EXPERIMENTAL HYBRIDIZATION OF XANTHIUM STRUMARIUM (COMPOSITAE) FROM ASIA AND AMERICA. I. RESPONSES OF F1 HYBRIDS TO PHOTOPERIOD AND TEMPERATURE

Responses to photoperiod and temperature were compared for hybrids between Asiatic plants in the indigenous strumarium morphological complex and plants in five American complexes: oviforme, italicum, chinense, cavanillesii, and pennsylvancium. The F₁ hybrids between Hong Kong plants and various American plants showed intermediacy in photoperiodic response; however, the hybrid night requirement was more similar to that of the American parent. The Hong Kong plants are difficult to evaluate photoperiodically but showed a night length requirement of 9.25–9.50 hr. Day-neutral plants from India in the strumarium complex produced day-neutral F₁ hybrids in crosses with most American plants having night requirements less than 10 hr. The F₁ hybrids involving the day-neutral Indian plants and either Indian or Australian chinense plants showed a night requirement of 8.25–8.75 hr. The chinense parental plants had apparent critical nights of 10.25–10.50 hr. Crosses between the day-neutral Indian plants and Mexican plants with apparent critical nights of 10.75–11 hr produced F₁ hybrids requiring nights of 8.75–9.00 hr. The various hybrids tended to show the broader temperature tolerances of the American parents. The ripeness-to-flower (maturity) responses of seedlings tended to show the genetic controls of the parent with the faster developmental rate. The hybridization evidence suggests that photoperiodic responses are quantitatively controlled and inherited independently of morphology and ripeness-to-flower responses. The populations of India are highly diverse and probably reflect recombinations of photoperiodic and temperature responses between indigenous day-neutral plants and photoperiodic chinense plants introduced from North America.     

VARIATION AMONG POPULATIONS OF XANTHIUM STRUMARIUM (COMPOSITAE) FROM NATURAL AND RUDERAL HABITATS

We assessed the mechanisms underlying the ability of cocklebur to spread from its natural riverside habitats and establish weedy populations in urban waste areas. We collected fruits from plants growing in natural and urban ruderal habitats and planted 2 maternal families from each of 9 habitat populations in 3 experimental gardens. The gardens were all in full sunlight but differed in the availability of water and nutrient resources. Plant performance in the gardens was measured by numbers and size of fruits produced. Traits known to be associated with cocklebur reproductive success were also measured: times to emergence and anthesis, photosynthetic capacity, mean stomatal conductance, relative growth rate, and biomass allocation to leaves and stems. Although there were significant differences among populations in the tradeoff between rate of growth in height and timing of anthesis, these population differences were not associated with habitat. Apart from a tendency to produce larger fruits in ruderal populations, there were no detectable differences in the characteristics of plants from natural vs. ruderal habitats. Plants from both habitats did have substantial and significant plastic responses to growth environment. In the three experimental gardens, fruit numbers increased with resource availability but fruit size did not differ significantly. As resource availability increased, plants from both habitats sustained growth longer and became more branchy. Canonical discriminant analysis of all the somatic and reproductive traits together supported the idea that natural and ruderal populations do differ in their overall plastic response to resource availability. The subtly different plastic responses of plants from the two habitats do not arise by substantial adjustments in a few dominant traits, but instead by relatively minor adjustments in a host of functionally interrelated phenological, morphological, and physiological traits. It is these small but coordinated differences in the plastic responses of many traits that appear to differentiate cocklebur from natural vs. weedy urban habitats.     

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.     

苍耳X anthium strumarium L.染色体数目和核型的观察

苍耳Xanthium strumarium L. (X. sibiricum Patr. ex Widder, X. iaponicum Widder) 为菊科苍耳属的一年生草本,广布于欧亚大陆和北美洲;我国各地也均有分布。苍耳为常用中药,其果实入药,能发汗通窍、祛风湿,解表镇痛,用于治风湿性关节痛及鼻炎等症。     

PHOTOPERIOD EVIDENCE IN THE INTRODUCTION OF XANTHIUM STRUMARIUM TO TAHITI AND HAWAII FROM MEXICO

Successful invasion of islands in the Pacific by Xanthium probably resulted from the chance introduction of a morpho-physiological form adapted to subtropical coastal habitats in North America. This hypothesis was tested by comparing naturalized plants from Tahiti in the Society Islands and Oahu in the Hawaiian Islands with collections from various coastal sites in Mexico. Plants from Tampico and Ciudad Mante, Tamaulipas; Veracruz (Antigua), Veracruz; and Salina Cruz, Oaxaca have morphological and physiological similarity to the Pacific introductions; however, each collection has certain niche relations that are slightly different from each other. All of the collections are in the same morphological complex and have apparent critical nights of 10.75–11 hr for reproduction. This morpho-physiological form is probably indigenous to Mexico, and it is widely distributed on the northeastern coastal plain of Tamaulipas.     

中国蒲公英属植物的初步整理

报导了我国蒲公英属植物的分类、分布及主要经济用途,也略讨论了该属植物的起源与区系地理等。     

INTERGENERIC HYBRIDIZATION,INDUCED POLYPLOIDY,AND THE ORIGIN OF THE HAWAIIAN ENDEMIC LIPOCHAETA FROM WEDELIA (COMPOSITAE)

The genetic relationships of the Hawaiian endemic genus Lipochaeta and the nearly cosmopolitan but extra-Hawaiian genus Wedelia were assessed by way of experimental hybridization. Hybrids between Wedelia biflora and diploid species of Lipochaeta consistently exhibited 15 pairs of chromosomes at meiosis, whereas the modal and maximum configuration seen in hybrids between W. biflora and tetraploid species of Lipochaeta was 15_II + 11_I. Meiotic pairing in microsporocytes of the intergeneric hybrid combinations involving Wedelia trilobata and both sections of Lipochaeta is lower than in intergeneric hybrids involving W. biflora. All of the intergeneric F, hybrids produced had relatively low pollen stainabilities ranging from less than 1–16% and, although they were vegetatively vigorous, they failed to produce viable achenes. The effects of chromosome doubling by colchicine in one species and several hybrids in Lipochaeta and in one intergeneric hybrid between Lipochaeta and Wedelia were studied. Chromosome doubling of the diploid species caused a decrease in pollen stainability and resulted in lack of achene production, whereas doubling the chromosomes of sterile intersectional and intergeneric hybrids restored female fertility and effected a dramatic increase in pollen stainability. Cytological analysis of the microsporocytes at diakinesis and metaphase I revealed pairs and quadrivalents in the induced polyploids. Pollen measurements indicated that pollen size in Lipochaeta is correlated with doubling of the chromosomes. Statistical analyses revealed that there is no significant difference in pollen size between the diploids and natural tetraploids, whereas both groups have pollen grains that are significantly smaller than the induced autotetraploid, the intergeneric allotetraploid, and the intersectional allohexaploid. The cytogenetic evidence indicates a relatively close genetic relationship between Wedelia and Lipochaeta and supports the view that the ancestry of the diploid section Aphanopappus of Lipochaeta (n = 15) is to be found in a species akin to Wedelia biflora (n = 15) and that this or a similar 15-paired wedelioid species and an unknown 11-paired wedelioid taxon are involved in the alloploid hybrid origin of the tetraploid section Lipochaeta (n = 26) of Lipochaeta.     

CHROMOSOME ANALYSIS OF HYBRIDS BETWEEN 4N GRINDELIA CAMPORUM AND 2N G. GRANDIFLORA (COMPOSITAE)

Morphology and meiosis are described in four progeny plants resulting from tetraploid Grindelia camporum Greene (2n = 24) from California pollinated by diploid G. grandiflora Hook. (2n = 12) from Coahuila, Mexico. Three of the four progeny were tetraploid, morphologically like the pistillate parent, and had metaphase I chromosome configurations which included quadrivalents and a complementary number of bivalents. They are considered to have resulted from selfing. The fourth plant was triploid (2n = 18) andmorphologically intermediate between the parents. Chromosome configurations in the triploid were variable with univalents, ring and rod bivalents, trivalents and pentavalents. These two species are considered related through an ancestor with a basic genome, but are separated cytologically by polyploidy and by two distinct chromosomal interchanges that explain the configurations observed in the triploid hybrid.     

CYTOGENETICS OF HYBRIDS OF CARTHAMUS SPECIES (COMPOSITAE) WITH TEN PAIRS OF CHROMOSOMES

Plants thought to be typical of 7 species were chosen to represent the various taxa of Carthamus species with 10 pairs of chromosomes. These entities were crossed in all possible combinations and 20 of the possible 21 interspecific hybrids were obtained after 3 seasons of crosses. Analyses of the hybrids included studies of microsporocytes, pollen stainability, achene fertility, rudimentary ovaries, and other morphological characteristics. Pairing of chromosomes at metaphase I indicated no translocations were present in hybrids between C. tenuis from Israel, C. alexandrinus from Egypt, C. glaucus from northern Israel, and C. syriacus from Jordan. Members of this group are assigned the standard arrangement. Hybrids of C. glaucus from Iran, C. glaucus from Syria, and C. dentatus from Turkey always showed a translocation or chromosomal interchange when crossed with any member having the “standard” arrangement. The last 3 species are considered to have the “non-standard” chromosomal arrangement. The parental species used in this study can be regarded as a set of testers which will allow identification of chromosomal differentiation in additional Carthamus materials as they are collected.     

横断山脉地区蒲公英属植物资料

对我国横断山脉地区的蒲公英属植物进行了总结,收载了该地区产的18种蒲公英属植物。     

PHYLOGEOGRAPHY OF BATRACHOSPERMUM MACROSPORUM (BATRACHOSPERMALES,RHODOPHYTA) FROM NORTH AND SOUTH AMERICA1

Phylogeographic trends in Batrachospermum macrosporum Mont. were investigated using the mitochondrial intergenic spacer between the cytochrome oxidase subunit 2 and 3 genes (cox2‐3). A total of 11 stream segments were sampled with seven in the coastal plain of North America and four in tropical areas of South America. Fifteen thalli were sampled from seven streams, 14 thalli from two streams, and eight thalli from two streams. There were 16 haplotypes detected using 149 individuals. Of the eight haplotypes from locations in North America, all were 334 base pairs (bp) in length, and of those from South America, five were 344 bp, and three were 348 bp. Two individual networks were produced: one for the haplotypes from North America and another for those from South America, and these could not be joined due to the large number of base pair differences. This split between haplotypes from North and South America was confirmed with sequence data of the rbcL gene. There was very little genetic variation among the haplotypes from the North American locations, leading us to hypothesize that these are fairly recent colonization events along the coastal plain. In contrast, there was high variation among haplotypes from South America, and it would appear that the Amazon serves as a center of diversity. We detected considerable variation in haplotypes among streams, but frequently, a single haplotype in an individual stream segment, which is consistent with data from previous studies of other batrachospermalean taxa, may suggest a single colonization event per stream.     

CHARACTERIZATION OF POLLEN ANTIGENS FROM AMBROSIA L. (COMPOSITAE) AND RELATED TAXA BY IMMUNOELECTROPHORESIS AND RADIAL IMMUNODIFFUSION

The pollen antigens of various Ambrosia and related species were studied to learn whether substances closely related to antigen E (the major allergen of Ambrosia artemisiifolia) were present. After conventional immunoelectrophoresis, pollen extracts from six Ambrosia species each produced at least one pronounced precipitin line with antiserum for purified antigen E. Electrophoretic mobility was the same for several species (A. artemisiifolia, A. bidentata, A. psilostachya, and A. trifida) but was relatively lower for A. acanthicarpa and A. ambrosioides. Precipitin rings were also produced when pollen extracts of the various Ambrosia species were subjected to radial immunodiffusion in agarose which contained antiserum for purified antigen E. There was great variation among the Ambrosia species with respect to precipitin ring diameters. The variation may be due to differences among species in content of the antigen E-like substances or to altered interaction with the immobilized antibody. Crossed (2-dimensional) immunoelectrophoresis was shown to be useful for characterizing Ambrosia pollen antigens. Pollen extracts from A. artemisiifolia produced eight pronounced precipitin bands and at least eight faint, relatively fast-moving bands after crossed immunoelectrophoresis with antiserum against a whole pollen extract from the same species. One of the pronounced bands contained antigen E.     

东亚和北美当归属（广义）的果实解剖和演化

应用比较解剖研究了广义当归属在世界主要分布区－东亚和北美种类的果实结构的特征。根据分生果背腹压扁的程度、棱翅与槽的形状、果皮层次和结构、油管数目、大小和位置等稳定性状,可将果实分为５个类型,与《中国植物志》将其分为５个属,即古当归属,当归属,高山芹属,柳叶芹属和山芹属的意见相一致。古当归属为原始的类型,山芹属具有高度演化的特征。且对一些种类有分类位置提出了调整的意见和各属的演化趋势。     

HYBRIDS AND PHYLOGENETIC SYSTEMATICS II. THE IMPACT OF HYBRIDS ON CLADISTIC ANALYSIS

I examined three aspects of the cladistic treatment of a set of 17 F₁ hybrids of known parental origin: (1) impact of hybrids on consistency index (CI) and number of most parsimonious trees (Trees), (2) placement of hybrids in cladograms, and (3) impact of hybrids on hypotheses of relationship among species. The hybrids were added singly and in randomly selected sets of two to five to a data set composed of Central American species of Aphelandra (including the parents of all hybrids). Compared to analyses with the same number of OTUs all of which were species, the analyses with hybrids yielded results with significantly higher CI. There was no difference in Trees between analyses with hybrids versus species. There was thus no evidence that hybrids would appear to be more problematic for cladistic methods than species. Accordingly, hybrids will not be readily identifiable as taxa that cause marked change in these indices. About % of the hybrids were placed as the cladistically basal members of the lineage that included the most apomorphic parent. Relatively apomorphic hybrids were placed proximate to the most derived parent (ca. 13% of hybrids). Other placements occurred more rarely. The most frequent placements of hybrids thus did not distinguish them from normal intermediate or apomorphic taxa. When analyses with hybrids yielded multiple most parsimonious trees, these were no more different from each other than were the equally parsimonious trees that resulted from analyses with species. Most analyses with one or two hybrids resulted in minor or no change in topology. When hybrids caused topological change, they frequently caused rearrangements of weakly supported portions of the cladogram that did not include their parents. When they disrupted the cladistic placement of their parents, they often caused their parents to change positions, with at least one topology bringing the parental lineages into closer proximity with the hybrid placed between them. Hybrids between parents from the two main lineages of the group caused total cladistic restructuring. In fact, the degree of relationship between a hybrid's parents (measured by both cladistic and patristic distance) was strongly correlated with CI (negatively) and with the degree of disturbance to cladistic relationships (positively). Thus, hybrids between distantly related parents resulted in cladograms with low CI and major topological changes. This study suggests that hybrids are unlikely to cause breakdown of cladistic structure unless they are between distantly related parents. However, these results also indicate that cladistics may not be specially useful in distinguishing hybrids from normal taxa. The applicability of these results to other kinds of hybrids is examined and the likely cladistic treatment of hybrids using other sources of data is discussed.     

NORTHERN AND SOUTHERN HEMISPHERE HYBRIDS OF MACROCYSTIS (PHAEOPHYCEAE)1

All combinations of individuals of Macrocystis pyrifera (L.) C. Agardh, M. integrifolia Bory, and M. angustifolia Bory hybridized. Gametophyte isolates obtained from 18 individuals were used, including M. pyrifera and M. integrifolia from the extremes of their Northern Hemisphere ranges along the Pacific Coast of North America and M. pyrifera and M. angustifolia from Tasmania, Australia. All combinations of gametophytes produced sporophytes of normal morphology, with the exception of crosses involving three gametophyte isolates. One female (M. integrifolia) and two male (M. pyrifera and M. angustifolia)gametophyte isolates were unable to produce normal sporophytes in combination with gametophytes of the opposite sex. Some cultures of female gametophytes produced abnormally shaped parthenogenetic sporophytes. Gametophytes and sporangia of M. pyrifera had n= 16 chromosomes. The M. integrifolia female gametophyte that was unable to produce normal sporophytes had n = ca. 32 chromosomes. These results show that these species of Macrocystis have not become reproductively isolated. Although these species may be considered conspecific according to the biological species concept, we recommend that they continue to be recognized as separate species based on morphological differences.     

MORPHOMETRIC ANALYSIS OF POLLEN FROM FOUR SPECIES OF AMBROSIA (COMPOSITAE)

A morphometric approach proved useful for characterizing pollen from the Ambrosia species that are responsible for most ragweed hayfever and for correctly identifying individual pollen grains. Pollen was harvested from 4 species (A. trifida, A. artemisiifolia, A. bidentata, A. psilostachya) grown in Champaign County, Illinois, and observed with a scanning electron microscope. Quantitative measurements of 6 different exine characters were made on micrographs of 96 pollen grains from each species. Discriminant analysis based on these characters was used to obtain a population centroid for each species and indicated that these centroids were distinctly different. This analysis also included a species classification for individual pollen grains and indicated probability of species membership. About 86% of all individuals were correctly classified when the 4 species were included, with a range from 80% (A. artemisiifolia) to 95% (A. psilostachya). When three species (A. trifida, A. artemisiifolia, A. bidentata) were analyzed, about 87% of the individuals were correctly classified, and almost all individuals (97%) were classified correctly when only A. trifida and A. artemisiifolia were included. The validity of the derived discriminant functions was also tested by treating herbarium specimens as unknowns. The species and percentages of these individuals correctly classified were A. trifida (100%), A. artemisiifolia (58%), A. bidentata (100%), and A. psilostachya (31%). The classification results using three species were A. trifida (100%), A. artemisiifolia (75%), and A. bidentata (100%); and using two species were A. trifida (100%) and A. artemisiifolia (92%). Hence, the disciminant functions do seem valid for 3 species, but further investigation of A. psilostachya is needed. This type of approach may be useful in pollen analysis, especially where these species significantly contributed to aeroallergens.