Pollen wall development in Sciadopitys verticillata (Sciadopityaceae)

Pollen wall development of Sciadopitys verticillata was observed by transmission electron microscopy. The pollen of S. verticillata is non-saccate and spherical, and the exine consists of the outer thick, sculptured ectexine and the inner lamellated endexine. At the early tetrad stage, the initial ectexine and lamellae of the initial endexine begin to form on the microspore plasma membrane. The ectexine granules gradually swell. Deposition of sporopollenin materials on the ectexine granules then results it their becoming partially connected to each other. Identification of the original small ectexine granules then becomes difficult, and, finally, the ectexine appears as a homogeneous, partially discontinuous layer. The granules of the early ectexine cannot be identified. At maturity, there are four to five endexine lamellae. Recent molecular data have shown that Sciadopitys first branches off from the Cupressaceae plus Taxaceae clade, which is characterized by granular exine. Although the ectexine of Sciadopitys is similar to that of the Cupressaceae during initial development, the morphology of the ectexine is significantly different in the mature pollen. The initial stage of pollen development clearly shows the structural homology of the granular ectexine. Divergence of the exine structure occurs in the later stages.     

Pollen wall development in Cryptomeria japonica (Taxodiaceae)

Pollen wall development in Cryptomeria japonica was observed by scanning and transmission electron microscopy. The pollen of C. japonica is characterized by a non-saccate, projecting papilla. The exine of C. japonica consists of the outer granular ectexine and the inner lamellated endexine. At the tetrad stage, the initial granular layer of the pro-ectexine first forms on the microspore plasma membrane. The tripartite lamellae of the pro-endexine form under the pro-ectexine. The prosporopollenin material is deposited on the pro-ectexine and pro-endexine at the free spore stage. The ectexine granule increases its volume and the endexine lamellae thicken. The papilla protrudes during the tetrad stage. The tip of the papilla bends laterally where the exine is thinner. Exine construction in C. japonica is similar to that of Cunninghamia; however, the amount and size of the granular ectexine and lamellated endexine differ. The conspicuous papilla protrudes and bends during the tetrad period.     

I. Primexine development in Passiflora racemosa Brot.: overlooked aspects of development

Developmental stages during the tetrad period were examined in detail by transmission electron microscopy with an emphasis on substructure. Our purpose was to find out whether the sequence of sporoderm developmental events provides additional evidence for our recent hypothesis on the underlying cause of exine ontogeny as a sequence of self-assembling micellar mesophases initiated by genomically given physicochemical parameters. Osmiophilic globules encrusting the surface of postmeiotic microspores and tapetal cells are temporary prepattern units which come first. The second prepattern structures are highly ordered bundles of microfilaments and microtubules which determine the position of microspore surface invaginations and clusters of the glycocalyx inside them. The first glycocalyx units are microgranules which during the middle tetrad stage rearrange into radially oriented rod-like units. The latter form lens-like clusters of the glycocalyx-1, located inside the invaginations. These clusters predestine the position of the future luminae in the exine reticulum. The second glycocalyx layer is laid down as a continuous layer over the whole microspore surface and has similar substructure, that is radial rods. Glycocalyx-2 is a framework for procolumellae which appear at the late tetrad stage. Therefore, the sequence of substructural units in the primexine is: globules, microgranules, rod-like units, and layers of radially oriented rods tightly packed in the periplasmic space. This sequence corresponds to the first three mesophases of self-assembling micelles: spherical micelles, cylindrical micelles, and layers of hexagonally packed cylindrical micelles (middle mesophase). We observed the same sequence in other species during primexine development, and the findings of this study provide new evidence for our hypothesis.     

A new look at sporoderm ontogeny in Persea americana and the hidden side of development

Background and Aims

The phenomenon of self-assembly, widespread in both the living and the non-living world, is a key mechanism in sporoderm pattern formation. Observations in developmental palynology appear in a new light if they are regarded as aspects of a sequence of micellar colloidal mesophases at genomically controlled initial parameters. The exine of Persea is reduced to ornamentaion (spines and gemmae with underlying skin-like ectexine); there is no endexine. Development of Persea exine was analysed based on the idea that ornamentation of pollen occurs largely by self-assembly.

Methods

Flower buds were collected from trees grown in greenhouses over 11 years in order to examine all the main developmental stages, including the very short tetrad period. After fixing, sections were examined using transmission electron microscopy.

Key Results and Conclusions

The locations of future spines are determined by lipid droplets in invaginations of the microspore plasma membrane. The addition of new sporopollenin monomers into these invaginations leads to the appearance of chimeric polymersomes, which, after splitting into two individual assemblies, give rise to both liquid-crystal conical ‘skeletons’ of spines and spherical micelles. After autopolymerization of sporopollenin, spines emerge around their skeletons, nested into clusters of globules. These clusters and single globules between spines appear on a base of spherical micelles. The intine also develops on the base of micellar mesophases. Colloidal chemistry helps to provide a more general understanding of the processes and explains recurrent features of pollen walls from remote taxa.     

Sporoderm development in Acer tataricum (Aceraceae): an interpretation

For the first time, the developmental events in the course of complicated exine structure establishment have been traced in detail with transmission electron microscope in the representative of Acer. A new look at unfolding events is suggested using the knowledge of a boundary field, colloid science. Our purpose was to find out whether the sequence of sporoderm developmental events represents, in essence, the sequence of self-assembling micellar mesophases, initiated by genomically given physicochemical parameters and induced by surfactant glycoproteins at increasing concentration. Indeed, the first units observed in the periplasmic space are globular ones (=spherical micelles) which become arranged into rod-like units (=cylindrical micelles). Then, twisted clusters of rodlets form a layer of procolumellae (middle micellar mesophase). The tectum emerges as an untwisting and merging of distal ends of procolumellae (distal untwist of micelle clusters). In the end of tetrad period, when a hydrophilic–hydrophobic switch occurs in the periplasmic space, the contrast reversal of the columellae corresponds to the change of normal micelles to reverse ones. The initiation of the foot layer and the endexine lamellae, with their typical central “white lines”, corresponds to the next—“neat”—mesophase, with its typical central gaps between layers. Aperture sites during development show all the main micellar mesophases and their transitional forms. The data received have supported our previous hypothesis.     

Exine ontogeny in Borago officinalis pollen

The primexine matrix is finely granulo-fibrillar up to callose digestion; it becomes distinctly fibrillar at the free microspore stage. The columellae and the tectum are initiated at the middle tetrad stage, the foot layer and the endexine are initiated when the callose wall digestion begins. The columellae are initiated by the deposition of spiral elements around a clear central zone. This hollow aspect of columella disappears when thickening. The foot layer and the endexine are built by the expansion of plasmalemma derived components. The foot layer appears first at the poles, then at the interapertural levels and at last at the apertures while the endexine appears first at the mesoapertures, then it spreads laterally towards the interapertural levels and, at last, at the poles. The gemmae are formed at the free microspore stage over all the tectum. The thickening of the exine takes place essentially during the free microspore stage and continues during the vacuolate microspore one. Apertures are entirely formed before the complete digestion of the callose wall. The ectoapertures are determined by the lacking of the columellae; the sites of the pericolpal cavities and the mesoapertures result from the plasmalemma retraction even before the setting up of the foot layer and the endexine by which they will be delimited respectively afterwards. The endoapertures are determined by the lacking of compact endexine at their level, and merge into a continuous equatorial belt.     

Sporoderm development in Liriodendron chinense (Magnoliaceae): a probable role of the endoplasmic reticulum

The developmental events in the sporoderm and the cytoplasm of Liriodendron chinense microspore from the early tetrad stage until late free microspore stage were observed. Various forms of the endoplasmic reticulum (ER) and surprising unusual aggregates of ER seen during microspore development attract special attention. Being scanty at early and middle tetrad stage, while primexine matrix (glycocalyx) acquires well-defined form, the ER becomes distinct in unusual forms at the late tetrad stage. Thin long tubules with an osmiophilic contents, which cannot be compared with the tubular smooth endoplasmic reticulum (SER), undulate through the cytoplasm. Towards the end of the tetrad period when callose begins to disintegrate, and a distinct tectate-columellate pattern of the ectexine becomes evident, two new forms of the SER occur in the cytoplasm. Instead of single tubules observed previously, 3–tubuled aggregates meander through the cytoplasm, the middle tubule contains an osmiophilic substance. The second form of the SER looks like an ordinary tubular SER, but has ampoule-like dilations with dark granular contents. Later on the tubules undergo major changes: multi-tubuled aggregates of parallel tubules overcrowd the cytoplasm, the outer tubules of each aggregate carrying ribosomes. These aggregates undulate through the cytoplasm, branch, and are associated with lipid globules. The tips of many aggregates are pressed to the plasmalemma. The ontogenetic period of time of the presence of these ER aggregates, their structure and localization in the microspore cytoplasm allow me to assume that these ER aggregates synthesize sporopollenin precursors.     

Cytochemistry of pollen development in Campsis radicans (L.) Seem. (Bignoniaceae)

In this study, cytochemical staining methods were used to follow the cytochemical modifications of microspore cytoplasm and sporoderm in Campsis radicans (L.) Seem. from tetrad stage to mature pollen. Flower buds were collected at different stages of development, and the anthers were fixed and embedded in Araldite. To make cytochemical observations under light microscope, semithin sections were cut and stained with different dyes. Cytochemical methods provided the opportunity to localize the reserve material in the microspore and pollen cytoplasm, to distinguish the different layers of the sporoderm, and to determine its chemical structure at different developmental stages. Microspore cytoplasm contains variable amounts of proteins, lipids, and insoluble carbohydrates at different stages of microsporogenesis. Sporoderm formation starts at tetrad stage by the formation of primexine and is completed at vacuolated microspore stage by the addition of sporopollenin from tapetum. During the vacuolization and enlargement of the microspores, the structure and the chemical composition of the exine are modified. The endexine becomes chemically different from the ectexine. The ectexine is composed of sporopollenin and a small amount of protein, whereas the endexine is composed of sporopollenin, proteins, and traces of polysaccharides.     

Exine and tapetum development in Symphytum officinale (Boraginaceae). Exine substructure and its interpretation

After detailing the exine ontogeny, our purpose was to find out whether the sequence of sporoderm developmental events corresponds to self-assembling micellar mesophases, initiated by genomically determined physicochemical parameters and induced by surfactant glycoproteins at increasing concentrations. Indeed, a scaffolding of the future exine, i.e., the glycocalyx, initiates with scattered clots, which then appear as clusters of spherical and worm-like micelles, derived from surface-active glycoproteins. At the middle tetrad stage, a continuous layer of the glycocalyx emerges, consisting of parallel, tightly packed cylinder-like units, which we interpret as a layer of cylindrical micelles, the so-called middle mesophase. These units bear dark-contrasted particles, arranged in strings or columns. These sites of the glycocalyx units?Cmicelles accumulate initial sporopollenin, hence the term ??sporopollenin acceptor particles?? (SAPs). This process leads to the appearance of procolumellae at the late tetrad stage. The glycocalyx units are rooted into callose and into the microspore cytoplasm. After formation of the tectum and the foot layer, the endexine initiates as a thin layer, and the latter develops into a very thick layer in the post-tetrad period. When callose disintegrates, ??bouquets?? of SAPs become evident on the tectum, which were evidently hidden inside the callose layer; these structures self-assemble into supratectal gemmae. An unusual, ??hybrid?? type of tapetum was observed. What is observed in Symphytum exine development allows us to obtain more evidence for the hypothesis of the participation of micellar self-assembly in sporoderm development and to bring together the concepts of micelles and of SAPs.     

Development of structure-less pollen wall inCeratophyllum demersum L. (Ceratophyllaceae)

Developmental process of structure-less exine is studied in a hydrophilous plant,Ceratophyllum demersum L., with electron microscopy. The plant shows a characteristic feature in tetrad formation. A callose wall is not synthesized and exine initiation does not occur during the tetrad stage. After release of microspores, a trilaminar layer with two electron-dense lines is formed in the surface of each microspore. The trilaminar layer develops to a thin structure-less exine that is considered to consist of only an endexine. The unusual exine would be an adaptive feature for submersed pollination in fresh water.     

Wall development and its autofluorescence of sterile and fertileVicia faba L. Pollen

Summary The ultrastructural changes of the pollen wall of three types of fertile and one of sterileVicia pollen were related to the autofluorescence of the pollen wall, measured by a microspectroscopic method. Till the liberation of the microspores from the tetrad, the spectrum of the ectexine shows sometimes two maxima and has a very low intensity. After this period the endexine is formed and its spectrum has one maximum with a high intensity. The differences of the pollen wall between the sterile and fertile pollen exist of the presence of one spectral maximum during the tetrad stage, a thick endexine and the absence of the intine in the sterile pollen. The different types show much differences during the tetrad stage in the callose wall as well as the ectexine. The autofluorescence illustrates the complexity and specificity of the pollen wall development.     

Pollen wall ultrastructure and ontogeny in Heliotropium europaeum L. (Boraginaceae)

The pollen grains of Heliotropium europaeum are heterocolpate, with alternation of 3 colpori and 3 pseudocolpi. The exine is characterized by a scabrate and thick tectum, massive columellae with a granular appearance and a thick nexine. The thickening of the intine at the apertural level makes the interpretation of this zone difficult. The ontogenetic study helped to understand the ultrastructure of the exine and the apertures. The different steps are as follows. The primexine matrix is formed during the beginning of the tetrad stage; it consists of an outer thick and electron dense zone and an inner one, less dense to electrons. The tectum and the infratectum begin to form in the outer zone of the matrix, towards the middle of the tetrad stage. The infratectum consists of a network of columellae variable in thickness and oriented in different directions. The foot layer is lacking. The endexine is formed on a lamella system during the callose loss and microspore separation. The endexine becomes compact very early on its inner part. The apertures are initiated during the tetrad stage; a granulo-fibrillar oncus develops. At the free microspore stage, the oncus gets fibrillar and is bordered by endexine lamellae on its outer side and by endexine granulations on its inner one and laterally. The intine is set at the end of this stage. At the vacuolated microspore stage, the intine shows three layers: two thin, clear and homogeneous layers, one outside and the other inside, and a thick middle layer that forms the zwischenkörper, crossed by trabecula, in the apertural areas.     

Pollen wall ontogeny in <Emphasis Type="Italic">Polemonium caeruleum</Emphasis> (Polemoniaceae) and suggested underlying mechanisms of development

By a detailed ontogenetic study of Polemonium caeruleum pollen, tracing each stage of development at high TEM resolution, we aim to understand the establishment of the pollen wall and to unravel the mechanisms underlying sporoderm development. The main steps of exine ontogeny in Polemonium caeruleum, observed in the microspore periplasmic space, are spherical units, gradually transforming into columns, then to rod-like units (procolumellae), the appearance of the initial tectum, growth of columellae in height and tectum in thickness and initial sporopollenin accumulation on them, the appearance of the endexine lamellae and of dark-contrasted particles on the tectum, the appearance of a sponge-like layer and of the intine in aperture sites, the appearance of the foot layer on the base of the sponge-like layer and of spinules on the tectum, and massive sporopollenin accumulation. This sequence of developmental events fits well to the sequence of self-assembling micellar mesophases. This gives (together with earlier findings and experimental exine simulations) strong evidence that genome and self-assembly probably share control of exine formation. It is highly probable that self-assembly is an intrinsic instrument of evolution.     

Pollen and anther development in Nelumbo (Nelumbonaceae)

The Nelumbonaceae are a small family of aquatic angiosperms comprising Nelumbo nucifera and Nelumbo lutea. Historically, the genus has been considered to be closely related to Nymphaeales, however new systematic work has allied Nelumbo with lower eudicots, particularly Platanus. In recent years, studies of pollen development have contributed greatly to the understanding of phylogenetic relationships, but little has been known about these events in Nelumbo. In this paper, pollen and anther development are morphologically described for the first time in N. lutea. A comprehensive ontogenetic sequence is documented, including the sporogenous tissue, microspore mother cell, tetrad, free spore, and mature pollen grain stages. The deposition of a microspore mother cell coat and callose wall, the co-occurrence of both tetrahedral and tetragonal tetrads, the formation of a primexine in tetrads, and primexine persistence into the late free spore stage are shown. The majority of exine development occurs during the free spore stage with the deposition of a tectate-columellate ectexine, a lamellate endexine, and an unusual granular layer below and intermixed with the endexine lamellae. A two-layered intine forms rapidly during the earliest mature pollen stage. Major events of anther development documented include the degradation of a secretory-type tapetum during the free spore stage and the rapid formation of U-shaped endothecial thickenings in the mature pollen grain stage. The majority of mature pollen grains are tricolpate, however less common monosulcate and diaperturate grains also develop. Co-occurring aperture types in Nelumbo have been suggested to be an important transition in angiosperm aperture number. However, aperture variability in Nelumbo may be correlated with the lateness of aperture ontogeny in the genus, which occurs in the early free spore stage. This character, as well as other details of pollen and anther ontogeny in Nelumbo, are compared to those of Nymphaeales and Platanus in an effort to provide additional insight into systematic and phylogenetic relationships. Although Nelumbo is similar to both groups in several characters, the ontogenetic sequence of the genus is different in many ways.     

Microsporogenesis in Pinus sylvestris. VI. Exine and tapetal development during the tetrad period

Coverage is of microspore tetrad period from end of cytokinesis to introduction of endexine in Pinus sylvestris. The ectexine of aperture, cap zone and sacci and the endexine are initiated while microspores are in the tetrad condition and enveloped in callose. Ectexine patterning including considerable expansion of sacci develops prior to the initiation of the endexine. Alveoli, sacci and alveoli within sacci are initiated by cytoplasmic invaginations which are sites of uptake of cell surface coat (glycocalyx) along with nutrients bound to the glycocalyx. Applications of tracers show that glycocalyx elements bind to cations and transport them to the cytoplasm. From the beginning of exine formation these invaginations are largest in the regions of future sacci and very small in the aperture. As growth progresses cytoplasm surrounding invaginations partially retracts, but callose contact is retained. Thus, these invaginations become callose covered hemispheroids (alveoli) that are “open” to the cell surface proximally and covered by callose distally but only partially so at the sides of the “cup‐shaped” alveoli. Until introduction of the endexine part of the alveolar‐sides are made up of cytoplasmic protrusions which contact the callose protrusions, even across sacci expanded more than 3 μm. Glycocalyx elements become aligned on the inner surface of the callosic alveoli and are sites for sporopollenin accumulation. The template for endexine components consists of glycocalyx elements that become aligned near the plasma membrane. Our observations indicate that uptake from the loculus to the microspore cytoplasm changes after introduction of the endexine. Henceforth, uptake is assisted by the endexine, as shown by tracers. Tapetal cells undergo two periods of hyperactivity during the period covered. Hyperactivity took place at the beginning of uptake by microspores and during endexine formation. The extra tapetal lamellation and its tapetal markers begin to exhibit the intense staining, after endexine initiation.     

An integral insight into pollen wall development: involvement of physical processes in exine ontogeny in Calycanthus floridus L., with an experimental approach

We aimed to understand the underlying mechanisms of development in the sporopollenin-containing part of the pollen wall, the exine, one of the most complex cell walls in plants. Our hypothesis is that distinct physical processes, phase separation and micellar self-assembly, underpinexine development by taking the molecular building blocks, determined and synthesised by the genome, through several phase transitions. To test this hypothesis, we traced each stage of microspore development in Calycanthus floridus with transmission electron microscopy and then generated in vitro experimental simulations corresponding to every developmental stage. The sequence of structures observed within the periplasmic space around developing microspores starts with spherical units, which are rearranged into columns to then form rod-like units (the young columellae) and, finally, white line centred endexine lamellae. Phase separation precedes each developmental stage. The set of experimental simulations, obtained as self-assembled micellar mesophases formed at the interface between lipid and water compartments, was the same: spherical micelles; columns of spherical micelles; cylindrical micelles; and laminate micelles, separated by gaps, resembling white-lined lamellae. Thus, patterns simulating structures observed at the main stages of exine development in C. floridus were obtained from in vitro experiments, and hence purely physicochemical processes can construct exine-like patterns. This highlights the important part played by physical processes that are not under direct genomic control and share influence on the emerging ultrastructure with the genome during exine development. These findings suggest that a new approach to ontogenetic studies, including a consideration of physical factors, is required for a better understanding of developmental processes.     

Sporoderm ultrastructure and development in Aristolochia manshuriensis Komarov (Aristolochiaceae)

Pollen ontogeny and sporoderm development in Aristolochia manshuriensis were studied for elaboration of the inaperturete pollen ontogeny in Aristolochia. Despite the formation of apertures in the tetrad period, the sporoderm in A. manshuriensis becomes inaperturate at the end of the free microspore period. A similar immature exine is also detected in A. macrophylla. Variants of aperture formation in the tetrad period in A. manshuriensis or formation of a polar aperture in the free microspore period in A. clematitis are associated with types of microsporogenesis. The ectexine and endexine in A. manshuriensis are formed over a longer time and reached much greater thickness than those in A. clematitis. The endexine and intine in A. manshuriensis do not reach a mature state, similar to A. clematitis. The exine of A. manshuriensis cracks, releasing a pollen tube enveloped by the intine. This fact does not hinder the functioning of the male gametophyte of A. manshuriensis.     

Comparative exine development from the post-tetrad stage in the early-divergent lineages of Ranunculales: the genera <Emphasis Type="Italic">Euptelea</Emphasis> and <Emphasis Type="Italic">Pteridophyllum</Emphasis>

Studies of pollen wall development produce a great deal of morphological data that supplies useful information regarding taxonomy and systematics. We present the exine development of Euptelea and Pteridophyllum, two taxa whose pollen wall development has never previously been studied using transmission electron microscopy. Both genera are representatives of the two earliest-diverging families of the order Ranunculales and their pollen data are important for the diagnosis of the ancestral pollen features in eudicots. Our observations show these genera are defined by having microechinate microreticulate exine ornamentation, perforate tectum, columellate morphology of the infratectum and the existence of a foot layer and endexine. The presence of lamellations is detected during the early stages of development in the nexine of both genera, especially in the apertures. Euptelea presents remains of the primexine layer during the whole maturation process, a very thin foot layer, and a laminate exinous oncus in the apertural region formed by ectexine and endexine elements. Pteridophyllum has a thicker tectum than Euptelea, a continuous foot layer and a thicker endexine. In the apertures, the exinous oncus is formed by islets and granules of endexine, in contrast to the Euptelea apertures. The secretory tapetum produces orbicules in both genera, but they have different morphology and electron-density. Comparisons with pollen data from related orders and families confirm the ancestral states for the pollen of eudicots proposed in previous studies: reticulate and echinate surfaces, columellate infractectum and a thin foot layer relative to the thickness of the ectexine. According to our observations, we propose considering the possibility of a polymorphic state for the aperture number in the ancestor of Ranunculales, and suggest the development of orbicules as the ancestral state in this order.     

The Structure of Pollen Wall and Its Relation to Po41en Aggregation in Cymbidium goeringii (Rchb. F.) Rchb. F.

The pollen wall of tetrads located in different positions of a mature pollinium of Cymbidium goeringii was examined with the electron microscope, and the compositions of wall materials were also tested with different histochemical methods. In all tetrads of a pollinium, the pollen wall can be distingished into an exine and an intine, but the exine may be varied greatly according to the tetrad position in a pollenium. The part of the pollen wall (the outer wall) of the external tetrads, lying close, to the tapetum, is composed of two layers, i.e. the exine, and the intine. Theexine consists of tectum, granulate ectexine and endexine, without foot layer. The intine is cellulose in nature. In the outer wall between different groups of: tetrads and in the inner wall within an individual tetrad, the structure of ectexine becomes simple and the deposition of sporopollenin is roduced The degree of reduction of ectexine nicreases from the outer to inner tetrads in several external layers of a pollinium, and even the internal tetrads have a reduced ectexine or lack of it. The present study also demonstrates that the mechanism of pollen aggregation into a pollinium is built on a combined effect of the following features: (1) connected bridges formed' by intine between two pollens within a tetrad, (2) formation of cytoplasmic channels between two pollens within a tetrad, (3) incomplete cell wall formation within a tetrad, (4) little size of tetrads and compact arrangement of mature tetrads and (5) a sticky viscin material surrounded on the outside of a pollinium.     

The fine morphology of pollen grains from the pollen chamber of a supposed ginkgoalean seed from the Middle Jurassic of Uzbekistan (Angren locality)

Pollen grains of Cycadopites-type were found in the pollen chamber of a supposed ginkgoalean seed Allicospermum sp. from the Middle Jurassic deposits of Uzbekistan (Angren locality). The pollen grains were studied with help of LM, CLSM, SEM, and TEM. All pollen grains show the identical morphology and exine ultrastructure allowing us to suppose the same botanical affinity. The pollen morphological data do not contradict the ginkgoalean interpretation of the seed; therefore, the pollen grains and the seed most probably did belong to the same parent plant. The pollen grains are monosulcate, the non-apertural surface is nearly psilate, with low short elements, which are occasionally scattered over the surface or more densely distributed. The aperture and adjacent areas appear to bear more distinct sculpturing. The ectexine is composed of a prominent solid tectum, a thin infratectum, and a thin foot layer. The infratectum is formed of one row of alveolae, which are more voluminous laterally, where the ultrastructure is more easily understandable. The endexine is multilamellate, although it is evident only in some regions of stained sections. Towards the aperture the ectexine becomes gradually thinner; over the aperture no sublayers can be discerned within the ectexine. The ectexine of the apertural region repeatedly varies in thickness, reflecting a sculpturing surface of this region. The obtained data contribute to the knowledge about the exine ultrastructure of ginkgoaleans; nonetheless, a TEM study of ginkgoalean pollen grains extracted from pollen organs is still highly desirable. We also considered pluses and minuses of CLSM: it failed to substitute SEM, since the surface pattern under study was too fine, but demonstrated the general morphology of the pollen grains under study better than conventional LM. The possibility of viewing virtual sections of any area of the pollen grain was profitable for later interpretation of TEM sections. CLSM would give better results in interpreting relatively large palynological objects with distinct sculptural elements, a complicated architecture, variously arranged appendages, or possessing cameras.