Clonal repeatability of in vitro pollen tube growth rates in Oenothera organensis (Onagraceae)

The concept that pollen selection is an important force driving angiosperm evolution has received a lot of attention in recent years. In order for pollen competition to have evolutionary significance, heritable variation for pollen tube growth rate must exist. In this study, I tested the clonal repeatability, a measure of broad-sense heritability, of in vitro pollen tube growth rate in Oenothera organensis. Although I found significant phenotypic differences in pollen tube growth rate, the calculated clonal repeatability value of 9.4% indicates that heritability of pollen tube growth rate in this species is quite low. In contrast, both flower identity and plant identity had highly significant effects on pollen tube growth rate, suggesting that the environment may be more important than genotype in determining pollen performance in this species.     

Flavonoids of Heterogaura (Onagraceae)

Heterogaura is a monotypic genus of the tribe Onagreae of the Onagraceae. It is endemic to south western Oregon and California. Four flavonol glycosides, kaempferol 3-O-rhamnoside, quercetin 3-O-glucoside, quercetin 3-O-rhamnoglucoside and myricetin 3-O-glucoside, were found to occur in methanolic leaf extracts of each of the populations sampled. The presence of only flavonols is consistent with flavonoid analyses from other genera of the Onagreae, including Clarkia, the closest relative of Heterogaura.     

Temporal variation in the pollen:ovule ratios of Clarkia (Onagraceae) taxa with contrasting mating systems: field populations

Among plants, pairs of selfing vs. outcrossing sister taxa provide interesting systems in which to test predictions concerning the magnitude and direction of temporal changes in sex allocation. Although resource availability typically declines towards the end of the growing season for annual taxa, temporal changes in mating opportunities depend on mating system and should change less in selfing taxa. Consequently, given that the pollen:ovule (P:O) ratio of flowers reflects the investment in (and potential fitness pay-off due to) male vs. female function, we predicted that the P:O ratio should also be less variable among and within selfers than in closely related outcrossers. To test these predictions, we measured temporal changes in sex allocation in multiple field populations of two pairs of sister taxa in the annual flowering plant genus Clarkia (Onagraceae). In the outcrossing Clarkia unguiculata and the selfing Clarkia exilis, ovule production declined similarly from early to late buds, whereas pollen production remained constant or increased in the outcrosser but remained constant or decreased in the selfer. Consequently, the P:O ratio increased within unguiculata populations but marginally increased or stayed constant in exilis populations. In all populations of the selfing Clarkia xantiana spp. parviflora and the outcrossing C. x. spp. xantiana, both ovule and pollen production per flower declined over time. The effects of these declines on the P:O ratio, however, differed between subspecies. In each xantiana population, the mean P:O ratio did not differ between early and late flowers, although individuals varied greatly in the direction and magnitude of phenotypic change. By contrast, parviflora populations differed in the mean direction of temporal change in the P:O ratio. We found little evidence to support our initial predictions that the P:O ratio of the selfing taxa will consistently vary less than in outcrossing taxa.     

Chromosomal evolution inEpilobium sect.Epilobium (Onagraceae)

WithinEpilobium sect.Epilobium, a cytological analysis of 121 experimental hybrids, involving 40 species, indicates the presence of a widespread BB chromosome arrangement in Eurasia, Africa, and Australasia, as well as in North and South America less commonly. The AA chromosome arrangement, which differs from BB by one reciprocal translocation, occurs in North America, South America, and in at least three European species. The CC arrangement, which differs from AA by two reciprocal translocations, characterizes theAlpinae, a circumboreal group. Distinctive or only partly worked out chromosome arrangements occur in the EuropeanE. duriaei andE. nutans and in the North AmericanE. luteum, E. obcordatum, E. oregonense, andE. rigidum. With earlier results, the chromosome arrangements of some 65 of the estimated 185 species of the section have been established fully or partly.     

Developmental characteristics of floral organs and pollen of Chinese cabbage (Brassica campestris L. ssp. chinensis)

Morphological and cytological studies are complementary approaches to understand the molecular mechanisms that regulate floral developmental pathways. To better understand abnormal mutant phenotypes in floral development, we conducted detailed observations and investigations of the morphology, cytology, and cell ultrastructure of wild-type Chinese cabbage (Brassica campestris L. ssp. chinensis Makino and syn. B. rapa ssp. chinensis) flowers when they developed from primordia to anthesis. First, we measured bud and organ length with a stereo microscope and observed the developmental status and characteristics of the floral organs using a scanning electron microscope; then we made thin slices of anthers to observe the developmental stage and characteristics of pollen using an optical microscope; and finally, we made super-thin slices of anthers to observe the ultrastructure of pollen during its development with the aid of a transmission electron microscope. In this study, the floral developmental continuum was divided into 17 stages based on significant changes in the shape of floral primordia, and the pollen developmental continuum was divided into 14 stages based on the developmental characteristics. The results could provide the morphological basis for further research on the molecular mechanisms that regulate development of the floral organs and/or pollen of Chinese cabbage and their allied species.     

Cytogenetic investigations onOenothera wolfii (Onagraceae)

Oenothera wolfii, endemic in coastal Northern California and Southern Oregon, has been analysed cytogenetically. It is a permanent structural heterozygote composed of two nearly identical genomic complexes which are closely related to the neighboring maritime ecotype of the homozygousOe. elata subsp.hookeri. — Oe wolfii is believed to have evolved recently from its homozygous ancestor by the accumulation of reciprocal translocations and the acquisition of balanced lethals. Forms such asOe. wolfii represent an important connecting link in our understanding of the evolution of the complexheterozygous species ofOenothera. Dedicated to Prof. Dr.Josef Straub for His 75th birthday.     

Self-sterility in Epilobium obcordatum (Onagraceae)

Greenhouse pollinations among 23 field-collected plants and their progeny revealed a high and variable degree of self-sterility in Epilobium obcordatum. Field-collected plants averaged 5% seed-set when selfed and 43% seed-set in crosses; greenhouse progeny 11% and 49%, respectively. In two of three pollen chase tests, previously or simultaneously applied self-pollen reduced seed-set to that resulting from pure self-pollination. Based on the relative survivorship of self- and outcross embryos, the mean number of lethal equivalents was determined to be 11.0 (range 2.9 to 17.6) in the field-collected plants, and 7.1 (range 0.62 to 21.8) in the greenhouse progeny. Pollinations among 91 individuals showed a significant negative correlation between seed-set and inbreeding coefficients of the resulting zygotes. Intrapopulation crosses yielded significantly fewer seeds than interpopulation crosses. Early embryonic mutational load, in combination with some level of biparental inbreeding, appears to be responsible for the level of self-sterility in natural populations of this species. A pollen dosage test for an interpopulation cross resulted in a maximum seed set of 47% at 16 tetrads. Additional unmeasured characteristics govern the probability of ovules becoming mature seeds.     

Cytogenetic investigations onOenothera nutans (Onagraceae)

Oenothera nutans, common to the Appalachian Mts between 650 and 1 700 m altitude, was investigated cytogenetically and taxonomically. The species is permanently structurally heterozygous. It consists of two genomes of the B-type which are more or less indistinguishable phenotypically. Nearly all of the strains investigated possess a self-incompatibility factor in one of the two complexes. Both complexes show a close relationship to the predominantly homozygousO. grandiflora, a native of the southern lowlands.O. nutans andO. grandiflora possess the same plastid type, plastome III. Probably,O. nutans evolved by an accumulation of reciprocal translocations within an originally structurally homozygous population, which must be regarded ancestral to the present forms ofO. grandiflora.     

Size-dependent pollen:ovule ratios and the allometry of floral sex allocation in Clarkia (Onagraceae) taxa with contrasting mating systems

Multiple field populations of two pairs of diploid sister taxa with contrasting mating systems in the genus Clarkia (Onagraceae) were surveyed to test predictions concerning the effects of resource status, estimated as plant size, on pollen and ovule production and on the pollen:ovule (P:O) ratio of flowers. Most theoretical models of size-dependent sex allocation predict that, in outcrossing populations, larger plants should allocate more resources to female function. Lower P:O ratios in larger plants compared to smaller plants have been interpreted as supporting this prediction. In contrast, we predicted that P:O ratio should not vary with plant size in predominantly selfing plants, in which each flower contributes to reproductive success equally through male and female function. We found that, in all four taxa, both ovule and pollen production per flower usually increased significantly with plant size and that the shape of this relationship was decelerating. However, ovule production either decelerated more rapidly than or at the same rate as pollen production with plant size. Consequently,the P:O ratio increased or had no relationship with plant size. This relationship was population-specific (not taxon-specific) and independent of the mating system. Possible explanations for the increasing maleness with plant size are discussed.     

The status of Clarkia australis (Onagraceae)

Clarkia australis and C. virgata grow on the western slope of the central Sierra Nevada of California. Clarkia australis was established to accommodate populations of C. virgata from south of the Tuolumne River that could not be successfully hybridized to populations north of the river. Although the species is maintained in the new Jepson Manual, its validity has been questioned because only two populations were originally tested, and they had no useful morphological traits that distinguished them from C. virgata. We report here the results of a large program of interpopulation hybridizations that show that C. australis is distinct and that its reproductive isolation from C. virgata is complete and absolute and reflects a compatibility block that apparently causes abortion of hybrid seeds in early development. Both species include populations north and south of the Tuolumne River and, in general, those of C. australis occupy higher elevations. Morphologically, the species are extremely similar though the mean values of several dimensions of the petals are different. However, significant variation among their populations has the consequence that, at present, the only certain way to assign particular populations to species is to test their compatibility with previously tested populations.     

Morphological and pollen differentiation in Solanum cardiophyllum ssp. cardiophyllum and S. cardiophyllum ssp. ehrenbergii

A multivariate analysis to differentiate morphologically the populations of wild potatoes Solanum cardiophyllum ssp. cardiophyllum and S. cardiophyllum ssp. ehrenbergii was carried out. An analysis of the morphology and the viability of pollen of these potato populations was also made. The results of the morphometric analysis indicate that both subspecies are phenetically different. The pollen grain shape and size in ssp. cardiophyllum are different in northern and southern populations. Pollen diameter is significantly different between subspecies. Based on these results we propose that these taxa should be considered as two separate species.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 415–426.     

Ovule fates in Epilobium obcordatum (Onagraceae)

We investigated the fates of ovules following self- and cross-pollination of the variably self-sterile Epilobium obcordatum by observing thin sections of ovaries 4, 5, and 10 d postpollination (PP). Three types of unfertilized ovules were observed: apparently normal ovules not entered by pollen tubes, nonfunctional ovules with a nucellus but lacking an embryo sac, and empty ovules with integuments but lacking a nucellus. The average frequencies for all three types were not different for selfs and crosses. Fertilization frequencies did not differ among pollination treatments, averaging 61.4% of the total sample of 4 and 5 d PP ovules. The phenotypes of failing postfertilization ovules at 4 and 5 d PP included those containing preglobular and globular embryos lacking endosperm, ovules with endosperm but no embryo present, and failing embryos in the presence of endosperm. Self- and cross-fertilized 10 d PP ovules also possessed heart-stage and torpedo-stage failing embryos. Early embryo failures (5 d PP) were distributed throughout the length of the ovary, as expected if failures are the consequence of factors intrinsic to each ovule. The sizes of normally developing self- and cross-fertilized embryos were not different at 4 d PP. The 5 d PP cross embryos were larger than selfs. A comparison of 10 d samples was not possible due to attrition of self embryos. Although ovular position did not affect the probability of early embryo failure, it did affect the size of normally developing embryos at 4, 5, and 10 d PP, with embryos at the stylar end larger than those at the basal end.     

Observations of meiotic chromosomes in Gaura (Onagraceae)

Observations of meiotic chromosomes are reported for all 21 species and 3 additional sub species ofGaura (Onagraceae), based upon a study of 647 individuals from 509 naturally occurring populations throughout the range of the genus. The basic chromosome number for the genus isx = 7, and 18 species are diploid withn = 7. Among these, the self-incompatible ones are often highly chromosomally heterozygous, with no homozygous individuals having been found in nature in the perenrennialsGaura lindheimeri andG. villosa, and two-thirds or more of the individuals apparently heterozygous in the following well-sampled species:G. calcicola, G. longiflora, andG. suffulta subsp.suffulta. In contrast, the autogamous species are entirely chromosomally homozygous or nearly so. Two species ofGaura are reported as chromosomal structural heterozygotes, with about 50% pollen abortion:G. biennis andG. triangulata; the translocation systems originated independently of one another. Two of the three polyploid species,G. sinuata andG. drummondii (G. odorata of many authors), are consistently tetraploid (n = 14) and, despite their cytological autotetraploidy, are thought to have originated following interspecific hybridization. They are the only rhizomatous species in the genus and may have had one ancestor in common. The remaining polyploid,G. coccinea, includes populations withn = 7, 14, 21, and 28, as well as evident interploid hybrids and, frequently, supernumerary chromosomes. The relationship among these populations is close and is maintained by frequent hybridization and exchange of genetic material. No other species seems to have participated in their origin, and the association of their chromosomes is consistently that characteristic of autopolyploidy in plants with tetraploid and higher chromosome numbers.     

The systematics of autopolyploidy in Epilobium latifolium (Onagraceae)

Diploid (2n = 36) plants ofEpilobium latifolium L. have been found in Alaska and the western Cordillera of North America, and tetraploids in Iceland, western Greenland, and southeastern Quebec. Study of the morphology of these chromosome races revealed a character, the number of pores in the pollen grains, by which they could be distinguished with an average probability of 75%. Tetraploids ofE. latifolium tend to have high percentages of 4-pored pollen, while the diploids usually have only 3-pored pollen. On this basis, the ranges of the races were extrapolated from herbarium material. Detailed comparisons of eight other features of the presumptive diploids and tetraploids showed no significant difference between them. The races appear to form worldwide, allopatric, ecogeographic phases. A high level (57.4%) of quadrivalent formation observed during tetraploid meiosis, and other considerations, indicate that the tetraploids arose by autopolyploidy. The recommendation is made that the races not be given formal taxonomic recognition.     

Unique tetrads of <Emphasis Type="Italic">Epilobium luteum</Emphasis> (Onagraceae: Onagreae) pollen from Alaska

Aperture morphology of tetrad pollen of Epilobium luteum (Onagraceae: Epilobieae) from three Alaskan collections is highly variable. The first collection appears to lack apertures altogether and is presumed to consist of immature pollen gains in a genus known to achieve mature size before the apertures become distinctly protruding. A second collection has tetrads with 3- and 4-apertured grains, the apertures in the latter are often irregularly spaced and not in apposition with the apertures of neighboring members. The third collection consists of the more typical 3-apertured members that characterize the majority of Epilobium pollen grains. In all of these collections individual pollen grains (monads) are interspersed among the tetrads. The variations in the number of apertures emphasize the importance of having a comprehensive understanding of the stage of development of the pollen (taxon) examined when describing pollen collections. In the first collection this would mean the recognition that in Onagraceae apertures occur in the later stages of microspore ontogeny. In the latter two collections a thorough background of the literature of the pollen morphology on this largest Onagraceae taxon is useful for the understanding of the significance of a range of aperture numbers on Epilobium pollen grains.