A unique pollen wall mutation in the family Compositae: ultrastructure and genetics

During a routine screening of pollen fertility in the n = 2 chromosome race of Haplopappus gracilis, a spineless pollen wall mutation was discovered that renders the otherwise functional pollen grains completely unrecognizable as Compositae pollen. Normal Haplopappus pollen is characterized by an outer layer, the ektexine, consisting of large spines supported by a roof (tectum), which in turn is supported by collumellae that are joined basally. A large cavity (cavea) stretches from aperture to aperture and separates columellae bases from the final ektexine unit, the foot layer. The spines, tectum, columellae, and columellae bases are filled with perforations (internal foramina), while the foot layer is without them. Immediately underlying the foot layer is a thickened, lamellate, disrupted, internal foramina-free second exine layer, the endexine. In contrast, the mutant pollen ektexine is a jumble of components with randomly dispersed spines as the only clearly definable unit. The endexine layer is similar to the endexine in normal pollen. The mutation apparently disrupts only the organization of ektexine units, and mutant pollen appears to be without the caveae and foot layer characteristic of normal pollen. In genetic tests, the mutant allele is recessive. There is a simple Mendelian pattern of inheritance of the mutant gene, and its phenotype is under sporophytic control.     

Palynological survey of the subtribe Elephantopinae (Asteraceae, Vernonieae)

Despite the taxonomic value of pollen morphology within the tribe Vernonieae, a detailed study of the pollen of the subtribe Elephantopinae is still lacking. The pollen morphology of ten species, representing three of the four genera of the subtribe, Elephantopus, Pseudoelephantopus and Orthopappus, was studied with LM, SEM, and TEM. The pollen of all the species studied was found to be echinolophate, although the differences in aperture features and both sculpture and exine structure allowed recognizing two pollen types. The species of Elephantopus (except for E. elongatus) and Pseudoelephantopus share the regular or more or less regular ridges pattern, the colporate condition but with the ectoapertures little developed, and the acaveate exine structure. From the sculpture and the apertures, the pollen of Caatinganthus harleyi was found to be similar to that of Elephantopus and Pseudoelephantopus. The pollen morphology of the monotypic Orthopappus, which is shared with that of E. elongatus, was further investigated for the first time. We found that it differs from that of the other species in having an irregular pattern of ridges, colporate condition but with a well-developed ectoapertures, and caveate exine. Additional studies of the exine structure and apertures features, coupled with molecular phylogeny, are needed to understand the evolution of pollen characters and re-evaluate the intergeneric relationships within the tribe.     

POLLEN MORPHOLOGY AND DETAILED STRUCTURE OF FAMILY COMPOSITAE,TRIBE CICHORIEAE. II. SUBTRIBE MICROSERIDINAE

A survey of pollen morphology of 40 species representing eight genera of the primarily North American subtribe Microseridinae reveals seven of the eight genera to have caveate, echinolophate, tricolporate grains, Picrosia being the only taxon with echinate pollen. Sectioned grains reveal the exine to consist of an ektexine and endexine. The ektexine, composed of spines, columellae, and foot layer appears to be of two basic types, one with six or seven levels of horizontally anastomosing columellae which are reduced to a single columellar layer under the paraporal lacunae and the second, a bistratified ektexine not reduced to a single layer below the paraporal lacunae. Sectioned exines of Pyrrhopappus are unusual, having very large columellae fused to the foot layer below ridges and highly reduced columellae under lacunae. Endexine organization is similar in most of the genera. Exceptions to this are Pyrrhopappus and some species of Agoseris, which have an “endexine 2” layer. Subtribe Microseridinae is essentially stenopalynous. The pollen data support most of the relationships suggested by Stebbins in his classification. The genera Agoseris, Microseris, Nothocalais, and Phalacroseris seem to form a natural group while Krigia and Pyrrhopappus form another cohesive series. The position of Picrosia, as an advanced offshoot of Pyrrhopappus, is not supported by the pollen data.     

第四纪地层中常见的菊科植物花粉及其起源与分布

文中描述近年来在不同地区第四纪地层中发现的20多种类型菊科植物花粉形态特征,主要有3大类型:外壁具刺型、外壁内部具基柱型和大网胞型。并对它们的植物亲缘关系进行分类探讨。这些类型的化石花粉的植物关系都可归属到相应的现代菊科植物的族或属。据记载,早始新世古地中海至美国西北部亚热带略干旱温暖环境中出现的菊科祖先,可能是菊科植物的起源中心。根据现代植物属(或族)的相关资料,文中讨论了主要类型花粉的起源或分布。绝大多数类型分布于欧亚大陆及北美,在亚洲主要集中于喜马拉雅山及中国西南部。     

POLLEN MORPHOLOGY AND DETAILED STRUCTURE OF FAMILY COMPOSITAE,TRIBE CICHORIEAE. I. SUBTRIBE STEPHANOMERIINAE

A survey of pollen morphology in 20 species representing the 11 genera of the North American subtribe Stephanomeriinae by light, scanning electron, and transmission electron microscopy revealed 10 of the 11 genera to have echinate, tricolporate pollen grains, Lygodesmia being the only genus with echinolophate pollen. Sectioned exines of most of the species examined are similar, being composed of ektexine and endexine. The ektexine surface is composed of spines which typically have globose perforate bases. A cavus occurs as a separation between the basis (foot layer) and the columellae in all of the genera examined except Chaetadelpha. Pollen of the two species of Glyptopleura were found to be strikingly different in exomorphology. Pollen of the putatively self-fertile G. marginata has much shorter spines than the closely related G. setulosa. Atrichoseris, Anisocoma, Calycoseris, Glyptopleura, Pinaropappus, Prenanthella, and most species of Malacothrix have pollen which lack paraporal ridges. The remaining genera, Chaetadelpha, Lygodesmia, Rafinesquia, and Stephanomeria have well-developed ridges of fused spine bases around the apertures. Pollen characters, particularly those of the aperture region, have been found to be systematically useful in the subtribe, therefore acetolyzed material gives more useful information than untreated pollen.     

Pollen morphology in thePortulacaceae (Centrospermae)

Pollen of 110 species from 18 genera in thePortulacaceae has been examined by light and scanning electron microscopy, and a representative number by transmission electron microscopy. Three basic pollen types were found: 3-colpate with thick tectum and foot layer with prominent unbranched columellae and an extremely thin endexine; pantoporate with thick tectum and foot layer with branched columellae enclosing pores and an endexine that is one to two layers thick; pantocolpate with thin tectum and foot layer with broad, short unbranched columellae and an inconspicuous endexine. All pollen types, however, have a spinulose and tubuliferous/punctate ektexine. Also, all the genera except three,Calandrinia H.B.K.,Montia L. andTalinum Adanson are stenopalynous. There is, however, no absolute correlation between pollen morphology and geographical distribution, although both the major centre of palynological diversity and the majority of all species with tricolpate grains occur in South America.     

THE MORPHOLOGY OF THE EXINE IN NIGELLA (RANUNCULACEAE)

The pollen morphology of eight species of Nigella (Ranunculaceae) was examined by scanning and transmission electron microscopy. The exomorphology of all species was identical: 3-colpate, spinulose, and punctate, but thin sections revealed two structural patterns. The ektexine structure of Nigella integrifolia, consisting of thickened foot layer, columellae, and thin tectum, is typical for the family as well as the order Ranunculales in general. In contrast, the remaining seven species, N. arvensis, N. damascena, N. elata, N. hispanica, N. sativa, N. segetalis, and N. stellaris, have an ektexine with an additional unit, a horizontal layer with shorter columellae, placed between the foot layer and tectum. Of all genera examined in the Ranunculaceae, only Nigella had this unusual stratification. This difference in the exine structure would add support to the treatment of N. integrifolia as a monotypic genus, Komaroffia integrifolia (Regel) Lemos Pereira.     

Pollen characteristics of New Zealand Gnaphalieae (Compositae) and their taxonomic significance II. Tem

Pollen ultrastructure has been investigated in 35 taxa of Gnaphalieae (Compositae), predominantly from New Zealand. Pollen grains of all taxa examined are tectate‐columellate and caveate. The characters regarded as particularly distinctive include variability of columellar form and internal organisation of the columellae. The importance of pollen characters in the classification of the New Zealand Gnaphalieae is discussed. The species of Haastia, characterised by senecioid pollen and internal tecta, do not belong to the Gnaphalieae. In the species with helianthoid pollen and sub‐columellae five different columellae types can be distinguished.     

DNA content in species of Vernonia and Vernonanthura from South America: An approach to systematics and evolution of the Vernonieae (Asteraceae)

There are relatively few studies of DNA content in the Vernonieae (Asteraceae) tribe. The first studies were realized in the Lessingianthus genus and determined the DNA content of 25 species. After DNA content, ploidy level and the total karyotype were compared in 6 Chrysolaena species. The aim of this study was to present, for the first time, the DNA content values of Vernonanthura and Vernonia and to thereby expand knowledge of the Vernonieae tribe. A total of 19 natural populations belonging to the genera Vernonanthura and Vernonia were studied for the first time. The results were compared with other Vernonieae genera and with other Asteraceae tribes. Our results found that Vernonieae have the smallest range of 1C value variation in Asteraceae. Furthermore, there were differences in the DNA content of Vernonia and Vernonanthura. These results show that low DNA content and herbaceous habit in Vernonia are characters derived from the higher DNA content and woody habit present in Vernonanthura. These results could indicate a hybrid origin of one species and allow the determination of both the ploidy and chromosome number of other taxa. The results observed in Vernonanthura species showed a highly significant correlation between 1C-value and latitude.     

New species and combinations in Vernonia sections Leiboldia and Lepidonia (Asteraceae), with a revisional conspectus of the groups

The wholly North American sectionLeiboldia ofVernonia as revised by Jones (1979) included six species. The present treatment recognizes only three species inLeiboldia, including one newly described from recent collections. The remainder of what Jones consideredLeiboldia has been transferred into a new sect.Lepidonia (Blake) Turner which was first proposed as a monotypic genus. A key to the two sections and 10 species is provided along with an account of their interrelationships. Morphological evidence suggests that sect.Lepidonia is one of the more primitive sections ofVernonia, being as close to certain African sections as they are to the mostly American sect.Vernonia. Consideration is given to the phyletic significance and generic value of receptacular pales in the Vernonieae from which it is concluded that these have persisted in three or more quite unrelated lines of this tribe and undue weight should not be accorded these in generic considerations. Consequently, the paleaceous, monotypic generaLepidonia andBolanosa are sunk into synonymy underVernonia, the former as a distinct section, the latter into the sect.Vernonia.     

The Pollen Morphology and Exine Ultrastructure of Paeonia L in China

The pollen morphology of 9 species of Paeonia L. has been investigated with both light microscope and scanning electron microscope. In addition, the exine structure of pollen grains of Paeonia suffruticosa and P. lactiflora was examined by transmission electron microscope. Tricolporoidate aperture is an important character of the pollen grains of the Paeonia. The surface of the exine is characterized by reticulate, foveolate and irregularly tuberculate-foveolate sculpture under the SEM. Thin sections of the pollen of this genus shows that the layers of exine are complete i.e. a perforate rectum to semitectum, columellae and foot layers. The endexine is continuous, considerably thickened in the aperture areas and relatively thin or indistinct in the mesocolpia. Paeonia has been placed in Ranunculaceae. But since the beginning of this century many authors have suggested to separating Paeonia from Ranunculaceae. Pollen marphology supports such separation. In Ranunculaceae most pollen grains are tricolpate or have other types of aperture, and exine with spinules and perforations between them. In electron microscopy, the ektexine contains a foot layer, columellae, and perforate rectum, the columellar layer with two types of columellae; the endexine is generally thin. However, the columellar layer of Paeonia has only monomorphic columellae. Some authors considered that there is a close relationship between Paeonia and the Dilleniaceae, but these also differ in the characters of the pollen grains. In Paeonia the constriction of the colpus in equator is in some degree similar to that of Theaceae (Camellia sasanqua Thunb.), Guttiferae (Hypericum L.), Actinidiaceae and Rosaceae. But in the other respects they are quite different. In sum, the pollen morphology of Paeonia is unique. So the palynological information supports Takhtajan's view that Paeonia should be elevated to a family (Paeoniaceae) or order (Paeoniales).     

Pollen wall development in Vigna vexillata I. Characterization of wall layers

Light and electron microscope observations characterized the layers that comprise Vigna vexillata L. pollen walls, and identified the timing of their development. Exine sculpturings form an unusually coarse ektexinous reticulum. The structure of the ektexine is granular; this differs from the columellate/tectate type of structure typical of most angiosperm pollen. The ektexine overlies a homogeneous-to-lamellar, electron-dense endexine, which in turn surrounds a thick, microfibrillar intine. Pollen grains are triporate and operculate, with Zwischenkörper and thickened intine underlying the apertures. The ektexine forms during the tetrad period of microspore development, the endexine and Zwischenkörper during the free microspore stage, and the intine during the bicelled (pollen) stage. Coarsely reticulate exine sculpturings and the granular structure of the patterned exine wall of the pollen grains are features that make this species suitable for detailed studies of pollen wall pattern formation.     

Pollen and the Evolution of Arctotideae (Compositae)

Recent molecular studies have elucidated the phylogeny of Compositae tribe Arctotideae, and found it to contain two, well supported, monophyletic subtribes, Arctotidineae and Gorteriinae, as well as some polyphyletic and problematic genera. On the basis of this new information, it may now be possible to identify diagnostic characters and synapomorphies to support the groupings defined within Arctotideae. Pollen characters have been shown to be particularly variable in Compositae. This paper aims to investigate the utility of those characters in the context of recent molecular phylogenies, in order to determine synapomorphic and diagnostic characters in Arctotideae. The pollen of each genus is described, illustrated with scanning electron micrographs, and optimised on a phylogeny of the tribe. Many pollen characters were found to be very informative when considered in the context of the current best estimate of phylogenetic relationships. Pollen morphology provides synapomorphies for clades at a number of hierarchical levels within Arctotideae, including the two subtribes, Arctotidinae and Gorteriinae, the grouping of Eremothamnus and Hoplophyllum, and smaller clades. It also supports the exclusion of Platycarpha from the tribe. The plesiomorphic palynological state for the tribe is discussed. Particular attention is paid to the evolution of different patterns of lophae (surface ridges). A single origin for the lophate condition is proposed as the most parsimonious mode of evolution in Arctotideae.     

Pollen morphology of the subtribe Cuspariinae (Rutaceae)

The neotropical subtribe Cuspariinae (Rutaceae) comprises as many as 26 genera and over 125 species. Pollen grains from 111 collections representing 71 species and 24 genera were examined by LM, SEM, and TEM. The pollen morphology of this subtribe is very diverse. Grains are mostly 3–6-aperturate and colporate, rarely porate (Spiranthera) or pantocolporate (Almeidea). Exine sculpturing is most commonly reticulate, sometimes perforate, foveolate-perforate, foveolate, foveolate-reticulate, reticulate, striate-reticulate, echinate, clavate, or baculate. The exine structure is columellate and tectate-perforate, columellate and semitectate, or intectate and is stratified into ektexine and endexine. The exine ofLeptothyrsa is distinctive in that the ektexine of the mesocolpium is longitudinally deeply ridged. The pollen ofHortia, characterized by a psilate exine with rare perforations, a very thick foot-layer, and reduced columellae, is unlike that of any member of the Cuspariinae and offers no support for the transfer of this genus from the Toddalioideae. The pollen data correlate with macromorphological characters and are taxonomically useful.     

Chromosome Numbers and Karyotypes of Species of Vernonia sect. Lepidaploa (Asteraceae: Vernonieae)

Vernonia is the largest genus of the tribe Vernonieae (Asteraceae) and comprises more than 1,000 species. In the present study we explore chromosome number and karyotype variation of eight species treated within different subsections of the section Vernonia sect. Lepidaploa. We aimed to explore if these data support the recognition of a single large genus (sensu Baker) or favor its splitting into 22 small genera (sensu Robinson). The species were collected in ??cerrado??, rupicolous and disturbed areas in the states of S?o Paulo and Minas Gerais, Brazil. Chromosome numbers varied from 2n?=?32 to 60. Most chromosomes were small, and the karyotype analysis revealed a predominance of metacentric and some submetacentric chromosomes. The karyotype symmetry in Vernonia was moderate (TF% 32.2 to 45.9), with the most symmetrical karyotype observed in V. rubriramea. The results obtained here did not conclusively support any of the taxonomic proposals for Vernonia due to the absence of distinctive or characteristic karyotype patterns for any of the taxonomic groupings, i.e., sections and subsections (sensu Baker) or new genera (sensu Robinson). Nevertheless, a tenuous relationship was observed between the chromosome numbers reported in the literature, those recorded here, and the taxonomic alterations suggested by Robinson for the genera Lessingianthus, Chrysolaena, and Vernonanthura that were originally part of Vernonia sensu Baker.     

Pollen morphology and its systematic implications for the genera<Emphasis Type="Italic">Keiskea</Emphasis> miq. and<Emphasis Type="Italic">Collinsonia</Emphasis> L. (Elsholtzieae-Lamiaceae)

The pollen morphology of six species ofKeiskea and three representative taxa ofCollinsonia was studied in detail using LM, SEM, and TEM. In both genera, pollen grains are monad, hexa-colpate, and mostly medium in size [P = 28.0 to 37.0 μm, E = 24.3 to 30.7 μm (Keiskea); P = 30.0 to 45.0 μm, E = 26.0 to 39.0 μm (Collinsonia)]. Polar outlines are of circular or ellipsoid form. Shapes range from primarily oblate-spheroidal to prolate-spheroidal to subprolate, and rarely prolate in the equatorial view. Their exine, including the inline characters, are clearly distinct from each other:Keiskea, well-developed bi-reticulate, often forming large lumina by supratectal ridges, unbranched columellae, one-third to one-half of the total exine thickness; versusCollinsonia, mostly perforate without supratectal ridges or a faint/very weak bi-reticulate appearance without supratectal ridges, seemingly branched columellae, ca. two-thirds of the total exine thickness. As demonstrated by these current data, the pollen morphology of the two genera is well distinguished, easily supporting the separation ofKeiskea fromCollinsonia.     

POLLEN MORPHOLOGY OF THE DILLENIACEAE AND ACTINIDIACEAE

Pollen of 53 species of Dilleniaceae and Actinidiaceae was examined by light microscopy, scanning electron microscopy, and a selected group in transmission electron microscopy. Dilleniaceae pollen ranges from tricolpate, tricolporate, tetracolpate, and incipiently inaperturate. Tricolpate types occur only among the Old World subfamily Dillenioideae and the compound aperturate (3-colporate) condition is restricted to the subfamily Tetraceroideae. Within the Dilleniaceae the tricolpate pollen type with elongated apertures is considered primitive, having given rise to the 3-colporate and 4-colpate conditions. The striking pollen dimorphism in the Neotropical species of Tetracera, all of which are androdioecious, is documented; however, in contrast to previous reports, pollen from bisexual flowers appears to be incipiently inaperturate and not pantoporate. The inaperturate condition is interpretated as an early stage in the evolution of outcrossing. Pollen morphology does not support a close relationship between Dilleniaceae and Actinidiaceae. Pollen morphological differences that can be noted between these families are: tectum complete and predominantly psilate or psilate-granular in Actinidiaceae, tectum incomplete, punctate to reticulate in Dilleniaceae; an equatorial bridge of ektexine over the endoaperture usually present in Actinidiaceae, absent in Dilleniaceae; columellae reduced in Actinidiaceae, columellae usually well-developed in Dilleniaceae. Pollen morphology does not argue against a close relationship between Actinidiaceae and Theaceae.     

Pollen morphology of Gyrostemonaceae,Bataceae andKoeberlinia

Pollen morphology of Gyrostemonaceae, Bataceae, andKoeberlinia, which have been affiliated with glucosinolate-producing taxa, was examined by field emission scanning and transmission electron microscopy. Pollen grains of Gyrostemonaceae are tricolpate with scabrate-spinulate surface and have a thick, unstratified exine, while those of Bataceae are tricolporoidate with granular surface and have a thin exine with a single, outermost granular layer. Gyrostemonaceae and Bataceae, which had often been considered sister taxa based on palynological similarity and now are considered more distantly related, have a similar spongy ektexine, but differences between them are evident.Koeberlinia, which is recently considered a sister group to Bataceae+Salvadoraceae (with no spongy ektexine), has tricolporoidate pollen composed of a plesiomorphic, stratified exine with columellae. The totality of evidence indicates that, contrary to earlier observations, pollen of Gyrostemonaceae and Bataceae does not closely resemble each other, and that the spongy ektexine, which looks to be similar in TEM sections, is a homoplasy that evolved independently in the two families. Dedicated to the late Prof. Emer. Kankichi Sohma (August 28, 1926–June 26, 1995), who supervised us for our M. Sc. and D. Sc. programs at Tohoku University, Sendai. He died after his 40 years career in palynology; his wide range of interests and enthusiasm for research, and his unfailing encouragement for students are greatly missed.     

POLLEN ULTRASTRUCTURE OF THE TRIBE INGEAE (MIMOSOIDEAE: LEGUMINOSAE)

All genera within the Ingeae, excluding Wallaceodendron, were examined with the transmission electron microscope. Thin sections reveal two pollen types (Types I and II) distinguished primarily by differences in polyad cohesion and ektexine organization. Type I polyads (only eight-grained species of Calliandra) are calymmate and the ektexine of individual cells is continuous around the grain, organized into a thin, foraminate tectum, irregularly shaped, often basally flared, foraminate columellae and thin, discontinuous foot layer. Type II polyads (16-grained species of Calliandra and remaining Ingeae) are predominantly acalymmate with individual grains typically free from one another or rarely, partially calymmate, i.e., individual grains show limited forms of attachment through small endexinous bridges (Pithecellobium latifolium [Zygia], Lysiloma) or localized appression of adjacent endexines (Pithecellobium daulense [Cathormion]). The adhesion of individual grains through localized fusion of lateral-distal and proximal ektexine in Enterolobium is unique among the partially calymmate Type II polyads. Ektexine in Type II polyads, largely restricted to the distal face, is composed of a thick, channeled tectum, granular interstitium and when present, thin discontinuous foot layer. Lateral-distal and proximal areas exhibit only endexine and, occasionally, a foot layer. The occurrence of nondistal ektexine is restricted to Enterolobium. The pollen data suggest that the acalymmate Ingeae polyads composed of grains with porate apertures, thick, highly channeled tectum, granular interstitium and lack of, or greatly reduced foot layer, are clearly derived within the Mimosoideae. Type I calymmate polyads appear to be independently derived. Ultrastructural data suggest that the Ingeae, excluding the eight-grained Calliandra species, represent a natural grouping with a close affinity to the Acacieae.     

Sesquiterpene lactones of Vernonia — influence of glaucolide-A on the growth rate and survival of Lepidopterous larvae

Summary Sesquiterpene lactone glaucolide-A from Vernonia, incorporated in the rearing diets of five species of Lepidoptera, significantly reduced the rate of growth of larvae of the southern armyworm, Spodoptera eridania; fall armyworm, S. frugiperda; and yellowstriped armyworm, S. ornithogalli. Quantitative feeding tests demonstrated that decreased feeding levels and reduced growth resulted from ingestion of a sesquiterpene lactone. Ingestion of glaucolide-A increased the number of days to pupation in four of the species. In the southern armyworm, it significantly reduced pupal weight. Glaucolide-A decidedly reduced percentage of survival of southern and fall armyworms. Yellow woollybear, Diacrisia virginica, and cabbage looper, Trichoplusia ni, larvae were essentially uneffected by the ingestion of the sesquiterpene lactone. Sesquiterpene lactones adversely affect growth rate and survival of certain insects that feed upon plants containing them. They apparently function as defensive products, screening out a portion of the potential herbivores.Vernonieae: Compositae