regions of the apex, in which elaboration of the hyphal elements by means of elongation and secondary crosswall formation takes place. From these meristematic zones the tissues of the mature rhizomorph are derived and include: (a) peripheral hyphae, (b) cortex, (c) subcortex, and (d) primary and secondary medulla. The manner of differentiation of an apical initial appears unique and involves synchronous nuclear divisions accompanied by segmentation in many planes. The result of this activity is the formation of multinucleate hyphae. Apical initials are usually highly cytoplasmic and possess peculiar non-membrane-bound fibrous bundles, but in all other respects they resemble the hyphae of most Basidiomycetes thus far examined with the electron microscope.     

INTERSPECIFIC INCOMPATIBILITY IN SOLANUM SPECIES

Pandey , K.K. (Crop Res. Div., D.S. & I.R., Lincoln, Christchurch, New Zealand.) Interspecific incompatibility in Solanum species. Amer. Jour. Bot. 49(8): 874–882. Illus. 1962.—A diallel cross involving 11 self-incompatible and 3 self-compatible species of Solanum was made to study the genetic basis of interspecific incompatibility. Interspecific incompatibility was not limited to crosses in which a self-compatible species was used as the male parent onto a self-incompatible species (unilateral incompatibility). A number of crosses between self-incompatible species were incompatible. In one cross, Q vernei X verrucosum, a self-compatible species was successful as a pollen parent with a self-incompatible species. Unlike other hybrids between self-compatible and self-incompatible species, which are self-incompatible, these F₁ hybrids were self-fertile, and cross-fertile among themselves and with both parents. The self-fertile S. polyadenium was cross-incompatible as a female as well as a male parent with all other species. It is suggested that the unilateral incompatibility is a property of the allele S_C which originated as a consequence of one kind of breakdown of the S_I gene; the S_C allele produces “bare” pollen growth substances which are inactivated in an incompatible style. It is proposed that the failure of the principle of unilateral interspecific incompatibility in solanaceous species may be due to the action of alleles at the second incompatibility locus revealed in certain Mexican species. It is assumed that the South American species are selected intraspecifically only for the action of S alleles but that in certain interspecific crosses and rarely in intraspecific crosses the alleles at the second locus may be expressed, thus interfering with the usual action of S alleles. The F₁ hybrids Q verrucosum (self-fertile) X simplicifolium (self-sterile) were self-incompatible at the tetraploid as well as the diploid level, and their cross-compatibility behavior was consistent with the expected activity of the S_C and S_I alleles of the 2 parents respectively.     

REPRODUCTION IN ZONARIA FARLOWII II. CYTOLOGY AND ULTRASTRUCTURE1

The development of spores and eggs of Zonaria farlowii was studied with light and electron microscopy. Oogonia require a lunar month to develop and are produced in periodic crops. The sporangia, on the other hand, are longer in developing and on a particular plant do not become mature all at the same time. Differences in size, content, and products of sporangia and oogonia have been found. One of the quantifiable differences is in the amount of osmiophilic substance, presumably oil, present in the 2 cells. This substance is more abundant in the sporangium, a cell which produces 8 reproductive spores. These differences are particularly interesting since the embryology of sporic and zygotic germlings has been found to be identical, while ontogeny of spores and eggs differs. The findings of this study are discussed in view of possible relationships between microstrtrclitral features of spores and fertilized eggs and their subsequent development.