POLLEN WALL ULTRASTRUCTURE OF SELECTED DISPERSED MONOSULCATE POLLEN FROM THE CENOMANIAN,DAKOTA FORMATION,OF CENTRAL USA

Seven dispersed monosulcate pollen taxa from the Dakota Formation of Minnesota, Nebraska, and Kansas were examined ultrastructurally. Rugubivesiculites rugosus has gymnosperm affinities based on its anasulcate aperture and the presence and nature of the formation of sacci. Stellatopollis sp. has exine sculpturing restricted to taxa with angiosperm affinities and is monosulcate. The affinities of the other five monosulcate taxa are uncertain and the exines are tectategranular. The sulcus in many of the remaining five taxa are flanked by small flange-like sacci. These five taxa have features found in gymnosperms and also some features of primitive extant angiosperms. The combination of characters of the pollen types presented here does not entirely agree with our current concept of primitive pollen characters as understood from extant ranalean angiosperms.     

Studies of Pollen Morphology and Its Systematic Position in the Order Piperales

The pollen morphology of 25 species and 10 genera of Piperales (Chloranthaceae, Piperaceae and Saururaceae) has been examined under light microscope, of which 7 species were observed under scanning electron microscope and 1 species, Hedyosmum orentale Merr. & Chun transmision electron microseope. Three principal types of pollen were found: anasulcate (mostly) (sometime trichotomosulcate), inaperturate (partly) and multicolpoidate (partly). The present article has discussed the palynological data mainly in relation to the classification and the systematic position of Cbloranthaceae and also deals with the systematic position of the order Piperales. The present author agrees to put the family Chloranthaceae into the order Piperales. Because this family differs from Piperaceae and Saururaceae in pollen morphology, therefore, Chloranthaceae should raise to the level of suborder. Among three families of the order Piperales, the present author considers Chloranthaceae to be the most primitive family, on account of the following reasons: 1. The family Chloranthaceae shows the characteristics of primitive entomophilous plants in the sculpture of exine, while in the other two families, Piperaceae and Saururaceae, their exine is almost smooth and represents wind-pollenated plants; 2. Pollen of the family Chloranthaceae are larger than those of Piperaceae and Saururaceae; 3. The fossil pollen Clavatipollenites has been proved to be one of the most primitive angiosperms on the earth, that it is known, it occurred in the early Cretaceous, and at that time ferns and gymnosperms were predominant, while the Chloran- thaceae has already existed at that time; 4. Sarcandra of Chloranthaceae possesses the characters of a vesselless secondary xylem and a delayed development of embryo. Thus, Chloranthaceae would be considered as the most primitive family in the order Piperales. The systematic position of the order Piperales is also discussed. Itutehinson makes a point that order Ranales is more primitive than Piperales, and his system is arranged in the following order: Ranales → Piperales → to climax family Chloranthaceae. This view-point, however, is net supported by the palynological data. Pollen morphology shows that Piperales is more primitive than Ranales, because the pollen in Piperales possess the ancient aperture type of Pteridospermes, i.e., the type of anasulcate aperture is prevailing in Piperales, moreover, pollen grains of Ranales are mainly tricolpate type, and tricolpate pollen is a characteristic of typical angiosperms. In addition, the Piperales possesses a series of characters that are common among monocots, but rare among dicots. As the divergence between dicer and monocot took place in the early Cretaceous, their ancestor possesses common chararcters both of dicots and monocots while the extant Piperales still possess many characters of monocots that indicate it is much nearer to the point of divergence, and it explains that the Piperales is closely related to the ancestor of monocots and dicers Piperales, therefore, is more primitive than Ranales.     

Why does some pollen lack apertures? A review of inaperturate pollen in eudicots

Apertures are key characters of pollen grains with systematic importance in angiosperms. They function as sites for pollen tube exit, water uptake, transfer of recognition substances and accommodation of volume changes. Not all pollen has apertures; inaperturate pollen (lacking obvious apertures) characterizes many angiosperm groups, especially in early divergent angiosperms and monocots, but also eudicots. In order to expand our knowledge of the systematic distribution, possible functional significance and development of inaperturate pollen in angiosperms, this review focuses on inaperturate and cryptoaperturate (with hidden apertures) pollen in the large eudicot clade, which comprises about 75% of present‐day angiosperm species. It includes new TEM observations of inaperturate pollen from four exemplar taxa selected from different parts of the eudicot phylogeny. Two categories of inaperturate (including cryptoaperturate) pollen occur in eudicots. (1) Sterile attractant or feeding pollen associated with functional dioecy has evolved iteratively at least six times in conjunction with complex breeding systems in the core eudicots. (2) Fertile pollen has evolved numerous times independently throughout eudicots, though generally in a relatively small number of individual taxa. Notable exceptions are the petaliferous crotonoid Euphorbiaceae s.s., in which fertile inaperturate pollen occurs in c. 1500 species, and two subfamilies of Apocynaceae s.l. (Secamonoideae and Asclepiadoideae) with c. 2500 species with fertile inaperturate pollen in pollinia. Fertile inaperturate pollen is sometimes (but not always) associated with an aquatic habit, parasitism, insectivory, heterostyly, anemophily or pollinia. Most fertile inaperturate pollen has a thin exine, or the exine is largely restricted to isolated components (muri, protuberances, subunits) separated by thinner areas which probably function as apertures. In cryptoaperturate pollen, the aperture is covered by continuous exine which probably has a protective function, similar to an operculum. Developmentally, inaperturate pollen is not associated with any particular tetrad type or meiotic spindle orientation (unlike some apertures) due to the absence of a colpal shield of endoplasmic reticulum or other organelles and hence is independent of microsporogenesis type. The lack of a colpal shield during the tetrad stage of development permits complete deposition of first primexine and then exine around each microspore, possibly mediated by the action of the DEX1 protein. © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 155 , 29–48.     

EVOLUTION OF EXINE STRUCTURE IN THE POLLEN OF PRIMITIVE ANGIOSPERMS

Diversity in the structure of the exine in 35 families of the ranalean complex is compared through a series of representative scanning electronmicrographs, and evolutionary trends in exine structure of primitive angiosperms are outlined, along with discussion of the significance of these data for understanding the evolution of exine structure in flowering plants as a whole. In order to reduce ambiguity in the palynological literature, it is suggested that persons undertaking light microscope studies of unstained, acetolyzed pollen grains adopt the morphological terms sexine-nexine in describing pollen wall layers while restricting their use of the chemically defined terms ektexine-endexine largely to pollen studies carried out with the transmission electron microscope. This study emphasizes that a clear understanding of the palynological concept of structure versus sculpturing is a necessary prerequisite for the taxonomic/ phylogenetic use of pollen wall morphology. Finally, data from investigation of a number of ranalean families of primitive angiosperms support the conclusion that the direction of a recurrent and major evolutionary trend in exine structure of flowering plants proceeds from pollen that is tectate-imperforate to tectate-perforate pollen to semitectate pollen, and more rarely, to pollen grains that are intectate.     

我国罂粟科(Papaveraceae)植物的花粉形态

本文用光学显微镜对我国16属60种5变种罂粟科植物花粉形态进行了系统地观察研究。其中对15属27种进行了扫描观察。本科花粉形态为多类型。3沟为原始的、基本的花粉类型。根据萌发孔的形状、数目和特征,将罂粟科的花粉分为2沟,3沟,4-6沟、散沟、散孔和无萌发孔6个花粉类型。根据孢粉学资料,本文对绿绒蒿属(Meconopsis Vig.)花粉粒和其它器官的进化趋势,罂粟科在进化上的亲缘关系和本科花粉萌发孔的演化趋向等问题进行了讨论。     

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.     

Palynological data on Illiciaceae and Schisandraceae confirm phylogenetic relationships within these two basally-branching angiosperm families

Illiciaceae and Schisandraceae, together with other members of Austrobaileyales have been identified as one of the earliest diverging lineages of angiosperms, within the ANITA grade. The specialized Illiciaceae and Schisandraceae comprise a clade defined by apomorphic characters including pollen grains with three or six colpate apertures. In both these families, pollen apertures and exine sculpture were found to be very informative when considered in the context of recent understanding of evolutionary patterns. In the current study, pollen grains of 21 taxa from Illiciaceae and Schisandraceae were investigated. These data, together with palynological data for taxa previously studied, were mapped into recent molecular phylogenetic trees to re-evaluate the existing classification and phylogenetic relationships in the two families. Palynological data were found to be relatively congruent with recent molecular phylogenies, while traditional delimitations of infra-generic taxa were somewhat conflicting and did not reflect phylogeny and evolution. The evolution of pollen morphology in the two families, together with other members of Austrobaileyales, is discussed in comparison with the molecular phylogenies.     

Aperture Orientation in Laurelia Pollen (Atherospermataceae Syn. Subfamily Atherospermoideae of Monimiaceae)

The development of the encircling aperture of pollen grains or the New Zealand species Laurelia novae-zelandiae is described by means of light and scanning electron microscopy. Some observations are also reported from pollen development and structure in the South American species Laurelia sempervirens, mature pollen of which can be distinguished from that of L. novae-zelandiae. The aperture begins to develop while pollen of L. novaezelandiae is still in tetrads, and passes through the distal and proximal poles of each grain. The widest parts of the mature aperture are at and near what are morphologically the distal and proximal poles of each grain. No such meridionosulcate type of pollen was recorded in a recent survey of pollen aperture types of primitive angiosperms and contradicts a recent report that pollen in the Atherospermataceae had equatorially aligned apertures.     

中国紫葳科花粉形态的研究

对紫葳科18属27种花粉进行了光镜和扫描电镜的观察,其中有些种是第一次观察或报道。紫葳科花粉明显地属多型花粉,萌发孔类型和纹饰特征多种多样。萌发孔基本上可分为3（－4）沟,3（－4）孔沟,3孔,多沟及散沟等类型;纹饰则可分为皱波状饰,穴状纹饰,穴－网状纹饰,网状纹饰,刺状纹饰以及表面光滑等。值得注意的是在菜豆树属Rademoachera Zoll.et Moritzi中,菜豆树R．sinica和滇菜豆树R.yunnanensis虽为同一个属,但其萌发孔类型和纹饰特征截然不同,菜豆树花粉为3孔,具网状纹饰;而滇菜豆树花粉为3孔沟,具穴状纹饰。就花粉形态学而言,紫葳科也许是多元发生的。     

蜘蛛抱蛋亚族的花粉形态

本文研究并报道了百合科蜘蛛抱蛋亚族的花粉形态,主要结论如下：1．开口箭属含两种花粉类型即：(1)单槽、椭圆形;(2)无萌发孔、球形。具单槽、椭圆形花粉的种类含单型性核型,花形较小,柱头小而花柱短,雄蕊高于柱头;无萌发孔、球形花粉的种类则为单型性核型,花形较大,柱头常高度膨大,雄蕊常着生在花被筒基部。2．万年青属花粉为椭圆形,具单槽;蜘蛛抱蛋属花粉则为球形,无萌发孔;开口箭属花粉一部分与万年青属相似,另一部分则与蜘蛛抱蛋属雷同。3．蜘蛛抱蛋亚族中最原始的花粉类型为单槽椭圆形,具孔状或网状外壁纹饰。球形,无萌发孔,外壁具皱波状、瘤状或芽孢状纹饰的花粉为派生类型。4．蜘蛛抱蛋亚族种下花粉形状、萌发孔类型和外壁纹饰基本稳定,可作为分种特征。     

Pollen evolution in cordaites and early conifers

Changes in three pollen grain characters: number of sacci, infrastructure of sacci, and number and position of apertures, are considered against a phylogenetic background of cordaites and early conifers reconstructed on the basis of female fructifications. Bisaccate pollen grains with double germination developed most probably independently in Majonicaceae and Voltziaceae, whereas the most primitive conifers produced monosaccate prepollen with a single proximal aperture. Another lineage resulting in true pollen with a single distal aperture can be observed within cordaites. The development of saccus infrastructure among early pinopsids was directed from eusaccus to protosaccus rather than vice versa. In any case, the eusaccus of modern conifers is not homologous to that of cordaites and its origin is to be elucidated. The pollen characters suggest a very early separation of conifers from cordaites, approximately simultaneous with the division of cordaites into three families.     

Formation and function of a new pollen aperture pattern in angiosperms: The proximal sulcus of Tillandsia leiboldiana (Bromeliaceae)

Pollen grains are generally surrounded by an extremely resistant wall interrupted in places by apertures that play a key role in reproduction; pollen tube growth is initiated at these sites. The shift from a proximal to distal aperture location is a striking innovation in seed plant reproduction. Reversals to proximal aperture position have only very rarely been described in angiosperms. The genus Tillandsia belongs to the Bromeliaceae family, and its aperture pattern has been described as distal monosulcate, the most widespread aperture patterns recorded in monocots and basal angiosperms. Here we report developmental and functional elements to demonstrate that the sulcate aperture in Tillandsia leiboldiana is not distal as previously described but proximal. Postmeitotic tetrad observation indicates unambiguously the proximal position of the sulcus, and in vitro germination of pollen grains confirms that the aperture is functional. This is the first report of a sulcate proximal aperture with proximal germination. The observation of microsporogenesis reveals specific features in the patterns of callose thickenings in postmeiotic tetrads.     

FINE STRUCTURE OF MISTLETOE POLLEN. III. LARGE-FLOWERED NEOTROPICAL LORANTHACEAE AND THEIR AUSTRALIAN RELATIVES

Pollen of all large-flowered neotropical loranthaceous genera and related Australian taxa (Nuytsia, Atkinsonia) were examined in the light, scanning, and transmission electron microscopes. Trilobate, deeply concave, peroblate or oblate pollen shapes are basic within the complex; triangular, slight convex, suboblate shapes are more derived features. The non-fixiform pollen of Atkinsonia is unique within the family. Most of the large-flowered genera possess exclusively syncolpate aperture types. However, among Psittacanthus species tricolpate, diplosynrugate and diplorugate apertures are also present, representing derived types for the complex. The inaperturate pollen of Atkinsonia appears to have developed independently within the family. Pollen sculpturing is typically non-uniform, i.e., there are pronounced sculpturing differences in polar and equatorial regions. Uniformly sculptured pollen is restricted to Atkinsonia, Ligaria and some species of Psittacanthus. Ultrastructurally, most exine modifications have occurred in the equatoral ektexine. Here the basic organization ranges from essentially columellaless to columellate, the latter the more derived condition. Endexine is typically thick and stratified in polar areas, thin and lamellate in peripheral and apertural regions. Similar pollen morphologies of Nuytsia and Gaiadendron support the idea of a transoceanic evolutionary connection between the Old and New Worlds. Pollen characters show Gaiadendron to be the most primitive and Psittacanthus the most advanced among the large-flowered neotropical genera.     

甘薯属10种植物花粉形态扫描电镜观察

对用于育种研究的甘薯属(Ipomoea)10种植物花粉形态进行扫描电镜观察研究,以补充该属植物花粉形态学信息. 结果表明,甘薯属植物花粉粒的形状均为近球形,体积较大;花粉萌发孔多为散孔,少数萌发孔不明显;表面纹饰有3种:刺状类型、网状带刺类型和网状类型.根据花粉粒大小、刺的长短、萌发孔特征和表面纹饰类型等可以把这10种植物区分开来,同时发现I.wrightii与已有报道存在较大差异.     

Pollen Morphology in the Subtribe Aspidistrinae (Liliaceae s. l.)

Pollen morphology in the subtribe Aspidistrinae is reported and the main viewpoints are summarized as follows: (1) Two major types of pollen grains, i.e., monosulcate,ellipsoidal pollen and inaperturate, spheroidal pollen, are identified in the genus Tupistra. The species with monosulcate, ellipsoidal pollen usually have monomodal karyotypes, brightcolored flower, smaller stigma, and stamens inserted at upper or middle part of perianth tube, while the inaperturate and spheroidal pollen is always found in the species with bimodal karyotypes, dingy-colored flower, swollen stigma and stamens inserted at the base of perianth tube. (2) Pollen grains in the genus Rhodea are monosulcate and ellipsoidal, while those in the genus Aspidistra are inaperturate and spheroidal, correspondingly similar to the second type of pollen grains in the genus Tupistra. (3) The most primitive pollen in the subtribe Aspidistinae is regarded as monosulcate and ellipsoidal, having perforate or reticulate exine sculpture. The inaperturate, spheroidal pollen with verrucate, gemmate or rugulate exine sculpture is considered derived; (4) Unlike those reported in other groups of the family Liliaceae, the infraspecific pollen shape, aperture type and exine sculpture in Aspidistrinae are basically stable and may serve as a taxonomic character.     

Comparative morphology of monocot pollen and evolutionary trends of apertures and wall structures

Data on pollen aperture and wall ultrastructure are reviewed for the monocots. New ultrastructural data on 30 taxa representing 18 monocot families are also presented. The evolutionary trends of apertures and pollen wall structure are discussed and it is proposed that the evolutionary trends of pollen apertures and wall structure in the monocots parallel those proposed by Walker for the dicots. However, the importance of these trends differ for monocots, suggesting that the selective pressures affecting pollen wall and aperture evolution in dicots and monocots have been similar, but with different emphasis.     

Pyrene and pollen observations in the pantropical genus Geophila (Rubiaceae-Psychotrieae)

Pyrene and pollen morphology of several neo- and paleotropical Geophila species is documented and discussed. Pyrenes show distinct morphological variation. They have one central or several adaxial ribs and are straight or twisted. Neotropical species only have twisted pyrenes, whereas Madagascan species have straight ones. Both pyrene types occur on the tropical African mainland and in tropical Asia. The genus is eurypalynous, including inaperturate, colpate and pantoporate pollen. The paper includes the first record of operculate pollen in the Rubiaceae, viz. in Geophila repens var. repens . Pyrenes as well as pollen morphology indicate that the varieties of the pantropical species G. repens need recognition at the specific level.     

APERTURE EVOLUTION IN THE GENUS PTYCHOPETALUM BENTH. (OLACACEAE)

The genus Ptychopetulum (Olacaceae), disjunct between South America (2 spp.) and Africa (2 spp. West Africa; I sp. Rhodesia), exhibits predominantly diploporate pollen among the West African species and predominantly triporate pollen among the South American taxa. An analysis of aperture evolution suggests two possible schemes of aperture evolution for the genus. Scheme I avers the independent origin of both triporate and diploporate apertures from a basic brevicolpate condition while scheme II proposes a basic diploporate condition in which the diploporate apertures have migrated to the equator, fused to form brevicolpi and have ultimately become reduced to pores. Pollen evidence in support of both schemes is discussed. The distribution of aperture types within the genus also suggests West Africa as the original site of aperture evolution and South America as the second area of aperture development with types traceable to West African stock.     

Release of developmental constraints on tetrad shape is confirmed in inaperturate pollen of Potamogeton

Background and Aims

Microsporogenesis in monocots is often characterized by successive cytokinesis with centrifugal cell plate formation. Pollen grains in monocots are predominantly monosulcate, but variation occurs, including the lack of apertures. The aperture pattern can be determined by microsporogenesis features such as the tetrad shape and the last sites of callose deposition among the microspores. Potamogeton belongs to the early divergent Potamogetonaceae and possesses inaperturate pollen, a type of pollen for which it has been suggested that there is a release of the constraint on tetrad shape. This study aimed to investigate the microsporogenesis and the ultrastructure of pollen wall in species of Potamogeton in order to better understand the relationship between microsporogenesis features and the inaperturate condition.

Methods

The microsporogenesis was investigated using both light and epifluorescence microscopy. The ultrastructure of the pollen grain was studied using transmission electron microscopy.

Key Results

The cytokinesis is successive and formation of the intersporal callose wall is achieved by centrifugal cell plates, as a one-step process. The microspore tetrads were tetragonal, decussate, T-shaped and linear, except in P. pusillus, which showed less variation. This species also showed a callose ring in the microsporocyte, and some rhomboidal tetrads. In the mature pollen, the thickening observed in a broad area of the intine was here interpreted as an artefact.

Conclusions

The data support the view that there is a correlation between the inaperturate pollen production and the release of constraint on tetrad shape. However, in P. pusillus the tetrad shape may be constrained by a callose ring. It is also suggested that the lack of apertures in the pollen of Potamogeton may be due to the lack of specific sites on which callose deposition is completed. Moreover, inaperturate pollen of Potamogeton would be better classified as omniaperturate.Key words: Alismatales, callose, microsporogenesis, pollen aperture, Potamogeton illinoensis, P. polygonus, P. pusillus, tetrad shape     

Pollen Morphology of the Family Zingiberaceae in China—Pollen Types and Their Significance in the Taxonomy

Pollen morphology of 89 species and 3 varieties belonging to 18 genera (out of 150 spcies in 20 genera) of Zingiberaceae in China was studied under both light microscope and scanning electron microscope. Pollen grains of Zingiberaceae are spherical, subspherical, ovoid and prolate, 36-225 μm in size, nonaperturate or aperturate (spiraperturate, porate). Pollen grains are almost not resistant to acetolysis. The wall is composed of a very thin exine and a thick intine. The exine is psilate, spinate, cerebelloid-areolate, striate, verrucate and foveolate. According to the presence or the absence of aperture and differential ornamentations, two types and six subtypes are recognized: I. The type Nonaperturate: (85 species and 3 varieties in 18 genera). Four subtypes can be recognized within the type based on the characteristics of the exine sculpture. These are: (1) The subtype Psilate, in which, the exine is nearly smooth (including: Hedychium, Curcuma, Kaempferia, Caulokaempferia coenobilis, Boesenbergia rotunda, Stahlianthus, Amomum compactum, Etingera, Hornstedtis, Rhynchanthus). (2) The subtype Spinate, which comprises two groups: (A) The group Short-spinate, pollen grains with smaller spines (Globba), (B) The group Long-spinate, pollen grains with longer spines (Alpinia, Amomum, Plagiostachys, Roscoea, Cautleya, Boesenbergia fallax, Caulokaempferia yunnanensis). (3) The subtype Cereblloid-areolate, pollen grains of which are spherical or subspherical, with cerebelloid sculpture (Zingiber Sect. Zingiber). (4) The subtype Striate, pollen grains of which are prolate or oliveshaped, and striate (Zingiber Setc. Cryptanthium). II. The type Aperturate, in which pollen grains are acetilysis-resistant and possess distinct apertures (mixed colpate-porate or forate), including two subtypes: (1)The subtype Mixed colpate and Porate. Pollen grains are both 3-colpate and 1-3-porate, and usually with one long spiral, two short (straight or slightly curved) colpi and 1-3-poris. The exine is verrucate or not, nearly sinuolate (Costus speciosus, C. tonkinensis, C. lacerus). (2) The subtype porate, whose grains are 6-8-porate and exine is foveolate (Costus megalobractea). The taxonomic significance of the pollen types in the family Zingiberaceae is also discussed.