DYNAMICS OF POLYPLOID FORMATION IN TRAGOPOGON (ASTERACEAE): RECURRENT FORMATION,GENE FLOW,AND POPULATION STRUCTURE

Polyploidy is a major feature of angiosperm evolution and diversification. Most polyploid species have formed multiple times, yet we know little about the genetic consequences of recurrent formations. Among the clearest examples of recurrent polyploidy are Tragopogon mirus and T. miscellus (Asteraceae), each of which has formed repeatedly in the last ～80 years from known diploid progenitors in western North America. Here, we apply progenitor‐specific microsatellite markers to examine the genetic contributions to each tetraploid species and to assess gene flow among populations of independent formation. These data provide fine‐scale resolution of independent origins for both polyploid species. Importantly, multiple origins have resulted in considerable genetic variation within both polyploid species; however, the patterns of variation detected in the polyploids contrast with those observed in extant populations of the diploid progenitors. The genotypes detected in the two polyploid species appear to represent a snapshot of historical population structure in the diploid progenitors, rather than modern diploid genotypes. Our data also indicate a lack of gene flow among polyploid plants of independent origin, even when they co‐occur, suggesting potential reproductive barriers among separate lineages in both polyploid species.     

A CYTOGENETIC ANALYSIS OF CERTAIN POLYPLOIDS IN GRINDELIA (COMPOSITAE)

Two diploid taxa, Grindelia procera and G. camporum, and 3 tetraploid ones, G. camporum, G. hirsutula, and G. stricta, have been studied to ascertain their interrelationships. Meiosis in diploid parental strains was regular, the common chromosome configuration being 5 rod bivalents and 1 ring bivalent. The average chiasmata frequency per chromosome was 0.60. Pollen fertility was about 90% in all strains examined. Diploid interspecific hybrids had normal meiosis with an average chiasmata frequency of 0.56 per chromosome. No heterozygosity for inversions or interchanges was detected, and pollen fertility was above 85%. Meiosis in parental tetraploid strains was characterized by the presence of quadrivalents in addition to a complementary number of bivalents. The average chiasmata frequency per chromosome was 0.59 and pollen fertility was generally about 80%. Tetraploid interspecific hybrids also had quadrivalents, normal meiosis, and high pollen fertility. Close genetic relationships between the diploids and between the tetraploids are indicated, and geographical, ecological, and seasonal barriers to gene exchange exist. Attempts to obtain hybrids between diploids and tetraploids were successful in a few cases. The hybrids were tetraploid and had normal meiosis and fertility similar to parental and F₁ tetraploids. Their origin was by the union of unreduced gametes of the diploid female parent and normal pollen from the tetraploid parent. On the basis of chromosome homology, normal meiosis, plus high fertility exhibited in the diploid, tetraploid, and diploid X tetraploid interspecific hybrids, these species of Grindelia are considered to be a part of an autopolyploid complex. Gene exchange between diploids and diploids, tetraploids and tetraploids, and diploids and tetraploids is possible. Tetraploid G. camporum may have originated by hybridization between G. procera and diploid G. camporum with subsequent doubling of chromosomes and selection for the combined characteristics of the diploids.     

DEVELOPMENTAL AND GENETIC SOURCES OF SEED WEIGHT VARIATION IN RAPHANUS RAPHANISTRUM L. (BRASSICACEAE)

Seed weight is known to have a marked impact on emergence and post-emergence productivity in wild radish (Raphanus raphanistrum). In this paper, I describe several levels of seed weight variation in plants taken from a natural population in Hamden, Connecticut. Six maternal plants from the 1981 season were analyzed in detail: the weights and positions of all seeds within a fruit were recorded, and some of these seeds were used the following summer for competition studies and progeny analysis. Within a plant, average seed weight decreased as the number of seeds within a fruit increased, suggesting that developing embryos compete for maternal resources. Seed weight also varied significantly among the six maternal plants used in the study. Comparison of the average weights of seeds produced by offspring of those six plants with the average weights of seeds borne by the maternal plant revealed a significant genetic component to seed weight variation. Seed weight varied up to six-fold within single fruits of R. raphanistrum; large seeds tend to occur near the pedicel or in the middle positions. Seed size variation seen within single fruits is of sufficient magnitude to result in differential reproductive output among closely related seeds under competitive field conditions.     

EFFECTS OF APEX REMOVAL AND NUTRIENT SUPPLEMENTATION ON BRANCHING AND SEED PRODUCTION IN THLASPI ARVENSE (BRASSICACEAE)

The shoot apex of a plant may be damaged by herbivory, weather conditions, or other factors, leading to growth of some axillary buds into branches. This alteration of branching pattern can affect the location and extent of seed production. I examined the effects of removal of the shoot apex and of mineral nutrient addition on branching and seed production in Thlaspi arvense. Individual plants received no, early, or late apex removal and no, early, or late nutrient addition in a two-way factorial experiment. Apex removal led to greater production of secondary branches. Total seed weight per plant was higher for early- than late-removal plants under all nutrient treatments, but intact controls produced the highest total weight of seeds in two of three nutrient treatments, because their seeds were heavier. Addition of nutrients increased the numbers of secondary branches, fruits, and seeds, and the average and total weight of seeds produced. Apex removal resulted in increased seed production only when it occurred early and when nutrients were added at the same time. Resource availability and plant phenology can be influential in determining the effects of apical damage on seed production, and their effects in combination may differ from their individual effects.     

EFFECTS OF PHOTOPERIOD AND VERNALIZATION ON THE NUMBER OF LEAVES AT FLOWERING IN 32 ARABIDOPSIS THALIANA (BRASSICACEAE) ECOTYPES

The extent of natural variation among wild type Arabidopsis thaliana L. Heynh for response to environmental stimuli that affect flowering is poorly documented. The effects of photoperiod and vernalization on the number of rosette leaves at the time of anthesis was measured for 32 Arabidopsis ecotypes. All ecotypes were vernalized 24 days at 4 C under continuous illumination. Vernalized and nonvernalized plants were transplanted into 8- (short-day) and 20-hour (long-day) photoperiods in controlled environment growth chambers. Two ecotypes failed to flower after 110 days. Mean leaf number was less for all ecotypes under long day compared to short day. Sixteen ecotypes responded to vernalization; eight had reduced leaf number regardless of photoperiod, and eight had reduced leaf number only under short day. Two ecotypes required vernalization to flower in this study. For three ecotypes, vernalization and short day resulted in a reduction in leaf number whereas vernalization and long day resulted in increased leaf number. Documentation of the effects and interactions of photoperiod and vernalization across many ecotypes provides a broader range of described natural variation for genetic and physiologic study.     

ECOLOGICAL CONSEQUENCES AND PHENOTYPIC CORRELATES OF PETAL SIZE VARIATION IN WILD RADISH,RAPHANUS SATIVUS (BRASSICACEAE)

In this paper we examine some ecological consequences and phenotypic correlates of flower size variation in wild radish, Raphanus sativus. Mean corolla diameter varied significantly among individuals within natural populations of R. sativus in California. On the average, almost 40% of flower biomass was allocated to corolla tissue. In field experiments, pollinator visitation increased significantly with corolla size. Large flowers also accumulated more nectar when pollinators were excluded from plants. In three populations, corolla size was positively correlated with allocation to pollen per flower (either anther weight or pollen grain number), but there was usually no phenotypic relationship between corolla size and several measures of female allocation (ovule number per flower, proportion fruit set, and total seed mass per fruit). Plants growing in the field produced fewer large flowers per unit of stem, and stem biomass was negatively related to corolla size for plants grown under controlled greenhouse conditions. Male and female fitness may covary differently with allocation to attractive floral features in species such as R. sativus, where seed production is often limited by resources rather than by pollen.     

ALLYL ISOTHIOCYANATE RELEASE AND THE ALLELOPATHIC POTENTIAL OF BRASSICA NAPUS (BRASSICACEAE)

The allelopathic potential of Brassica species has been attributed to release of the mustard oil glycosides which they produce in large quantities. Upon hydrolysis, these glucosinolates yield isothiocyanates, compounds with strong antibiotic properties. The objective of this study was to assess whether Brassica napus, a common and widespread crop and weed crucifer, is capable of allelopathic interference, and if so, whether its glycoside derivative, allyl isothiocyanate (AI), is capable of producing this interference. Wild type and low glucosinolate-mutant B. napus were grown in low organic content soil under greenhouse conditions, and AI release into soil was monitored. Most plants released low levels of AI, though approximately 10% released much higher levels. Wild type plants released more AI than mutants. Growth of the target species, Medicago sativa, was not affected by additions of AI to soils at concentrations equal to the median and 95% quantile from the B. napus soils. In replacement series experiments, the two B. napus genotypes suppressed growth of M. sativa equally despite differences in AI release rate. In an intraspecific replacement series experiment, the two B. napus genotypes were equal competitors. Under our experimental conditions, B. napus showed no indication of being allelopathic, and AI concentrations typical of soils around B. napus plants did not inhibit target plants.     

POLYPHYLY OF THE BLACKBIRD GENUS AGELAIUS AND THE IMPORTANCE OF ASSUMPTIONS OF MONOPHYLY IN COMPARATIVE STUDIES

A phylogeny for Agelaius blackbirds was constructed using sequence data from an 890 base-pair (bp) region of the mitochondrial cytochrome-b gene in nine species of Agelaius and a single species from all but 1 of the 28 described blackbird genera and subgenera. The genus was found to be polyphyletic with the South American members of Agelaius more closely related to other South American blackbird genera. Application of bootstrap and jackknife manipulations supports this conclusion. That this relatively well-known genus is polyphyletic represents a warning to those attempting to construct phylogenies without first demonstrating monophyly of the ingroup. The conclusion that Agelaius is polyphyletic necessitates (1) the reinterpretation of previous studies that assumed monophyly and (2) the initiation of a variety of new comparative behavioral and ecological studies suggested by this finding.     

TRANSMISSION GENETICS OF ISOZYME LOCI IN RAPHANUS SATIVUS (BRASSICACEAE): STRESS-DEPENDENT NON-MENDELIAN SEGREGATION

Deviations from Mendelian ratios are frequently treated as an intrinsic property of individuals, independent of the environment. We tested whether the environment of the parents could alter patterns of inheritance in the wild radish, Raphanus sativus. We demonstrated the genetic basis of 12 isozyme loci by controlled pollinations of unstressed plants. The frequency of deviant segregation detected was not different than that expected by chance. Controlled pollinations among stressed plants showed over 3 times as much deviant segregation as the unstressed controls. No genetic correlates of segregation bias were detected. Linkage was assessed for 64 of the 66 pairs of loci. Two linkage groups were detected, one involving four loci (PGM2–ACO–ACP–LAP), the other involving a single pair (EST-PRX). The second linkage group is apparently associated with a locus or tightly linked loci which may segregate for “balanced” lethals on the same chromosome. Deviant segregation did not appear to act primarily by selection on a particular gamete. Postzygotic selection was the probable source of at least some of the aberrant segregation. Because no particular allele was favored in such situations, selection is apparently operating on alleles at linked loci rather than on the allozyme loci per se. Data from other studies on wild radish support the suggestion that postzygotic selection might be an important influence on progeny segregation ratios. Because wild radishes often encounter a variety of stresses in the field, in this species, aberrant segregation may be common under natural conditions.     

RELATIVE SALT TOLERANCE OF CAKILE EDENTULA (BRASSICACEAE) FROM LACUSTRINE AND MARINE BEACHES

Seeds of Cakile edentula ssp. edentula var. edentula (hereafter edentula) were collected from the Pacific Ocean shore of Washington, and Cakile edentula ssp. edentula var. lacustris (hereafter lacustris) seeds were collected from the Lake Huron shore of Ontario. Germination rates of the two varieties showed equal tolerance of substrate salinities ranging from 0–10,000 ppm. Early root growth of edentula seedlings was significantly stimulated by 1,000 ppm salts, but otherwise both varieties showed equal tolerance to 10,000 ppm. Edentula plants exposed for 4 wk to salt spray intensities of 0, 20, and 90 mg dm^–2 d^–1, applied in a growth chamber environment, showed no significant changes in shoot or root biomass, plant morphology, or reproductive effort. Lacustris plants exhibited significant declines in stem length and reproductive effort with increasing levels of salt spray. Our results suggest that lacustris and edentula have diverged physiologically over the past 9,000 years, and field observations from the literature corroborate our conclusion that this divergence in salt tolerance has ecological significance.     

EMBRYO SAC LACKING ANTIPODAL CELLS IN ARABIDOPSIS THALIANA (BRASSICACEAE)

Ultrastructure of the embryo sac lacking antipodals in prefertilization stages in Arabidopsis thaliana has been examined 2 hr before and 5 hr after manual cross pollination. The cytoplasm of both synergids before fertilization is rich in ribosomes, mitochondria, and rough endoplasmic reticulum, and also contains several microbodies and spherosomes. The filiform apparatus includes electron-dense material and a fibrous part. Many cortical microtubules appear in the filiform apparatus area. One of the two synergids degenerates before fertilization. The synergids, the egg cell, and central cell have a rich cytoskeleton of microtubules; only the synergids appear to contain microfilaments. At the chalazal end, the antipodals are initially present but degenerate by the time of pollination in most embryo sacs in the starchless line studied. The embryo sac is completely surrounded by a wall containing an electron-dense layer, separating it from the nucellus, including the chalazal end. When the antipodals have degenerated, the electron-dense layer disappears at the chalazal end only, and the wall between the central cell and the nucellus is homogeneous. Between the central cell and nucellar cells no plasmodesmata are found. The membranes of both antipodal cells at the chalazal end of the embryo sac appear sinuous, like those of transfer cells. The central cell has plastids preferentially distributed around the nucleus, but the other organelles are randomly distributed. The central cell in the embryo sac and the adjacent chalazal nucellar cells show a transfer-cell function in the embryo sac after the antipodals degenerate.     

THE INTRAFLORAL PHENOLOGY OF STREPTANTHUS TORTUOSUS (BRASSICACEAE)

The sequence of floral events during anthesis was examined in Streptanthus tortuosus to determine the relationship between the male and female floral phases. The flowers are strongly protandrous. In the staminate phase, the anthers mature sequentially over a 3–4-day period. Because pollinators quickly remove pollen from the anthers, sequential anther maturation prolongs the male phase relative to what it would be if anthers did not mature sequentially. Pollen applied to the stigma during the staminate phase does not adhere readily and does not germinate. The length of the pistillate phase depends on pollinator activity, as pollination accelerates the abscission of floral parts. Unpollinated flowers remain pistillate for 3–4 days, during which time stigmatic receptivity declines gradually. In the field, 72% to 80% of flowers are staminate at any time, indicating that the staminate phase is three times longer than the pistillate phase when pollinators have access to the flowers. The consequences of the relative length of the floral phases and the schedule of stigmatic receptivity are discussed in terms of outcrossing mechanism, floral longevity, and sexual selection models.     

REVISION OF THE MASTOPHOROIDEAE (CORALLINALES,RHODOPHYTA) AND POLYPHYLY IN NONGENICULATE SPECIES WIDELY DISTRIBUTED ON PACIFIC CORAL REEFS1

The subfamily Mastophoroideae (Corallinaceae, Rhodophyta) is characterized by species possessing nongeniculate, uniporate tetrasporangial conceptacles without apical plugs, the presence of cell fusions, and the absence of secondary pit connections. However, molecular phylogenetic studies not including the type genus Mastophora indicated that the Mastophoroideae was polyphyletic. Our molecular phylogenetic analysis of the subfamily including the type genus using DNA sequences of SSU rDNA and plastid‐encoded gene of PSII reaction center protein D1 (psbA) revealed that Mastophora formed a robust clade only with Metamastophora. The other mastophoroid genera were divided into six lineages within the family Corallinaceae. Five supported lineages—(i) Pneophyllum; (ii) Hydrolithon gardineri (Foslie) Verheij et Prud’homme, Hydrolithon onkodes (Heydr.) Penrose et Woelk., and Hydrolithon pachydermum (Foslie) J. C. Bailey, J. E. Gabel et Freshwater; (iii) Hydrolithon reinboldii (Weber Bosse et Foslie) Foslie; (iv) Spongites; and (v) Neogoniolithon—were clearly distinguished by the combination of characters including the presence or absence of palisade cells and trichocytes in large, tightly packed horizontal fields and features of tetrasporangial and spermatangial conceptacles. Therefore, we amend the Mastophoroideae to be limited to Mastophora and Metamastophora with a thin thallus with basal filaments comprised of palisade cells, tetrasporangial conceptacles formed by filaments peripheral to fertile areas, and spermatangia derived only from the floor of male conceptacles. This emendation supports Setchell’s (1943) original definition of the Mastophoroideae as having thin thalli. We also propose the establishment of three new subfamilies, Hydrolithoideae subfam. nov. including Hydrolithon, Porolithoideae subfam. nov. including the resurrected genus Porolithon, and Neogoniolithoideae subfam. nov. including Neogoniolithon. Taxonomic revisions of Pneophyllum and Spongites were not made because we did not examine their type species.     

THE LIGHT FACTOR IN THE GERMINATION ECOLOGY OF DRABA VERNA

The majority of the seeds of the winter annual Draba verna L. require light for germination, but light can be given before they are fully after-ripened and as long as 4 months before temperature and moisture conditions are favorable for germination. Seeds that are exposed to light in late spring, or in late spring and early summer, and then removed to darkness can germinate under favorable temperature and moisture conditions in autumn, even after passing through a long unfavorable (for germination) wetting and drying period at high summer temperatures in darkness. The light requirement for germination probably is not an important factor restricting D. verna to open, well-lighted habitats.     

24种十字花科短命植物的扩散体特征与扩散对策

对准噶尔荒漠中24种十字花科短命植物的扩散体特征与扩散对策的观测结果表明:种间果实和种子的形态各异,角果开裂与否与其木质化程度有关。木质化程度较高的角果不开裂。其中长角果以果节为扩散单元自体扩散。而短角果有两种扩散单元,以果实为扩散单元的无辅助扩散结构或具翅或绵毛,分别进行自体或风媒扩散;以果序的一段为扩散单元的果实具坚硬呈勾状的喙,通过果序梗断裂或附在动物体表散布,兼具自体扩散和动物体外传播的二重性。木质化程度较低的角果开裂,以种子为单元扩散。其中微尘状种子无辅助扩散结构,在果皮开裂时产生的张力或风力的作用下散布;粘液种子以水媒为主,兼具风媒和动物传播的特点;粘液和翅并存的种子具有风媒和水媒扩散的二重性。果翅等辅助扩散结构有利于长距离散布,绵毛及种皮粘液使果实或种子与地表能更好地粘附,木质化结构及种皮粘液还具有保水和保护作用。从扩散的时空特征看,24种植物在扩散持续时间上有集中扩散和持续扩散两种方式,前者避免了种子干旱失水和被捕食,属逃逸对策;后者形成空气种子库逐批扩散,属保护对策。从空间上,异果芥(Diptychocarpus strictus)果序上、下部果实的开裂方式与时间、种子形态完全不同,导致其扩散时间与距离不同;离子芥(Chorispora tenella)等类群果实不同部位的果节非同步脱落以及爪花芥(Oreoloma sulfureum)果实上、下部木质化程度及开裂方式不同,导致其果节或种子的扩散时间不同。以上这些特征对于它们在荒漠环境中成功地扩散、定居,抵御干旱、逃避捕食,避免同胞种子间的竞争、保障物种延续并扩大种群等,具有重要的生态学意义。     

THE EFFECT OF GENETICALLY BASED DIFFERENCES IN SEED SIZE ON SEEDLING SURVIVAL IN ARABIDOPSIS THALIANA (BRASSICACEAE)

The relationship between genetically different seed sizes and seedling survival under severe nutrient deprivation was determined by comparing ten inbred lines of Arabidopsis thaliana. Seedlings were supplied with only sterile distilled water, and the number of days between germination and death (complete chlorosis) was recorded. Seedlings from genotypes with larger seeds survived longer than seedlings from genotypes with smaller seeds. These results suggest a genetically based adaptive significance of larger seed size resulting from a greater seedling tolerance of nutrient deprivation. This may confer a potentially important selective advantage when nutrient deprivation is the result of a low resource supply in the environment, or the result of nutrient depletion by neighbors.     

STRUCTURE AND FORMATION OF THE BYSSUS COMPLEX IN MYTILUS (MOLLUSCA, BIVALVIA)

A review is presented of the mechanism of byssus productionin the genus Mytilus. The pedal glands which secrete the byssusmaterial are described, followed by an account of the morphology,structure and chemistry of the byssus itself. Finally, the releaseor shedding of the byssus from the pedal tissues is discussed. (Received 10 November 1981;     

DNA SEQUENCE DATA DEMONSTRATE THE POLYPHYLY OF THE GENUS CYSTOSEIRA AND OTHER SARGASSACEAE GENERA (PHAEOPHYCEAE)1

Phylogenetic relationships in the Sargassaceae were explored using three DNA markers, and the monophyly of its genera was challenged. Nineteen out of 24 currently recognized genera were sampled, representing 63 species. The variable mt23S‐tRNA Val intergenic spacer could only be aligned within genera and could not be used to infer intergeneric relationships. The partial mt23S was also useful to delineate genera and was alignable at the family level but provided few informative characters. Analysis of mt23S DNA sequences together with chloroplast‐encoded psbA sequences resulted in a better resolved phylogeny. Hormophysa was the first genus to branch off within the Sargassaceae, followed by Myriodesma; then the three genera Caulocystis, Carpoglossum, and Scaberia in unresolved order; and then Acrocarpia. The other taxa studied here were divided over three major clades, but there was no branch support for the monophyly of two of these. The genera Bifurcaria, Cystoseira, Halidrys, and Sargassum appeared polyphyletic. The following taxonomic changes are proposed: a new genus Brassicophycus for Bifurcaria brassicaeformis (Kützing) E. S. Barton; reinstatement of the genus Sargassopsis for Sargassum decurrens (R. Brown ex Turner) C. Agardh; reinstatement of the genus Sirophysalis for Indo‐Pacific Cystoseira trinodis (Forsskål) C. Agardh; reinstatement of the genus Polycladia for the western Indian Ocean species Cystoseira indica (Thivy et Doshi) Mairh, Cystoseira myrica (S. G. Gmelin) C. Agardh, and Acystis heinii Schiffner; and reinstatement of the genus Stephanocystis for the North Pacific Cystoseira species and Halidrys dioica N. L. Gardner. The European Cystoseira species should be split into three genera, but no name changes are proposed yet, because diagnostic characters were found only for the clade including the type species. Some evolutionary trends could be discerned from the mt23S + psbA phylogeny.     

群心菜花蜜腺的发育解剖学研究

群心菜（Ｃａｒｄａｒｉａｄｒａｂａ（Ｌ．）Ｄｅｓｖ）花蜜腺６枚,包括４枚侧蜜腺的和２枚中蜜腺,属十字花科侧中蜜腺类型中的侧分离中间亚型,侧中蜜腺结构相同,都由分泌表皮,产蜜组织组成,分泌表皮顶部分布的有变态气孔器,产蜜组织中无维管束分布,属较原始的十字花科花蜜腺亚型类型,在花的各部分基本分化完成后,由花托表层细胞恢复分裂能力形成蜜腺原基,蜜腺原基经分裂,分化和形态建成,发育形成成熟蜜腺,侧中蜜腺发