APOMIXIS IN EUPATORIUM TANACETIFOLIUM (COMPOSITAE)

Cytogenetics and embryological studies of male sterility have been reported for the first time in Eupatorium tanacetifolium Gill, ex H. et A. (Gyptis pinnatifida Cass.). This species produces viable seeds but abnormal pollen that is not shed by the anthers. There are great abnormalities in karyokinesis and cytokinesis in microsporogenesis that result in irregular sporads formed by 5–10 cells of variable size, shape, and chromosome number. There is an irregular distribution of chromosomes, due to absence of regular pairing and disjunction, presence of chromatinic bridges in most of telophases, and to succesive aberrant cytokinesis without formation of cell plate and with variably oriented walls. The two chromatids of each chromosome presumably remain joined until the end of the process. Somatic chromosome number of 2n = 30 is reported for one population of this species, an apomict taxon of probable triploid origin. Embryo-sac ontogeny is of the Antennaria type of diplospory, wherein embryo and endosperm development are parthenogenetic.     

SEED FERTILITY IN TRAGOPOGON HYBRIDS (COMPOSITAE)

The seed fertility of 172 interspecific and 32 intraspecific F₁ hybrids in the genus Tragopogon was determined. Intrageneric barriers to interbreeding were variable; some were weak and others quite strong. Maternal influences on fertility were found in 18 pairs of reciprocal interspecific crosses. It was postulated that cytoplasmic and nuclear factors interacted to determine the level of seed fertility. Although F₁'s were generally sterile or semi-sterile, the proportion of fertile seeds produced in most crosses increased in the F₂ generation. In two cases, however, a significant decrease was observed.     

CYTOTAXONOMIC STUDIES IN CHRYSOTHAMNUS (ASTEREAE,COMPOSITAE)

The basic chromosome number for Chrysothamnus is x = 9 as determined by 234 original counts representing all species. Polyploidy is frequent only in C. viscidiflorus where diploids, tetraploids, and hexaploids are found. Pollen size in the genus is basically correlated with style length rather than ploidy level. Most aspects of meiosis appear normal except for the tendency for nucleolar fragmentation. To reduce chromosome clumping during meiosis, an ice-water pretreatment was utilized. Karyotypes in Chrysothamnus reflect similar trends of specialization noted in chromosome structure for other Compositae. The taxa are predominantly self-fertilized; however, one instance of interspecific hybridization is noted. Taxonomic significance of the chromosome numbers, karyotypes, and breeding system in Chrysothamnus is discussed.     

GENETIC DIVERGENCE AND GEOGRAPHIC SPECIATION IN LAYIA (COMPOSITAE)

Electrophoretic variability was examined in six species of Layia (Compositae), native to California, which have previously been studied by Clausen, Keck, and Hiesey, and are regarded as a classic example of geographic speciation in plants. The study was carried out to test the hypothesis that the extent of divergence in structural genes coding enzymes is concordant with divergence in morphological characteristics, ecological traits, and reproductive isolation. Eleven enzymes specified by 17 loci were analyzed. The genetic identity values were consistent with those expected on the model that the species diverged gradually as they adapted to geographically separate habitats. Thus, the values between the three species complexes proposed by Clausen, Keck, and Hiesey (L. chrysanthemoides/L. fremontii; L. jonesii/L. leucopappa/L. munzii; L. platyglossa) were substantially lower than the values between species within the complexes. The results provide an important contrast to the very high genetic identities between species which originated rapidly from their progenitors. The electrophoretic results also provided evidence that the cytosolic isozyme of phosphoglucomutase and the cytosolic NADP-dependent isocitrate dehydrogenase in the six species are coded by duplicate genes.     

FLORAL FLAVONOIDS AND ULTRAVIOLET PATTERNS IN VIGUIERA (COMPOSITAE)

Variation occurs among species of Viguiera series Viguiera for ultraviolet (uv) absorption/reflection patterns of ligules. Floral flavonoids that cause uv absorption occur in epidermal papillae. Flavonoids are further localized to the proximal portion of the ligule in the seven taxa that have only proximal uv absorption. Floral flavonoids involved in uv absorption consist of flavone, flavonol, and anthochlor (chalcone/aurone) glycosides. Quercetin 3-methyl ether glycosides characterize the ligules of 10 taxa occurring in Baja California, Mexico, and nearby areas, and these taxa appear to form one taxonomic group. The anthochlor pair, marein/maritimein, characterizes V. dentata, and the lack of ligule flavonoids distinguishes V. potosina from the remaining taxa. The presence of the anthochlor pair, marein/maritimein, only in V. dentata and the lack of ligule flavonoids in V. potosina concur with other data to indicate that these species are not correctly placed with each other or with the other species currently included in series Viguiera.     

PHOSPHOGLUCOMUTASE AND ISOCITRATE DEHYDROGENASE GENE DUPLICATIONS IN LAYIA (COMPOSITAE)

Genetic analysis of isozyme segregation patterns in Layia (Compositae) showed that cytosolic phosphoglucomutase isozymes are encoded by duplicated genes, and that the cytosolic NADP-dependent isocitrate dehydrogenase isozymes are encoded by duplicated genes in species with haploid chromosome numbers of n = 7 and triplicated genes in those with n = 8. The duplicated genes specifying both isozymes assorted independently in all species tested. An electrophoretic survey of phosphoglucomutase in diploid species representing six additional genera of Madiinae, the subtribe to which Layia is assigned, revealed that Achyrachaena, Calycadenia, Hemizonia, Holocarpha, and Madia all possessed duplicated genes. In Lagophylla, one species also had duplicated genes for the isozyme but a second species did not, a loss probably resulting from mutation or chromosomal deletion. The phosphoglucomutase duplication characterizes nearly the entire subtribe and may prove useful to identify phylogenetic relationships between the Madiinae and other subtribes.     

POLYPLOIDY,DYSPLOIDY, AND CHROMOSOME PAIRING IN ECHEVERIA (CRASSULACEAE) AND ITS HYBRIDS

The 140+ species of Echeveria have more than 50 gametic chromosome numbers, including every number from 12 through 34 and polyploids to n = ca. 260. With related genera, they comprise an immense comparium of 200+ species that have been interconnected in cultivation by hybrids. Some species with as many as 34 gametic chromosomes include none that can pair with each other, indicating that they are effectively diploid, but other species with fewer chromosomes test as tetraploids. Most diploid hybrids form multivalents, indicating that many translocations have rearranged segments of the chromosomes. Small, nonessential chromosomal remnants can be lost, lowering the number and suggesting that higher diploid numbers (n = 30–34) in the long dysploid series are older. These same numbers are basic to most other genera in the comparium (Pachyphytum, Graptopetalum, Sedum section Pachysedum), and many diploid intergeneric hybrids show very substantial chromosome pairing. Most polyploid hybrids here are fertile, even where the parents belong to different genera and have very different chromosome numbers. This seems possible only if corresponding chromosomes from a polyploid parent pair with each other preferentially, strong evidence for autopolyploidy. High diploid numbers here may represent old polyploids that have become diploidized by loss, mutation, or suppression of duplicate genes, but other evidence for this is lacking. Most species occur as small populations in unstable habitats in an area with a history of many rapid climatic and geological changes, presenting a model for rapid evolution.     

STRUCTURE AND ONTOGENY OF LATICIFERS IN CICHORIUM INTYBUS (COMPOSITAE)

The branched anastomosed laticifer system in the primary body of Cichorium intybus L. originates in embryos from files of laticiferous members at the boundary between phloic procambium and ground meristem. Upon seed germination, laticiferous members develop perforations in the end walls which become entirely resorbed. Perforations also develop in the longitudinal walls of contiguous laticiferous members and from lateral connections between developing laticifer branches. Additional laticiferous members originate as procambium differentiation proceeds, and their differentiation follows a continuous acropetal sequence in leaf primordia of the plumule. In roots, laticifers closely associated with sieve tubes in the secondary phloem originate from derivatives of fusiform initials in the vascular cambium. These laticifers develop wall perforations and in a mature condition resemble laticifers in the primary body. As the girth of the root increases, laticifers toward the periphery, unlike associated sieve tubes, resist crushing and obliteration. Laticifers vary in width from about 4 to 22 μm; the widest ones occur in involucral bracts and the narrowest ones in florets. There was no evidence that intrusive growth occurs during development of the laticifer system, although such growth may occur during development of occasional branches which extend through ground tissue independent of phloem and terminate in contact with the epidermis. Presence of amorphous callose deposits is related to aging of laticifers and mechanical injury.     

INFRASPECIFIC VARIATION OF SESQUITERPENE LACTONES IN AMBROSIA PSILOSTACHYA (COMPOSITAE)

The distribution of fifteen sesquiterpene lactones in sixty-two populations of Ambrosia psilostachya ranging from Canada to Mexico, and cytological observations for seven populations are reported. Chemical data for four Mexican populations of A. cumanensis Kunth are presented. The origin of chemically unique populations of A. psilostachya which occur on the islands and peninsulas that line the Texas Gulf coast are discussed with respect to the chemical, morphological and cytological data.     

CHROMOSOME NUMBERS IN SOUTH AMERICAN HAPLOPAPPUS CASS. (COMPOSITAE)

Chromosome counts are reported for 33 species from all four sections of the genus Haplopappus in South America. These include first reports for 28 species and two putative hybrids. All chromosome numbers reported herein are 2n = 5_II, with the exception of H. prunelloides with 2_n = 6_II. Unlike the North American species, the morphological diversity of South American taxa is not concomitant with chromosomal variation.     

VARIATION AND LOCALIZATION OF FLAVONOID AGLYCONES IN HELIANTHUS ANNUUS (COMPOSITAE)

Flavonoid aglycone variation within Helianthus annuus, a species widely distributed throughout North America, was analyzed. Flavonoid aglycones of H. annuus consist of two types, flavones and chalcones. The flavone aglycones are sequestered in glandular trichomes that occur on both leaf surfaces, whereas the chalcone aglycones appear to be incorporated in the waxy leaf cuticle. Considerable variation in flavonoid profile was observed with some plants exhibiting as few as one, and others as many as seven of the eight aglycones detected. No definable phytogeographic patterns were observed for this flavonoid variation. Flavonoid aglycone variation also did not differentiate the infraspecific taxa within H. annuus.     

CHROMOSOME STUDIES IN COMPOSITAE

Original chromosome observations, including number, from meiotic preparations from 65 collections representing 57 species in 43 genera of Compositae are reported. Of these, eight are first reports. Meiotic irregularities were noted in several species; pollen stainabilities are given for these. Observations that differ from earlier reports are discussed.     

FLAVONOID EVOLUTION IN ROBINSONIA (COMPOSITAE) OF THE JUAN FERNANDEZ ISLANDS

Leaf flavonoids were isolated and identified from 54 populations representing all seven species of Robinsonia, a genus of dioecious rosette trees endemic to the Juan Fernandez Islands. Fourteen compounds were detected consisting of flavonols, flavones, flavanones and dihydroflavonols. The distribution of these compounds in Robinsonia largely corresponds to specific and sectional limits based on morphological data. The morphologically similar species, R. gayana and R. thurifera, have identical flavonol profiles (derivatives of quercetin). Likewise, the closely related R. evenia and R. masafuerae are unique in the genus by possessing flavones. The inclusion of Rhetinodendron (i.e., R. berteroi) in Robinsonia is supported by its strong flavonoid similarity with species in two other sections of the genus. The morphologically diverse section Eleutherolepis exhibits the greatest flavonoid variation of any section, and only here are found flavones, flavanones and dihydroflavonols. The direction of flavonoid evolution in Robinsonia is hypothesized to be from fewer to more classes of compounds. Biosynthetic considerations suggest that this gain in compounds is due both to a gain of an additional enzymatic step and to the sequestering of precursors. This interpretation of direction of flavonoid evolution is in agreement with several lines of evidence including the flavonoid chemistry of the hypothesized outgroup (i.e., species of Senecio on mainland Chile), the ages of the two islands, and morphological trends.     

DEVELOPMENT OF NORMAL AND SEEDLESS ACHENES IN CICHORIUM INTYBUS (COMPOSITAE)

Cross- and partially cross-pollinated capitula of Cichorium intybus (Compositae, Lactuceae) were examined for a study of normal and seedless fruit development respectively. Embryos develop according to the Asterad pattern, and the free-nuclear endosperm becomes cellular 15–17 hrs after pollination. A zone of disorganized cellular material surrounds the embryo sac at anthesis, and, in normal achenes, this zone expands as the seed develops. Initially the developing seed elongates and comes into contact with the top of the ovary by 48 hrs. In contrast to this pattern, the ovule in developing seedless achenes degenerates within 72 hrs. Irregularities, such as an abnormally proliferating endothelium, embryo formation without endosperm, and endosperm formation without an embryo often accompany this degeneration. Differentiation of the pericarp in seeded achenes begins between 48 and 72 hrs, starting at the apex and proceeding basipetally; in seedless fruits the process is similar though initiated somewhat later. The normal pericarp at maturity exhibits a pigmented exocarp, a broad mesocarp of thick-walled lignified cells, and a tenuous endocarp. In seedless achenes the fruit coat is similar except that the exocarp is colorless and the cells of the mesocarp are relatively small.     

THE FLAVONOIDS OF LASTHENIA (COMPOSITAE)

Lasthenia (Compositae: Helenieae), a western North American-Chilean genus of 16 species, produces 22 flavonoid glycosides. Flavonoids are the chalcones butein and okanin, the aurones maritimetin and sulfuretin, the flavone luteolin, and the flavonols kaempferol, quercetin, and patuletin. The presence or absence of various of these classes of compounds in general follows sectional alignments in the genus, confirms affinities based on morphological and cytological evidence, and suggests relationships of problematical species. Intraspecific variation in flavonoid constituents occurs in several species, and in one taxon intrapopulation variation seems to exist as well. Evolution within Lasthenia has been associated with a loss of the ability to produce or accumulate luteolin, chalcones, and aurones; an increase in diversity of quercetin glycosides; acquisition of the ability to produce patuletin; and an elaboration of glycosylation patterns of patuletin.     

DEVELOPMENT OF THE PHYTOMELANIN LAYER IN FRUITS OF AGERATUM CONYZOIDES (COMPOSITAE)

The development of the phytomelanin layer in the achenes of Ageratum conyzoides (Compositae, Eupatorieae) was studied using light and electron microscopy. At the level of the embryo sac, the young ovary wall contains an outer zone, consisting of an epidermis and two hypodermal layers, and an inner zone, consisting of developing fiber cells and 3–5 layers of parenchyma. A schizogenous space forms between the developing fibers and the inner hypodermis at about the time that the embryo sac is fully organized. At this stage, the developing fibers contain papilla which are outgrowths that connect the fibers to the inner hypodermal cells. After fertilization, phytomelanin accumulates on the cell walls lining this space. Subsequently, by the time the fruit matures, the phytomelanin fills the space completely and forms a solid, black layer. The surface of the inner hypodermis that faces the space forms a mold; the characteristic peglike projections of the mature phytomelanin layer develop by filling the invaginations between the hypodermal cells. During phytomelanin accumulation, abundant smooth endoplasmic reticulum is present in the hypodermis, especially in the outer layer. It is hypothesized that the precursors of the phytomelanin are synthesized in this endoplasmic reticulum and that these precursors migrate into the space where the phytomelanin is polymerized.     

ASSOCIATIONS BETWEEN ALLOZYME FREQUENCIES AND SOIL CHARACTERISTICS IN GAILLARDIA PULCHELLA (COMPOSITAE)

Selection favoring different alleles in different environments frequently has been suggested as an explanation for allozyme variation within and among populations. This hypothesis predicts that allozyme frequencies will be correlated with environmental variables. Previous studies on allozyme frequency-environment covariation in plants often have relied on qualitative assessments of the environment and have emphasized highly autogamous species. We have examined allozyme frequency-soil associations in Gaillardia pulchella, an obligately outcrossed annual plant, by regressing the frequencies of 15 common allozymes representing six polymorphic enzyme loci on principal components from a set of 20 quantitative soil variables. Fifty-one populations, representing four taxonomic varieties, were included in the analysis. Among the 26 populations representing the var. pulchella, allozymes Adh-2f and Pgm-1c were significantly associated with a block of highly inter-correlated soil characteristics which serve to discriminate between soils derived from calcareous vs. non-calcareous rock types. This geographically complex pattern of allozyme frequency-soil covariation is not likely to be spurious and, thus, indicates the presence of adaptively differentiated soil races, or ecotypes. However, these results are not sufficient to conclude that the allozyme frequency divergence between ecotypes was mediated by selection, either directly or through genetic hitchhiking. The pattern of allozyme frequency-soil covariation within var. pulchella was not found among the other taxonomic varieties. Patterns of genotype-environment covariation often may be recognizable only within geographically or environmentally restricted groups of populations because of the confounding influences of other environmental variables.