A SIX-YEAR STUDY OF FLAVONOID DISTRIBUTION IN A POPULATION OF LASTHENIA CALIFORNICA (ASTERACEAE)

A population of the annual Lasthenia californica in the Jasper Ridge Biological Preserve (of Stanford University) showed three major flavonoid pigment types (A, B and C; a fourth was seen in very low frequencies). The three more common types exhibited a suite of aurone and chalcone glucosides and a set of highly polar flavonoid glucuronides. Type C consisted solely of the base array; type B added luteolin 7-glucoside to the base array, while type A exhibited the base array plus eriodictyol 7-glucoside and flavonol 3,7-diglycoside bisulfates. The rare type D profile resembled type A except that the diglycosides were not sulfated. Collections were made along one of several fixed transects annually from 1982 to 1987. The main transect studied (Transect No. I) can be divided into two distinct parts on the basis of a changeover of flavonoid pattern from a mixture of types B and C at one end to exclusively (or very nearly) type A at the other. The frequencies of types B and C varied from year to year with type C predominating every year. The frequency of type A plants remained remarkably stable over the six-year period. Collections along other transects showed a similar constancy of both frequencies of the flavonoid types and location of the types along the transects. Growth experiments in a greenhouse showed that seeds from type A parents harvested in the field produced only type A progeny, while type B and C plants gave only type B and C progeny with type C predominating (a single type A progeny plant was obtained from a type C parent). Flavonoid diversity in L. californica appears to be genetically controlled and is influenced significantly by the flavonoid chemistry of the seed parent.     

VARIATION OF NUCLEAR DNA CONTENT IN HELIANTHUS ANNUUS (ASTERACEAE)

Nuclear 2C DNA content was determined by laser flow cytometry for 13 diploid (2n = 34) lines (cultivated varieties and inbred lines) of Helianthus annuus. Mean DNA amount of second leaf nuclei varied from 6.01 to 7.95 pg (32%) among lines. Mean DNA content varied up to 19% within lines. Variability in mean DNA content exceeding 27% and 48% was detected among leaves from different nodes of plants of the open-pollinated variety, Californicus, and the inbred line, RHA 299, respectively. The root tip and shoot tip nuclei of H. annuus have been reported to consist of a mixture of aneuploid (17 to 33 chromosomes) and diploid (34 chromosomes) cells, a condition called aneusomaty. Chromosome counts of root tips and an analysis of the distribution of DNA content of large numbers of nuclei from leaves indicate that aneusomaty either does not occur, or is not common, among the lines investigated. The intraspecific, intraline, and intraplant variation in DNA content in H. annuus support the concept that a sizable portion of a plant genome is unstable and subject to rapid changes in DNA amount.     

FLAVONOID DIVERSIFICATION IN THE ACHILLEA AGERATIFOLIA AND A. CLAVENNAE GROUPS (ASTERACEAE)

The Achillea ageratifolia and A. clavennae groups (Asteraceae, tribe Anthemideae) were investigated for intra- and interspecific leaf flavonoid variation. Numerous flavonoid glycosides, mostly unidentified, and variously methylated free aglycones based on scutellarein, 6-hydroxyluteolin, quercetagetin, and 6-hydroxy-kaempferol were detected. The distribution of methylated aglycones among individuals of A. ageratifolia was used to model the putative biosynthetic pathways. Several steps in the biosynthesis probably involve enzymes (methyltransferases) with strict positional specificities and broader substrate specificities. Geographical patterns in the flavonoid content of A. ageratifolia were only partly consistent with the current concept of A. ageratifolia. Different flavonoid profiles in A. ageratifolia do not appear to be of crucial importance in its adaptation to different habitats. The species of the A. clavennae group were clearly separable on flavonoid profiles and the presence of a hybrid population was supported by the flavonoid data. Interspecific differences in number of glycosides in relation to the number of free aglycones are discussed.     

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.     

PHYLOGENETIC IMPLICATIONS OF CHLOROPLAST DNA RESTRICTION SITE VARIATION IN THE MUTISIEAE (ASTERACEAE)

Phylogenetic relationships among 13 species in the tribe Mutisieae and a single species from each of three other tribes in the Asteraceae were assessed by chloroplast DNA restriction site mapping. Initially, 211 restriction site mutations were detected among 16 species using 10 restriction enzymes. Examination of 12 of these species using nine more enzymes revealed 179 additional restriction site mutations. Phylogenetic analyses of restriction site mutations were performed using both Dolio and Wagner parsimony, and the resulting monophyletic groups were statistically tested by the bootstrap method. The phylogenetic trees confirm an ancient evolutionary split in the Asteraceae that was previously suggested by the distribution of a chloroplast DNA inversion. The subtribe Barnadesiinae of the tribe Mutisieae is shown to be the ancestral group within the Asteraceae. The molecular phylogenies also confirm the paraphyly of the Mutisieae and provide statistical support for the monophyly of three of its four currently recognized subtribes (Barnadesiinae, Mutisiinae, and Nassauviinae). The fourth subtribe, Gochnatiinae, is shown to be paraphyletic. Within the subtribes, several closely related generic pairs are identified. Chloroplast DNA sequence divergence among genera of the Asteraceae ranges between 0.7 and 5.4%, which is relatively low in comparison to other angiosperm groups. This suggests that the Asteraceae is either a relatively young family or that its chloroplast DNA has evolved at a slower rate than in other families.     

EVOLUTIONARY IMPLICATIONS OF INTRASPECIFIC CHLOROPLAST DNA VARIATION IN DWARF DANDELIONS (KRIGIA; ASTERACEAE)

Intraspecific chloroplast DNA polymorphisms were examined for 51 populations of seven species in the genus Krigia. A total of 1,100 restriction sites was surveyed and 46 of these were variable at the intraspecific level. Twenty-two of the variable sites were found within K. virginica, giving this species one of the highest levels of intraspecific chloroplast DNA divergence of any examined species. In contrast, no restriction site variation was detected within K. dandelion, K. wrightii, and K. occidentalis. Five polymorphisms were identified from the 16 populations of the K. cespitosa-gracilis complex, but no mutations distinguished the K. cespitosa and K. gracilis types. Krigia montana and K. biflora showed 11 and eight restriction site polymorphisms, respectively. The chloroplast genome of the hexaploid K. montana was derived from the diploid K. biflora rather than the tetraploid K. montana. High levels of polymorphism were found in species having different ploidy levels, such as K. virginica, K. biflora, and K. montana. Furthermore, most mutations found in these three species were recorded from the tetraploid lineages. As a result, evolutionary rates between different ploidy levels differ significantly. The chloroplast DNA restriction site data suggest that all surveyed populations of the autotetraploid K. virginica originated from a common ancestor. Our results also indicate that certain regions of the chloroplast genome have changed more rapidly than others and have the potential to resolve evolutionary questions at the population level.     

SYSTEMATIC IMPLICATIONS OF CHLOROPLAST DNA VARIATION IN THE SUBTRIBE MICROSERIDINAE (ASTERACEAE: LACTUCEAE)

Phylogenetic relationships among six of the seven currently recognized genera of the subtribe Microseridinae were examined using comparative restriction site mapping of the chloroplast genome. Complete cleavage maps of 17 enzymes were constructed for 35 of the 46 species in the subtribe and six representative outgroup species from four other subtribes of the Lactuceae. A total of 373 restriction site mutations was detected, 180 of which were phylogenetically informative. The chloroplast DNA mutations were used to generate phylogenetic trees using Dollo, Wagner, and weighted parsimony, and the resulting monophyletic groups were evaluated by the bootstrap method. The results indicate that the Microseridinae is paraphyletic because Stephanomeria is nested within the six core genera of the subtribe. Five of the genera of the Microseridinae are monophyletic, whereas Microseris is paraphyletic because one of its species, M. lindleyi, is more closely related to Agoseris and Nothocalais. The distant relationship of M. lindleyi to the rest of the species of Microseris is enigmatic, especially in view of its known involvement in the origin of two allotetraploid species, M. decipiens and M. heterocarpa. The taxonomic implications of these results lead to a revised classification of Microseris in which the genus Uropappus is revived to accommodate M. lindleyi, and a new genus, Stebbinsoseris, is described to include the allotetraploids. The cpDNA phylogeny is used to evaluate the patterns of character evolution within the Microseridinae, as well as to test previous hypotheses that differences in generation time result in unequal rates of chloroplast DNA evolution.     

CHARACTERIZATION OF INVASIVE CONYZA SPECIES (ASTERACEAE) IN EUROPE: QUANTITATIVE TRAIT AND ISOZYME ANALYSIS

The genus Conyza (Asteraceae) represents one of the foremost examples of intercontinental plant invasions from the New World to the Old World that has resulted in a number of taxonomic problems owing to the dearth of knowledge concerning the biosystematics of the genus. In this study, quantitative trait and enzyme electrophoresis analysis was used to assess the pattern of variation within and among five short-lived taxa of Conyza (C. blakei, C. bonariensis, C. canadenis, C. floribunda, C. sumatrensis), introduced into Europe, and to test the hypothesis of a recent hybrid origin of C. floribunda. The five taxa exhibit marked differences in morphological and life-history characteristics that are concordant with divergence in genes encoding isozymes. In addition, a recent hybrid origin of C. floribunda is clearly not supported by either quantitative or electrophoretic data. These findings suggest that the five taxa differentiated genetically in their native range prior to their introduction in Europe. As formerly suggested by Arthur Cronquist (Bulletin of the Torrey Botanical Club 70: 629–632, 1943), C. canadensis appears to be more closely related to the genus Erigeron than the other taxa. The electrophoretic results also confirm that this particular species is diploid, while the presence of multiple bands and fixed heterozygosity demonstrates that the other Conyza taxa are all genetic allopolyploids. Both quantitative and electrophoretic data suggest the presence of two groups among the polyploid taxa: one comprising C. bonariensis and C. sumatrensis, the other C. blakei and C. floribunda. The latter grouping is supported by the observation that C. blakei and C. floribunda, both native to Argentina, exhibit a striking ability to shift from semelparous to iteroparous reproduction, while the other taxa are strictly semelparous.     

HERITABILITY OF CARBON ISOTOPE DISCRIMINATION IN GUTIERREZIA MICROCEPHALA (ASTERACEAE)

The carbon isotope composition (¹³C/¹²C) of C₃ plant tissues provides a long-term, integrated measure of photosynthetic metabolism. Quantitative genetic methods were used to study the inheritance of carbon isotope composition and several morphological characters in Gutierrezia microcephala, a short-lived desert perennial. Open-pollinated seed was collected from a population located near Lee's Ferry, Arizona, in an area that was disturbed approximately 20 years ago. Seeds were germinated and seedlings grown in a common greenhouse environment. Carbon isotope discrimination (Δ) and all morphological characters varied significantly among maternal families. Heritability of Δ was estimated as 81% in the greenhouse and as 92% using the regression of family mean Δ on parent Δ from field samples. Using both field and greenhouse data, we estimated a lower bound for heritability in nature of 54%. Offspring size and biomass both showed significant, positive correlations with Δ. The corresponding genetic correlations were similar in magnitude and direction, but did not differ significantly from zero. These results imply the existence of heritable differences in physiology associated with carbon assimilation and water loss within populations, and the potential for microevolutionary change through natural selection.     

VARIATION IN GERMINATION RESPONSE TO TEMPERATURE IN RUBBER RABBITBRUSH (CHRYSOTHAMNUS NAUSEOSUS: ASTERACEAE) AND ITS ECOLOGICAL IMPLICATIONS

Seed collections from 72 rubber rabbitbrush populations occupying a range of habitats in western North America were incubated at 3 C in the laboratory. Collections from warm desert habitats required less than 2 weeks to achieve 90% relative germination under these conditions, while collections from montane habitats showed delayed germination requiring up to 20 weeks. When 13 representative collections were incubated at constant temperatures from 5 to 30 C, all germinated completely at 30 C within 4 weeks. Collections from warm desert habitats germinated rapidly over the whole range of temperatures. Montane collections sometimes exhibited dormancy at intermediate temperatures (15 and 25 C) even though they were ultimately able to germinate at lower temperatures. Results suggest that dormancy is conditional and temperature-dependent in this species. Chilling the seeds extends the temperature range for germination downward to include the chill temperature itself. Germination response to temperature and its variation as a function of habitat are of apparent adaptive significance, serving to time germination so that the probability of seedling survival is maximized in each habitat. Within populations, response to temperature varied as a function of year of harvest and of within-year harvest date, indicating that germination patterns are probably not under rigid genetic control but represent an integration of genetic and environmental factors.     

LEAF HAIRS IN ENCELIA (ASTERACEAE)

Four different kinds of leaf hairs occur in Encelia species. These are unicellular-based and multicellular-based uniseriate hairs, moniliform hairs, and biseriate glandular hairs. The unicellular-based uniseriate hairs appear responsible for increased leaf spectral reflectance by species within the genus. In particular, it appears that elongation of the distal cell of the uniseriate hair is necessary for increased leaf reflectance.     

A COMPARATIVE ANALYSIS OF MORPHOLOGICAL VARIATION PATTERNS IN THE PAPIONINS

Comparative studies of intraspecific variation patterns are important in attempts to reconstruct the differential selection pressures experienced by related species and in assessing the resultant observed interspecific variation. Reconstruction of past selection depends on an assumption of relatively stable patterns of genetic variance and covariance through time and across related species. Models by Lande (1979) and Turelli (1988a, 1988b) lead to contrasting expectations of stability versus lability of variation patterns, respectively, at least for closely related species. I report on a comparative study of phenotypic variance and correlation patterns in seven species of papionins, including macaques, baboons, and mangabeys, in order to determine the stability of variation patterns in this group. The three-dimensional coordinates of 12 bony landmarks on the face were used in a finite-element scaling analysis in order to measure local size variation at each landmark. Variances and correlations for these local size metrics were calculated using pooled sex-specific values. Variation patterns were compared across species using vector and matrix correlations in combination with various randomization-based significance tests. Patterns of variation and correlation were generally quite similar across the seven species, although some differences were also apparent. The overall magnitude of correlation is also similar among species, as only a few interspecific comparisons showed significant differences. Thus, it is concluded that patterns of variation and correlation in facial morphology have tended to remain stable in this group of primates. This result should allow for reconstruction of past differential selection pressures in the clade. The pattern of similarity among correlation structures for these species showed no association with their phylogenetic relationships.     

PATTERNS OF ISOZYME VARIATION IN WESTERN NORTH AMERICAN ANTENNARIA (ASTERACEAE: INULEAE) II. DIPLOID AND POLYPLOID SPECIES OF SECTION ALPINAE

Antennaria is a genus of dioecious, perennial, herbaceous Composites that are especially diverse in the cordillera of western North America. Section Alpinae consists of about nine taxa, among them A. aromatica, A. densifolia, A. media, A. pulchella, and A. umbrinella. Although diploids are morphologically distinct, the polyploid derivatives of the diploids obscure the morphological distinctness of the groups. A survey of 19 putative isozyme loci indicates that the diploids have diverged only moderately from one another with respect to biochemical genetics (I = 0.838 to 0.961). Additionally, only moderate amounts of genetic diversity were detected. Isozyme data are supportive of a hypothesis of a rapid mode of speciation in Antennaria, where morphological differentiation has been accompanied by small amounts of allozyme divergence. Polyploids have significantly higher amounts of heterozygosity than diploids and tetrasomic inheritance is inferred. Evidence from morphology and biochemical genetics suggests that the polyploids represent a continuum between interracial autopolyploids and segmental allopolyploids. In light of the relatively low degree of genetic and morphological divergence among many Antennaria species, taxonomic conservatism is advocated.     

植物区系的数值分析

本文介绍了应用数值分析的某些方法和原理进行植物区系基本成分统计、地理成分划分及其分区。植物区系数值分析不仅具有分析严密、计算简单及结果直观等优点,而且它比经典方法更能反映其本质特征。     

RESTRICTION ENDONUCLEASE ANALYSIS OF RIBOSOMAL DNA SEQUENCE VARIATION IN LAMINARIA (PHAEOPHYTA)1

The taxonomy and evolutionary relationships of species in the genus Laminaria are poorly understood. Previous studies have demonstrated significant plasticity of morphological characters used to describe taxa, and interfertility has been reported among putative species. We analyzed nuclear ribosomal DNA (rDNA) sequence variation in eight species of Laminaria (L. agardhii Kjell., L. digitata (Huds.) Lamour., L. groenlandica Rosenv. [sensu Druehl 1968], L. longicruris De la Pyl., L. longipes Bory, L. saccharina (L.) Lamour., L. setchellii Silva, and L. yezoensis Miyabe) to elucidate evolutionary relationships in this genus. Restriction maps were constructed using a small subunit rDNA probe from Costaria costata (Turn.) Saunders, an rDNA repeat from the nematode Caenorhabditis elegans, and 11 hexameric restriction endonucleases in an annealing analysis of genomic DNA. Laminaria rDNA restriction maps were compared to each other and to that of the outgroup taxon, C. costata. rDNA restriction maps of Laminaria species and C. costata were similar. Restriction fragment length polymorphisms mapped to both the coding regions and the nontranscribed spacer of rDNA. Laminaria species were distinguished with this method. The restriction maps of L. agardhii, L. saccharina, and L. longicruris were identical, supporting a previous hypothesis that these species are conspecific. Comparison of restriction maps of Laminaria species suggested that the generic subdivision of Sections Simplices and Digitatae may be invalid.     

中国鼠兔亚属(Subgenus Ochotona)种系发生的探讨

本文对鼠兔亚属主要产于国内的14种,进行了严格的支序系统分析。通过对这14种形态特征分析、对比,找出了22个具有种系发生意义的性状,并用国内学者尚未采用的格局分支论的组分分析,重建了它们的种间分支亲缘关系。根据分支图及国内已有的化石和地史资料,对该亚属的演化作了初步探讨。