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.     

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 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.     

POLLEN APERTURE EVOLUTION AMONG THE SUBFAMILIES PERSOONIOIDEAE,SPHALMIOIDEAE, AND CARNARVONIOIDEAE (PROTEACEAE)

Pollen apertures were analyzed among the subfamilies Persoonioideae (seven genera; ca. 95 spp.), Sphalmioideae (one genus; one spp.), and Camarvonioideae (one genus; two spp.). Pollen was examined by light microscopy, cryosection, and transmission electron microscopy. Completed studies of pollen apertures among Grevilleoideae (ca. 40 genera; ca. 800 spp.), one of two major subfamilies in Proteaceae, provide a basis for comparison and analysis of aperture evolution among these subfamilies. Aperture characters within Persoonioideae are unique among Proteaceae examined to date. Five distinct aperture types occur among the three subfamilies, three of which (Placospermum, Persoonia, Bellendena) are restricted to Persoonioideae. Sphalmioideae and Camarvonioideae each exhibit a unique aperture organization. The most primitive aperture organization, and one unique to Placospermum, exhibits three main features: 1) a thin, granular endexine continuous around the grain; 2) a heterogeneous foot layer throughout the grain with increased disruptions at the aperture; and 3) only slight differences in exine characters between apertural and nonapertural regions. The Persoonia aperture type represents the next stage of aperture evolution which involves loss of endexine, restriction of a heterogeneous foot layer to the aperture, and marked differences in exine characters between apertural and nonapertural regions. The uniformly homogeneous ektexine in both nonapertural and aperture regions in Bellendena has developed independently. Sphalmium exhibits a primitively thin granular endexine though the restriction of endexine to the aperture is a derived condition. Carnarvonia exhibits several pollen characters also found among Grevilleoideae including: 1) a homogeneous nonapertural ektexine; 2) a slightly heterogeneous apertural ektexine; 3) a lamellate/granulate endexine organized into irregularly shaped “clumps” clustered around the aperture; and 4) a clear demarcation between apertural and nonapertural exine. These characters support the hypothesis that Carnarvonia may have diverged early from the pre-Grevilleoids.     

Conventional and novel modes of exine patterning in members of the Araceae – the consequence of ecological paradigm shifts?

Summary. In the family Araceae, the members of all subfamilies except Aroideae follow the conventional mode of exine formation pattern, which conforms with the textbook view of sporoderm stratification and chemistry (sporopollenin ektexine formed before the endexine). Only members of the subfamily Aroideae show a quite uncommon mode of exine formation pattern, with an endexine formed prior to the nonsporopollenin, polysaccharidic outer exine layer. The intine is formed simultaneously with this non-sporopollenin layer. From the differing timetable and especially from the different origin it is concluded that this outer exine layer is not homologous to the angiosperm ektexine. The fundamental question, why members of the Aroideae lack an elaborated sporopollenin ektexine, is discussed in terms of functionality of the nonsporopollenin outer exine layer. It seems that a major change in aroid evolution took place at the point when the family phylogenetically and ecologically shifted from bisexual (most subfamilies) to unisexual flowers (Aroideae only). The hypothesis is that ephemeral spathes and the absence of sporopollenin are the consequence of an adaptive syndrome for a short pollination time window in many members of the Aroideae, with short-lived pollen, an energetically not costly pollen wall, rapid germination of pollen tube, and brief receptivity of stigma. Correspondence and reprints: Institute of Botany, University of Vienna, Rennweg 14, 1030 Vienna, Austria.     

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.     

Pollen Organs and Seeds with Eucommiidites Pollen

The characteristic Mesozoic pollen genus Eucommiidites is described from pollen organs and seeds recovered in Cretaceous strata of North America and Europe. The pollen organs are from the lowermost Upper Cretaceous (Cenomanian) of Texas and are referred to Erdtmanitheca texensis gen. et sp. nov. They are spherical heads, composed of numerous, densely crowded, radiating pollen sacs that contain abundant well-preserved pollen. Combined LM, SEM and TEM investigations show that the pollen grains each have a distinct distal colpus flanked by two lateral colpi in an equatorial position. Pollen wall ultrastructure is gymnospermous with a thick lamellate inner layer (endexine) and an outer layer (ektexine) composed of a granular inner part and a homogeneous outer part. The endexine is thickened in the region of the colpi. Small seeds from the Lower Cretaceous (upper Berriasian to Valanginian) of Bornholm, Denmark contain abundant Eucommiidites pollen in their micropyles. The seeds are referred to Erdtmanispermum balticum gen. et sp. nov. They are ovoid, and weakly triangular in transverse section and gradually taper at the apex into an elongated tube. The megaspore membrane is granular and well developed, and apparently surrounded by three separate tissues interpreted as nucellus, a thin inner integument and a sclerified outer envelope. Eucommiidites pollen in the micropyles of the seeds has a laminated endexine and an ektexine comprising two homogeneous parts separated by a granular layer. Réévaluation of other seeds known to contain Eucommiidites pollen indicates that they share significant similarities with Erdlmanispermum and that they may have been produced by closely related plants. Comparison of “Eucommiidites plants” with other seed plants suggests that they are probably most closely related to the anthophytes comprising Bennettitales, Pentoxylales, Gnetales and angiosperms.     

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.     

Development of the pollen wall in Tsuga canadensis (Pinaceae)

In discussions of exine structural types, Tsuga is often mentioned as an exception, since no infratectal layer is present in the ektexine. The present investigation documents the formation of this pollen wall type at the ultrastructural level in T. canadensis . All layers of the exine are formed during the tetrad period, when the microspores are surrounded by a callose wall. The outer layer (ektexine) is elaborated on a fibrillar microspore surface coat, while the inner layer (endexine) is elaborated on lamellated structures. The deposition of the pretectum is followed by the appearance of endexine lamellae. In the initial stages, the two layers—pretectum and endexine—appear to be separated from each other only by a dense microspore surface coat. As additional wall materials are deposited, the tectal elements become convoluted and come to rest, in places, on the now recognizable footlayer. Upon release from the tetrad, intine formation begins and continuous accumulation of sporopollenin leads to an increase in ektexine thickness. The mature pollen wall of Tsuga canadensis , with a convoluted tectum resting directly on the footlayer, is characteristic of the genus.     

Lophate sculpturing of Vernonieae (Compositae) pollen

Vernonieae (Compositae) taxa Vernonia and Elephantopus were examined by scanning (SEM) and by transmission electron microscopy (TEM). All pollen grains of Vernonia have a subechinolophate sculpturing consisting of a wrinkled net-like ektexine delineating a system of thickened and randomly meandering ridges and corresponding valleys. Ektexine on the ridges radiates from near mid-regions of massive, individual columellae that terminate in sharp spines approximately 4.5 μm above the ektexine surface. Less massive secondary columellae without apical protrusions support the ektexine net in the spine areas. In the valleys, the ektexine is markedly depressed. Removal of the net by freeze-fracture exposes an exine surface that is clearly imprinted with distinctive lophate features. Elephantopus pollen is echinolophate and extends the morphological range for this little studied genus. Lophae are the thickened equivalents of ektexine tecta and represent the highly anastomosing and intertwined rod-like elements of forked distal parts of densely clumped, massive columellae. Lophae surfaces contain rare punctations that are randomly scattered among blunt spinules ranging from approximately 0.3 to 1.5 μm in height. It is clear that only one sculpture pattern, lophate, characterizes Vernonieae pollen and that it embraces two forms, subechinolophate and lophate. Of the two structural patterns in the Compositae, caveate and noncaveate, subechinolophate sculptured Vernonia pollen is noncaveate. Elephantopus pollen is also noncaveate but more generalized than Vernonia.     

FINE STRUCTURE OF MISTLETOE POLLEN. V. MADAGASCAN AND CONTINENTAL AFRICAN VISCUM L. (VISCACEAE)

Pollen characters of Madagascan and continental African Viscum are described and compared to those in Asia and Australia. The subprolate, tricolporate, nonuniformly sculptured pollen of Madagascan taxa is most similar to that of Asian species. Ultrastructurally, however, the completely granular equatorial ektexine of Madagascan Viscum is most similar to that of continental African taxa. Continental African Viscum, in contrast to Madagascan and Asian species, display a wide variation in pollen shape and apertures. Pollen shape ranges from subprolate to oblate, the latter unique to Africa. The most striking feature of continental Viscum is their variability in aperture number and aperture type. Aperture number varies at both the intra- and interpopulational levels with such variation resolvable to the individual flower—a condition unique to the continent. The only simple (colpate) aperture type in the genus is restricted to Africa. The continental species can be divided into two species groups based on pollen characters: Group I (4 spp.) characterized by strictly tricolporate rounded convex pollen with a rodlet/granular equatorial ektexine structure and Group II (most continental species) possessing multiapertures, concave lobate shape, uniform sculpturing and granular equatorial ektexine. The African V. menyharthii, V. fischeri, V. rotundifolium and V. minimum exhibit no clear Group I or II affinities. An analysis of overall pollen characters in Viscum indicates a trend towards spheroidal shape, multiapertures and uniform sculpturing and ektexine organization. Though pollen characters suggest ties between Australia, Asia and Madagascar, they indicate an even stronger relationship between Madagascar and continental Africa, particularly eastern Africa. The relationship of the majority of African Viscum, excluding those with obvious Madagascan affinities, remains obscure. The unique palynological features of Group II species coupled with their inflorescence structure suggest an independently evolving group.     

FINE STRUCTURE OF MISTLETOE POLLEN. IV. EURASIAN AND AUSTRALIAN VISCUM L. (VISCACEAE)

Fifteen Eurasian and Australian species of Viscum L. were examined by light, scanning, and transmission electron microscopy. Pollen characters divide the species into two groups, each correlated with differences in habit and inflorescence structure: Group I (12 spp.) characterized by psilate or slightly sculptured exines and a non-uniform ektexine pattern and Group II (3 spp.) possessing highly sculptured (echinate, rodlet) surfaces and uniform ektexine patterns. Within each of the groups, pollen characters divide the species into several subgroups. Among Group I, species V. nepalense, V. heyneanum and V. ovalifolium are particularly close. The Group I species, V. trilobatum, is placed in its own subgroup primarily because of its uniform ektexine pattern—a unique feature among Asian and Australian Viscum. Of the three Group II species, V. album and V. alniformosanae are palynologically almost indistinguishable. Pollen of the Group II V. cruciatum, though exomorphologically similar to V. album, is closest ultrastructurally to the Indian V. trilobatum. Overall, the most common and probably basic pollen characters among the Eurasian and Australian species include: subprolate, rounded convex almost spherical shapes; tricolporate apertures, and non-uniform sculpturing and ektexine patterns. Oblate-spheroidal or prolate-spheroidal shapes, prominent sculpturing, and a uniform ektexine pattern are derived characters largely restricted to the Eurasian and Asian Group II species (V. album, V. alniformosanae, V. cruciatum).     

On the Ultrastructure of Pollen Exine in Metasequoia Hu and Cheng

Metasequoia is endemic to China. Present study deals with ultrastructure of pollen exine of M. glyptostroboides Hu et Cheng, and in comparision with other genera of the family. Pollen grains of Metasequoia are spheroidal or subsphoroidal and 27.8(24.3–32.3) μm in diameter. There is a papilla in the distal face. The papilla is wide at the base, 3.5–5.2 μm high, with pointed and circular end and the base crooked toward one side. Exine is about L5 μm thick, layers distinct, Nexine is as thick as sexine. Surface weakly granulate. According to observation by SEM, exine is covered with fine granules and rather coarse tuberculae. The former can be easily separated from the latter. The loose and uneven tuberculae are provided with minute spinules on the surface and generally fall off after acetolysis. The fine and dense granulae, however, remain intact after acetolysis. The study by TEM shows that ektexine is made of granules densely arranged and connected with each other. In addition, sparse Ubisch bodies are unevenly distributed on granular layer with geminate surface. The thick endexine, is composed of 10–15 lamellae. It is worthy to note that all lamellae possess tripartite structure. But lamellae of endexine in other genera of Taxodiaceae have no tripartite structure except the lamella near ektexine. Number of lamella and thickness of endexine in Metasequoia differ from those of other genera in Taxodiaceae; for example endexine with 8–10 lamellae in Taxodium, 8–9 lamellae in Sequoia, 6–7 lamellae in Glyptostrobus, 6–8 lamellae in Cunninghamia, about 16 lamellae in Cryptomeria etc.     

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.     

FINE STRUCTURE OF PARKINSONIA ACULEATA POLLEN. I. THE POLLEN WALL

Larson , Donald A., and C. Willard Lewis , Jr . (U. Texas, Austin.) Fine structure of Parki nsonia aculeata pollen. I. The pollen wa ll. Amer. Jour. Bot. 48(10): 934–943. Illus. 1961.—Fresh pollen samples and chemically fossilized pollen samples were fixed in either potassium permanganate or osmium tetroxide and some sectioned material was poststained in an effort to demonstrate the various wall components existing in pollen of P. aculeata. The exine was observed to be consistently 2-layered with the layers corresponding to the endexine and ektexine demonstrated with basic fuchsin staining by optical microscopy. The exine also contained an extensive system of internal channels. Aperture membranes contained laminate-globulate fine structure, and this has been interpreted as a modification for the maintenance of membrane continuity through changes in pollen grain volume.     

Pollen morphology of the genera Polygonum s. str. and Polygonella (Polygoneae: Polygonaceae)

The pollen of 30 taxa (27 species, one subspecies and two varieties) in two genera, viz Polygonum s. str. and Polygonella was investigated with LM and SEM, and some selected taxa with TEM. In all genera investigated the pollen is prolate to spheroidal, and the aperture is mostly tricolporate, rarely panto-hexacolporate (especially Polygonum section Polygonum). The exine sculpturing pattern is the most variable feature. Three types of exine can be recognized. Type 1 (Avicularia-Type, sensu Hedberg) - All species of section Polygonum and section Tephis share the smooth tectate exine with spinules, sometimes the surface is more or less rough (Polygonum afromontanum in section Tephis). Type 2 (Pseudomollia-Type, sensu Hong) - Pollen of Polygonum molliaeforme (section Pseudomollia) has the exine, which is verrucose on both poles and nearby the mesocolpium, and mostly psilate around the ectoaperture. Type 3 (Duravia-Type, sensu Hedberg) - Pollen grains of Polygonum section Duravia and Polygonella have the exine which is semitectate-reticulate at the mesocolpium and the poles, and rugulate/reticulate or sometimes foveolate with microspinules around the ectoapertures. The pollen grains in four taxa (viz Polygonum section Pseudomollia, P. section Duravia and genus Polygonella) have a well-marked dimorphism of the ektexine, which is considered to be a synapomorphic condition. The differences of pollen grain between the genus Polygonella and Polygonum section Duravia are almost non existent and clearly interrelated. It is therefore postulated that the similarity in pollen of both taxa is not the result of convergency, but is interpreted as a homology. It is noteworthy that the pollen of Polygonum molliaeforme (section Pseudomollia) appears as intermediate between the Avicularia-type and the Duravia-type, and is well supported the value of separated section for its own. Additionally, in TEM, some exine ultrastructures (e.g. columellae, foot layer, endexine) appear to be valuable characters for comparison between/among taxa. The systematic potentialities of the pollen data of the studied taxa at various systematic levels are also discussed.     

Palynological variation in Balsaminoid Ericales. II. Balsaminaceae, Tetrameristaceae, Pellicieraceae and general conclusions

BACKGROUND AND AIMS: The objective of this study is to examine the palynological diversity of Balsaminaceae (two genera/+/-1000 species), Tetrameristaceae (two genera/two species) and Pellicieraceae (one genus/one species). The diversity found will be used to infer the systematic value of pollen features within the balsaminoid clade. METHODS: Pollen morphology and ultrastructure of 29 species, representing all families of the balsaminoid clade except Marcgraviaceae, are investigated by means of light microscopy, scanning electron microscopy and transmission electron microscopy. KEY RESULTS: Balsaminaceae pollen is small to medium sized with three to four apertures, which can be either colpate or porate, and a sexine sculpturing varying from coarsely reticulate to almost microreticulate. Tetrameristaceae pollen is small sized, 3-colporate, with a heterobrochate reticulate sculpturing and granules present in the lumina. Pellicieraceae pollen is large sized, 3-colporate with long ectocolpi and a perforate sexine sculpturing with large verrucae. Furthermore, Pelliciera is characterized by the occurrence of aggregated orbicules, while orbicules are completely absent in both Balsaminaceae and Tetrameristaceae. Balsaminaceae pollen differs from the other balsaminoid families due to the occurrence of colpate or porate grains with an oblate to peroblate shape, a very thin foot layer and a lamellated endexine. CONCLUSIONS: From a pollen morphological point of view, Balsaminaceae are completely different from the other balsaminoid families. Therefore, no pollen morphological synapomorphies could be defined for the balsaminoid clade. However, various pollen features were observed that could indicate a possible relationship between Tetrameristaceae, Pellicieraceae and Marcgraviaceae. Despite the palynological similarities in the latter three families, it remains unclear to what extent they are related to each other.     

Genetics and development of morphological and physiological characters of male sterility in Oenothera

Summary Male sterility in Oenothera is influenced by two nuclear genes,fr andster. Their function is independent of the plastomes. Development of anthers, fertile and sterile male, was studied by electron microscopy and histochemical methods. Both genes act on lipid metabolism but at different developmental stages. Infr/fr homozygotes the disturbance is expressed as a lack of sporopollenin in the exine, while amorphous lipid material is deposited in the loculus. Inster/ster homozygotes sporopollenin is formed normally in the endexine but the paracristalline structure of the ektexine is missing. In both mutants the disturbance leads to complete destruction of the pollen grain. The deviation from fertile pollen development is correlated with abnormalities of the tapetum and outer cell layers of the anther wall.     

ULTRASTRUCTURE OF PARKIA POLYADS (MIMOSOIDEAE: LEGUMINOSAE)

Pollen of 16 species of Parkia (ca. 30 spp.) was examined in the light, scanning and transmission electron microscopes (LM, SEM and TEM). Pollen is shed as large (61-236 μm), globose, 16–32-grained polyads that remain united following acetolysis. Thin sections reveal that polyad cohesion is maintained through fusion and/or adhesion and/or appression of adjacent ektexines in lateral and/or proximal and/or distal regions. In lateral and proximal regions the ektexine is often embedded in a dense granular endexinous matrix. Individual grains of the polyad are clearly outlined in most species but are completely obscured in several New World species. Sculpturing varies both in pattern and type of element. Nonuniformly sculptured polyads, in which the sculpturing of individual grains differs between the peripheral and central portions of the distal face, occur only among Old World species. Uniformly sculptured polyads occur among both New and Old World taxa. Two main sculpturing types are present among Parkia species: verrucate and tectate-perforate, the latter particularly well developed in the reticulate polyads of the neotropical P. ulei and P. multijuga p.p. Ultrastructurally, all Parkia polyads exhibit ektexine and endexine continuous around the individual grain. Distal ektexine is prominent and clearly stratified, usually exhibiting a thick, perforate tectum, columellate interstitium, and a thin, locally discontinuous foot layer. Lateral and proximal ektexine is significantly thinner and less structured. Individual grains within the polyad possess three to four peripheral-distal apertures usually at the points of contact between adjacent grains. Proximal and central grain apertures occur in most, but not all, species. Polyad characters suggest a particularly close relationship among the New World species P. decussata, P. discolor, P. gigantocarpa, P. igneiflora, and P. nitida. Parkia ulei, though sculpturally closest to P. multijuga p.p., is ultrastructurally most similar to the African P. biglobosa. The large tectate-perforate-fossulate polyads of P. pendula and P. platyparkia are unique in the New World.     

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).