Fossil floral structures of a basal angiosperm with monocolpate,reticulate-acolumellate pollen from the Early Cretaceous of Portugal

Distinctive monocolpate and reticulate-acolumellate pollen grains with a coarse, loosely attached reticulum have long been known as a conspicuous element of many palynological assemblages from the Early and mid-Cretaceous. These grains are now described in situ in staminate structures and on the surface of pistillate organs from two Early Cretaceous (Barremian or Aptian) mesofloras from Portugal (Vale de Agua and Buarcos). Staminate organs include a staminate axis with spirally arranged stamens and many isolated stamens. Stamens consist of a short filament, a dithecate, tetrasporangiate anther, and a short apical extension of the connective. Anther dehiscence is extrorse by longitudinal slits and in situ pollen is monocolpate, semi-tectate with a coarse, loosely attached reticulum composed of narrow muri with a spiny ornamentation. The infratectal layer of the pollen wall is thin, granular, and lacking columellae; and the foot layer is distinct. The endexine is thin, except under the aperture where it is thick. The pistillate organs are minute consisting of a simple unilocular ovary containing a single thin-walled seed. Associated with staminate and pistillate structures are many coprolites consisting almost exclusively of pollen grains of this distinctive type. The staminate and pistillate organs are not found in organic connection, and two new genera are established to accommodate the new floral structures: Pennistemon comprising the staminate structures and Pennicarpus comprising the pistillate structures. A new genus, Pennipollis, is also established for the dispersed grains, based on the type species Peromonolites peroreticulatus Brenner, since no appropriate genus has yet been described for these acolumellate grains. Features of the pollen grains strongly indicate affinity with members of the Alismatales and characters of the mesofossils also support this assignment. This is the first record of putative monocots in the early Cretaceous based on combined pollen and floral features.     

MORPHOLOGICAL AND EMBRYOLOGICAL STUDIES ON THE LEMNACEAE. I. THE FLORAL STRUCTURE AND GAMETOPHYTES OF LEMNA PAUCICOSTATA

Maheshwari, Satish C., and R. N. Kapil. (U. Delhi, Delhi, India.) Morphological and embryological studies on the Lemnaceae. I. The floral structure and gametophytes of Lemna paucicostata. Amer. Jour. Bot. 50(7): 677–686. Illus. 1963.—In Lemna paucicostata, a locally occurring member of the Lemnaceae, the plant body is represented by a frond which is devoid of lignified elements. The root shows a winged root sheath but does not have root hairs. There are no distinctive layers like the endodermis and xylem. The male archesporium is hypodermal and differentiates normally as in other angiosperms into parietal and sporogenous layers. The tapetum is single-layered and plasmodial. The partition walls of the anther are not derived by sterilization of the sporogenous cells as believed earlier. The microspore tetrads are isobilateral and decussate, the meiotic divisions being successive. The pollen grains are shed at the 3-celled stage. The ovary contains a single hemianatropous, bitegminal and crassinucellar ovule. The development of the embryo sac conforms to the Allium type.     

Vertical variation in urticaceae airborne pollen concentration

The pollen contents at different heights (1.5 and 15 m) of species of the Urticaceae family have been studied by sampling with Hirst type volumetric samplers. In order to achieve this, the two pollen types belonging to this family have been treated separately,Urtica urens-Parietaria sp. on the one hand andUrtica membranacea on the other, the latter having a smaller pollen grain. The results show that meteorological factors are bound to influence the behaviour of both these types of pollen in relation to height. With damp weather the pollen contents vary very slightly at different heights while when the weather is dry and calm, differences in pollen content at different heights become more significant. Nevertheless, when the atmosphere is stratified, the behaviour of each pollen type is different. The results show that, for most of the months considered, there is a higher pollen content ofU. membranacea at upper heights, whileU. urens-Parietaria sp. has higher levels of pollen content at a lower height. High temperatures, absence of rain and calm weather conditions favour the presence of convective phenomena which in turn create a favourable atmosphere for the vertical transportation of the small pollen grains ofU. membranacea, which are better represented in the samplers placed at 15 m.     

笔竹大小孢子发生及雌雄配子体发育研究

为探讨笔竹(Pseudosasa viridula)结实率低的原因,该文通过采用石蜡切片的方法结合显微技术对笔竹大小孢子发生及雌雄配子体的发育过程进行研究.结果表明:(1)笔竹的雄蕊多为3枚,极少有6枚,每枚花药具有4个花粉囊.(2)花药壁发育为基本型,由4层细胞构成,由外向内依次为表皮细胞、药室内壁细胞、中层细胞和绒...     

开口箭大小孢子发生及雌雄配子体发育

利用石蜡切片技术,对百合科植物开口箭(Tupistra chinensis Baker)大小孢子发生及雌雄配子体发育进程进行胚胎学观察分析,以明确开口箭胚胎发育的特征,为百合科植物的研究提供生殖生物学依据。结果表明:(1)开口箭花药具有4个药室,花药壁的发育方式为基本型,由表皮、药室内壁、中层及绒毡层组成;绒毡层发育类型为分泌型,到四分体花药阶段绒毡层细胞开始解体退化,花药成熟时完全消失。(2)花粉母细胞减数分裂为连续型,依次形成二分体、四分体,四分体为左右对称形;成熟花粉为2-细胞花粉,具单萌发沟。(3)子房3室,倒生型胚珠6枚,双珠被,薄珠心;在花部的分化早期,由珠心顶端表皮下方分化出雌性孢原细胞,孢原细胞经过一次平周分裂形成周缘细胞和造孢细胞,造孢细胞发育为大孢子母细胞;大孢子母细胞第一次减数分裂后形成二分体,珠孔端的二分体孢子退化,合点端的二分体孢子继续第二次分裂,形成两个子细胞依次发育为二核胚囊、四核胚囊和八核胚囊;开口箭的胚囊发育类型为葱型。     

Studies in the embryology ofHeliantheae (Compositae)

The embryology ofLagascea mollis, Zinnia angustifolia andGalinsoga parviflora has been studied. The anther archesporium is hypodermal and consists of a single row of 6–8 cells, there are two layers below the epidermis of the anther and a periplasmodial tapetum. Ripe pollen grains are tricolpate and 3-celled. The ovary contains a single ovule, but in a few cases ofGalinsoga parviflora two ovules have been found. The female archesporium is unicellular, but sometimes more than one archesporial cell occurs inLagascea mollis andZinnia angustifolia. The embryo sac development is of the Polygonum type, the synergids are hooked, antipodal cells show great variation. The endosperm development is cellular inGalinsoga parviflora and peripheral layer persists in the mature seed. The embryo development conforms to the Senecio variation of the Asterad type. The pericarp structure and cmbryological features support the disputed systematic position ofLagascea withinHeliantheae. Part of a Ph.D. thesis, accepted at Andhra University.     

Pollenkitt is lacking inGnetatae:Ephedra andWelwitschia; further proof for its restriction to the angiosperms

Three members of theGnetatae (Ephedra campylopoda, E. americana, Welwitschia mirabilis) were investigated by TEM and SEM with respect to their anther tapetum and pollen development. In all three species pollenkitt is lacking. The pretended pollen stickiness thus does not depend on pollenkitt. Considering former observations one can now clearly state that pollenkitt is missing in all recent gymnosperm classes (both anemophilous and ± entomophilous). Pollenkitt thus is restricted and ± omnipresent within the angiosperms, where it represents one of the most important components of the entomophily syndrome. This can be regarded as important proof for the hypothesis that the angiosperms are a single coherent phylogenetic group.     

A COMPARISON OF CLEISTOGAMOUS AND CHASMOGAMOUS FLORAL DEVELOPMENT IN COLLOMIA GRANDIFLORA DOUGL. EX LINDL. (POLEMONIACEAE)

An individual of Collomia grandiflora typically produces both closed or cleistogamous (CL) and open or chasmogamous (CH) flowers. The developmental origin of these dimorphic floral types within a plant was investigated using histological techniques, allometric relationships, and scanning electron microscopy. Prior to archesporal cell stage in the anthers, CL and CH meristems are indistinguishable. In the CL anther, an absence of ventral locule cell differentiation together with a shorter period of time between archesporial cell differentiation and meiosis in the two dorsal locules results in accelerated anther dehiscence and a smaller mature anther size and pollen grain number. Divergence between the CL and CH patterns of corolla development is coincident with microspore mitosis in the CH anther. At this point, there is an increase in growth in corolla length relative to growth in calyx length in the CH flower which does not occur in the CL flower. Calyx and ovary development are similar in the two floral forms; however, ovary expansion due to fertilization occurs earlier in the CL flower as a result of precocious anther development and stigma receptivity. The hypothesis that anther differentiation may trigger organ growth rate changes and differentiation events in the flower and hypothetical roles for abscisic acid and gibberellin in modifying floral development in C. grandiflora are discussed.     

葱兰大小孢子发生和雌雄配子体发育及其系统学意义

采用常规石蜡切片技术,对石蒜科葱兰的花药壁发育、大小孢子的发生和雌雄配子体的发生过程进行了研究,并对葱兰属、石蒜科、百合科以及葱科的胚胎学特征进行比较讨论。结果表明:(1)葱兰花药四室,药壁由表皮、药室内壁、中层和绒毡层组成;药壁发育类型为单子叶型,绒毡层的类型为分泌型;花粉成熟时药室内壁径向加长并纤维状加厚,表皮宿存;小孢子母细胞在减数分裂过程中胞质分裂为连续型,小孢子四分体排列方式主要为四面体型,还有少数一些为左右对称型,成熟花粉为2-细胞型。(2)葱兰的雌蕊3心皮合生,子房下位,中轴胎座,3室,每室具2列倒生胚珠;胚珠双珠被,厚珠心,具蓼型胚囊。(3)葱兰属的胚胎学特征与石蒜科的其他种类存在较大的差异,如葱兰属花药壁发育为单子叶型,而石蒜科花药壁发育主要为双子叶型,但葱兰属的这些胚胎学特征却和百合科较为相似。     

Precocious pollen germination in Arabidopsis plants with altered callose deposition during microsporogenesis

Pollination is essential for seed reproduction and for exchanges of genetic information between individual plants. In angiosperms, mature pollen grains released from dehisced anthers are transferred to the stigma where they become hydrated and begin to germinate. Pollen grains of wild-type Arabidopsis thaliana do not germinate inside the anther under normal growth conditions. We report two Arabidopsis lines that produced pollen grains able to in situ precociously germinate inside the anther. One of them was a callose synthase 9 (cs9) knockout mutant with a T-DNA insertion in the Callose Synthase 9 gene (CalS9). Male gametophytes carrying a cs9 mutant allele were defective and no homozygous progeny could be produced. Heterozygous mutant plants (cs9/+) produced approximately 50% defective pollen grains with an altered male germ unit (MGU) and aberrant callose deposition in bicellular pollen. Bicellular pollen grains germinated precociously inside the anther. Another line, a transgenic plant expressing callose synthase 5 (CalS5) under the CaMV 35S promoter, also contained abnormal callose deposition during microsporogenesis and displaced MGUs in pollen grains. We also observed that precocious pollen germination could be induced in wild-type plants by incubation with medium containing sucrose and calcium ion and by wounding in the anther. These results demonstrate that precocious pollen germination in Arabidopsis could be triggered by a genetic alteration and a physiological condition.     

Genetic and environmental variation in anther,pollen and pistil dimensions in sesame

Measurements of anther (length, width, depth), pollen grain (percent fertility, polar diameter, equatorial diameter, polar diameter/equatorial diameter ratio, volume) and pistil (stigma length, style length, ovary length, total pistil length, stigma width, style width, ovary width) were taken on 12 diverse sesame (Sesamum indicum L.) genotypes on each of four collection dates in 1994. Highly significant differences among genotype means were obtained for all characters except polar diameter. Highly significant differences among environment (collection date) means were found for ten of the 15 characters measured. Highly significant genotype x environment interactions were obtained for all characters except anther length. For the anther characters measured, relatively high repeatability values were found, ranging from 99.8% for length to 87.6% for depth. For the pollen grain characters measured, the repeatability values ranged from 67.6% for percent fertility to 23.1% for polar diameter. For the pistil characters measured, the repeatability values ranged from 94.0% for style width to 49.6% for total pistil length. These results indicate that genotype and environment influence anther, pollen grain and pistil characters. Variation in some of these morphological aspects could influence the consistency and interpretation of male transmission studies on both the applied and evolutionary levels.     

Embryological Studies on Narcissus tazetta var. chinensis

Chinese narcissus (Narcissus tazetta var. chinensis Roem) blooms but has no seeds. Embryological studies on the species were conducted to discover the causes of its sterility. Its anther wall is composed of four layers of cells, and its tapetum is of the secretory type. The cytokinesis of microspore mother cells is of the successive type, and the tetrad is tetrahedral. During meiosis of microspore mother cells, some chromosomes lagged, and several micronuclei were found in tetrads. Only 27.7% of the pollen grains contained full cytoplasm, and 1.3% of them germinated in culture medium. No pollen grain, however, could germinate on the stigma. The ovary is trilocular with axile placenta, and the ovules are bitegmic, tenuinucellate, and anatropous. Its embryo sac is of the polygonum type. Most embryo sacs degenerated, and only about 4.5% of the ovules contained a normal embryo sac with an egg cell, two synergids, three antipodal, and a central cell containing two polar nuclei. One reason for the sterility of Chinese narcissus is the abnormality of microsporogenesis and megasporogenesis, in which only a few functional pollen grains and embryo sacs are produced. The other reason is that the pollen grains cannot germinate on the stigma. This paper was translated from Journal of Xiamen University (Natural Science), 2005, 44(1) (in Chinese)     

Embryology and systematic position ofLiparis (Orchidaceae)

The embryology ofLiparis paradoxa andL. rostrata has been studied. The young anther wall consists of an epidermis, endothecium, three middle layers and secretory tapetum with uninucleate cells. In the mature anther, two or three sub-epidermal layers develop fibrous thickenings. The anther wall development corresponds to the massive type. Simultaneous cytokinesis results in decussate, isobilateral, linear, T-shaped and tetrahedral pollen tetrads. The pollinia are compact and at anthesis the pollen grains are 2-celled. Ovules are anatropous, bitegmic and tenuinucellate. Both the integuments are dermal in origin and 2-layered. Development of female gametophyte is of the monosporic, 8-nucleate type. Double fertilization occurs. The primary endosperm nucleus degenerates. The mature embryo is organized from the derivatives of tiers 1, 1, m, and n. Its development conforms to a variation of the Onagrad type. The seed coat is formed entirely by the outer layer of the outer integument. There are three sterile and three fertile valves in the ovary. In the prefertilization stages valves consist of parenchymatous cells. After fertilization, the sterile valves become sclerenchymatous whereas the fertile valves remain parenchymatous. The embryological characters support the disputed systematic position ofLiparis within subtribeLiparidinae ofEpidendreae.     

Anther, pollen and tapetum development in safflower, Carthamus tinctorius L

In safflower, the anther wall at maturity consists of a single epidermis, an endothecium, a middle layer and the tapetum. The tapetum consists mainly of a single layer of cells. However, this single-layer appearance is punctuated by loci having ‘two-celled’ groupings due to additional periclinal divisions in some tapetal cells. Meiotic division in microsporocytes gives rise to tetrads of microspores. The primexine is formed around the protoplasts of microspores while they are still enveloped within the callose wall. Just prior to microgametogenesis, the microspores enlarge through the process of vacuolation, and the exine wall pattern becomes established. Microgametogenesis results in the formation of 3-celled pollen grains. The two elongated sperm cells appear to be connected. The exine wall is highly sculptured with a distinct tectum, columellae, a foot layer, an endexine and a thin intine. Similar to other members of the Asteraceae family, the tapetum is of the invasive type. The most novel finding of this study is that in addition to the presence of invasive tapetal cells, a small population of ‘non-invasive’ tapetal cells is also present. The tapetal cells next to the anther locules in direct contact with the microspores become invasive and start to grow into the space between developing microspores. These tapetal cells synthesize tryphine and eventually degenerate at the time of gametogenesis releasing their content into the anther locules. A smaller population of non-invasive tapetal cells is formed as a result of periclinal divisions at the time of tapetum differentiation. These cells are not exposed to the anther locules until the degeneration of the invasive tapetal cells. The non-invasive tapetal cells have a different cell fate as they synthesize pollenkitt. This material is responsible for allowing some pollen grains to adhere to each other and to the anther wall after anther dehiscence. This observation explains the out-crossing ability of Carthamus species and varieties in nature.     

EVOLUTIONARY AND ECOLOGICAL SIGNIFICANCE OF STARCH STORAGE IN POLLEN OF THE ARACEAE

Starch content was qualitatively assessed for pollen of 79 of the 111 currently recognized genera of the family Araceae—one of three monocot families known to exhibit both starchy and starchless pollen. Although 73% of the genera investigated had exclusively starchy pollen, character correlation suggests that starchless pollen is the primitive type for the family Araceae, as well as for monocots in general. Pollen starch content is a highly conservative character at the generic level in Araceae; only a single genus (Schismatoglottis) clearly exhibits both character states. The distribution of starchy pollen among aroid genera is consistent with what have here been termed Bakers' Starch Laws. Aroid pollen below a certain critical diameter—17-25 μm—is almost invariably starchless. Larger pollen is nearly always starchy, except where insect pollinators may use pollen nutritionally. There is strong evidence that the trend from starchless to starchy pollen in Araceae is reversible, according to the constraints imposed by the aforementioned factors.     

REPRODUCTIVE FEATURES OF DENTARIA LACINIATA AND D. DIPHYLLA (CRUCIFERAE), AND THE IMPLICATIONS IN THE TAXONOMY OF THE EASTERN NORTH AMERICAN DENTARIA COMPLEX

Reproductive features including ovule development, megasporogenesis, megagametogenesis, microsporogenesis, microgametogenesis, pollen tube growth, embryogeny, and natural seed germination were studied in a single population each of Dentaria laciniata Muhl. ex. Willd. and D. diphylla Michx. to test for possible agamospermy. The population of D. laciniata studied is sexual. The archesporial cell functions directly as the megasporocyte. It undergoes two meiotic divisions, but the micropylar cell of the dyad fails to undergo meiosis II, and a linear triplet of three cells is formed. The chalazal megaspore divides to form an eight-nucleate, seven-celled megagametophyte of the Polygonum type. Simultaneous cytokinesis follows the second meiotic division of the microsporocyte yielding a tetrahedral tetrad of microspores. A three-celled pollen grain is formed prior to anther dehiscence. Following apparent fertilization, the Capsella-variation of the Onagrad type of embryogeny results in a conduplicate embryo. Endosperm is initially nuclear, but eventually becomes cellular. Seeds readily germinate in nature. Similar events are documented in one population of D. diphylla up to the organization of the embryo-sac, which disintegrates before cellularization. These reproductive events and other data indicate that the eastern North American species of Dentaria may form a sexual polyploid complex with some sexual populations and some sterile ones.     

Light and electron microscopies reveal unknown details of the pollen grain structure and physiology from Brazilian Cerrado species

Pollen grains have a relatively simple structure and microscopic size, with two or three cells surrounded by the protective sporoderm at maturity. The viability and efficiency of pollen transport from anther to stigma depends on pollen physiological properties, especially the relative water content of the vegetative cell. Pollen transport is a crucial fate for most angiosperms that depends on biotic pollinators and studies focusing on understanding the morpho-physiological properties of pollen grains are still scarce, especially to tropical open physiognomies as the Brazilian Cerrado. Therefore, we investigate some structural and physiological aspects of pollen grains from six native species naturally growing in one Cerrado area: Campomanesia pubescens (Myrtaceae), Caryocar brasiliense (Caryocaraceae), Erythroxylum campestre (Erythroxylaceae), Lippia lupulina (Verbenaceae), Pyrostegia venusta (Bignoniaceae), and Xylopia aromatica (Annonaceae). We selected dehiscent anthers and mature pollen grains to analyze (1) the anther wall and pollen microstructure, (2) the pollen water status at the time of anther dehiscence, and (3) the pollen chemical compounds. In all analyzed species, the anther and pollen developed in a successfully way, and except for Caryocar brasiliense, all species were able to emit pollen tubes in the germination tests. As expected for a dry and open environment, most species dispersed their pollen grains in a partially dehydrated form, as indicated by our harmomegathy experiment. As indicated by our study, the pollen ability in preventing dissection, maintaining its viability in a dry and hot environment during its transport from anther to stigma, may be related to the sporoderm apertures and to the reserve compounds, mainly carbohydrates in the form of hydrolysable starch grains.

     

Floral anatomy,embryology, seed,and fruit development in Cephalanthus (Naucleeae-Rubiaceae), with emphasis on C. glabratus

Information on the reproductive anatomy in genera of the tribe Naucleeae, particularly Cephalanthus, is scarce and fragmented. Of the six species in the genus, only the mature megagamethophyte of Cephalanthus occidentalis has been described. This study aims to provide information on embryological aspects in flowers of C. glabratus and to analyze the morphology and anatomy of the flowers, fruit, and seed in the six species of the genus. Cephalanthus glabratus have imperfect flowers: pistillate (PF) and staminate (SF). In the PF, the ovules are functional, while in the SF, they atrophy during the formation of the embryo sac. The mature ovule has a single integument, corresponds to the Phyllis type and the embryo sac is a Polygonum type, forming only in the PF. The presence of pollenkitt and secondary presentation of pollen were observed in the SF, as well as in the pollen formation previously described, whereas in the PF, they are absent, due to the collapse of the pollen grains inside the indehiscent anthers. The analysis of the ontogeny of the ovular excrescence in C. glabratus determined its funicular origin, calling it an aril. Its development is a pre-anthesis event, initiated during megasporogenesis. In seeds, the aril is a fleshy, white appendage which almost completely envelops the seeds of Cephalanthus, except for Cephalanthus natalensis where it is noticeably more reduced. Studies of the fruit in Cephalanthus species indicate that the infructescence is a dry schizocarp which separates into uni-seminated mericarps, except in C. natalensis that has fleshy indehiscent fruit.

     

山麦冬大小孢子发生及雌雄配子体发育研究

采用石蜡切片技术对百合科植物山麦冬大小孢子发生及雌雄配子体发育进行了观察研究。结果表明:(1)山麦冬花药具有4个花粉囊,花药壁的发育方式为基本型,花药壁完全分化时由表皮、药室内壁、中层及绒毡层组成。(2)绒毡层发育类型为分泌型,到四分体孢子彼此分离形成单细胞花粉阶段,绒毡层细胞开始解体退化,花粉成熟时绒毡层细胞完全消失;花粉母细胞减数分裂为连续型,四分体为左右对称形排列,成熟花粉为3-细胞花粉,单萌发沟。(3)子房3室,每室2枚胚珠,胚珠倒生型,双珠被,薄珠心,雌性孢原细胞不经过平周分裂而直接发育而成大孢子母细胞。(4)减数分裂后四分体大孢子呈线型或T型排列,合点端大孢子分化为功能大孢子,胚囊发育为蓼型;花粉母细胞减数分裂过程中,二分体、四分体细胞外方被胼胝质壁所包被,小孢子形成后胼胝质壁逐渐消失。该研究结果丰富了百合科植物生殖生物学研究的内容,也为探讨百合科植物的系统学研究提供了参考。     

ABNORMAL POLLEN VACUOLATION1 (APV1) is required for male fertility by contributing to anther cuticle and pollen exine formation in maize

Anther cuticle and pollen exine are the major protective barriers against various stresses. The proper functioning of genes expressed in the tapetum is vital for the development of pollen exine and anther cuticle. In this study, we report a tapetum‐specific gene, Abnormal Pollen Vacuolation1 (APV1), in maize that affects anther cuticle and pollen exine formation. The apv1 mutant was completely male sterile. Its microspores were swollen, less vacuolated, with a flat and empty anther locule. In the mutant, the anther epidermal surface was smooth, shiny, and plate‐shaped compared with the three‐dimensional crowded ridges and randomly formed wax crystals on the epidermal surface of the wild‐type. The wild‐type mature pollen had elaborate exine patterning, whereas the apv1 pollen surface was smooth. Only a few unevenly distributed Ubisch bodies were formed on the apv1 mutant, leading to a more apparent inner surface. A significant reduction in the cutin monomers was observed in the mutant. APV1 encodes a member of the P450 subfamily, CYP703A2‐Zm, which contains 530 amino acids. APV1 appeared to be widely expressed in the tapetum at the vacuolation stage, and its protein signal co‐localized with the endoplasmic reticulum (ER) signal. RNA‐Seq data revealed that most of the genes in the fatty acid metabolism pathway were differentially expressed in the apv1 mutant. Altogether, we suggest that APV1 functions in the fatty acid hydroxylation pathway which is involved in forming sporopollenin precursors and cutin monomers that are essential for the development of pollen exine and anther cuticle in maize.