Paleozoic seed fern pollen organs

Paleozoic pollen organs exhibit numerous morphological forms that have been arranged in categories based on their probable organization. Progymnosperm ancestors are characterized by three dimensional branching systems bearing pairs of terminal sporangia. Early Mississippian examples of seed fern fertile branches appear little modified from the progymnosperms. These pteridosperm microsporangia are nonsynangiate and thin walled with longitudinal dehiscence. By Upper Mississippian time all forms show sporangial clustering into large or small groups, with several taxa exhibiting radially symmetrical synangia. In the Lower Pennsylvanian all pollen organs are synangiate and appear to consist of a uniseriate ring of sporangia that either surround a central hollow, or are bilaterally flattened. Sporangial dehiscence in all forms is longitudinal and toward the center of the synangium. In bilateral synangia with no central hollow, the sporangia either separate laterally or effective dehiscence areas are restricted to the free apical portions of the sporangia. Callistophytacean synangia resemble the lyginopterid type, but are abaxial on laminar foliage. This family is thought to have evolved from the lyginopterids during the Early Pennsylvanian. Middle Pennsylvanian medullosan pollen organs are all radial and may be solitary, aggregated into groups, or fused into a large compound synangium. Several pollen organ types are reinterpreted, and the possible evolutionary relationships among the various Paleozoic pollen organ forms are discussed based on synangial organization, patterns of frond branching, and pollen or prepollen morphology.     

ACAULANGIUM GEN. N., A FERTILE MARATTIALEAN FROM THE UPPER PENNSYLVANIAN OF ILLINOIS

Material described by Graham as Cyathotrachus bulbaceus is believed to represent a new genus that is a common constituent of Upper Pennsylvanian coal balls. The sessile synangia of Acaulangium gen. n. are borne in a row on either side of the pinnule midrib and are composed of four to six short, tapering, laterally appressed sporangia. The sporangia have extended tips which curve over the inside of the synangium distally and delimit a small open area inside the synangium. The outer facing walls of the sporangia are two to three cells thick throughout while the inner facing walls are uniseriate. During dehiscence the sporangia separate laterally and spore release results from the rupture of a row of elongate cells along the inner sporangium midline. Among species of Scolecopteris the new genus resembles S. illinoensis and S. minor var. parvifolia but differs in its sessile synangial attachment. The additional parenchyma present between sporangial cavities in the synangia of Acaulangium, and the tendency toward bilateral symmetry suggests an early stage in the evolution of a bivalve synangium such as is present in Marattia.     

FERAXOTHECA GEN. N., A LYGINOPTERID POLLEN ORGAN FROM THE PENNSYLVANIAN OF NORTH AMERICA

The pollen organ Feraxotheca gen. n. is described from Pennsylvanian age coal balls from the Lewis Creek, Kentucky, locality. The fructifications consist of bilaterally symmetrical synangia composed of a basal pad supporting elongate sporangia that are laterally appressed for the entire length of the sporangial cavities. Sporangial tips extend over the center of the synangium and delimit a small open area, while the bases arise from a parenchymatous cushion that is bounded by short tracheid-like cells. Each synangium is borne on the surface of an expanded pinna tip and is surrounded by a small amount of laminar tissue that envelopes the base of the synangium. Ultimate pinnae are rectangular in transverse section, possess an elliptical vascular bundle surrounded by canals containing a yellow froth-like substance, and have a cortex of elongate cells that radiate from the center of the axis. Sporangia contain small (40–64 μm), radial, trilete spores ornamented by regularly spaced coni or blunt tipped grana. Feraxotheca is compared with the compression genus Crossotheca and some new ideas are advanced concerning the morphology of this compression genus. The obvious differences between Feraxotheca and other lyginopterid pollen organs strongly suggests that the Lyginopteridaceae, as it is currently interpreted, is an unnatural family.     

DEVELOPMENTAL STUDY OF BRANCHED RHIZOPHORES IN THREE SELAGINELLA SPECIES

A morphogenetic investigation was made of the rhizophore of three large-sized tropical Selaginella species. The rhizophores of Selaginella delicatula, S. caudata, and S. plana arise exogenously at the points of branching of the main stems. In S. delicatula they are initiated at the junction of the second youngest branching. The rhizophore apical meristem has a tetrahedral apical cell and is capless. The rhizophores are usually three or four times dichotomously branched in S. delicatula and S. plana and four or five times in S. caudata. In S. delicatula, dichotomous branching of the rhizophore involves formation of two new apical cells subsequent to loss of an original apical cell. A pair of roots is formed endogenously from inner cells below the dermal layer at the apex of ultimate rhizophore branches. The finding that the rhizophore is an autonomously branched, leafless, and capless axis leads us to argue that Selaginella rhizophores, like lepidodendrid rhizomorphs, are fundamental axial organs that coordinate with the stem and root.     

Anatomical inferences on aerial bud protection of three Eugenia shrub species from the Cerrado

• Location and degree of protection of aerial buds are important functional traits in disturbance- or stress-prone environments since aerial buds ensure the development of new organs under favourable growing conditions. This study was carried out in a Brazilian Cerrado area under regeneration after long-term Pinus cultivation, where the trees were clear-cut in 2012 and the remaining material was burned in 2014.
• After the fire treatment, several species resprouted from belowground organs and their aboveground organs were directly exposed to full sunlight. We collected 15 terminal branches with fully expanded leaves from three individuals of each of three Eugenia species to investigate if those with well-developed belowground organs invest in bark for aboveground bud protection. The samples were analysed using light and electron microscopy.
• In addition to terminal and axillary buds, all species presented accessory buds, and the number varied according to the node analysed. None of the aerial buds were protected by bark, but all were well protected by cataphylls and densely pubescent leaf primordia. There were also inter- and intra-petiolar colleters that released a mucilaginous protein exudate. The distance between the shoot apical meristem and the outer surface was longer in the terminal bud than in axillary buds. The bud leaf primordia covering the shoot apical meristem had a thick cuticle, unicellular non-glandular trichomes that accumulate phenolic and lipophilic compounds, and secretory cavities.
• Our study shows that all three Eugenia species studied here had highly protected aerial buds allocated from belowground organs. These morphological traits may improve the chances of the species' persistence in areas subjected to frost events, low relative humidity, high irradiance and harmful UV levels.

     

THE ORIGIN AND DEVELOPMENT OF INDUCED BRANCHES IN HELIANTHUS

The development of buds and their vascular connections are described for Helianthus annuus and H. bolanderi. Bud meristems of H. annuus usually become isolated by parenchymatization of the bud traces in the cortical zone. If the buds are induced to grow as a result of decapitation of the terminal meristem, a continuous range between typical primary connections and pseudo-adventitious connections are made between the main axis and the lateral buds. Considerable growth of the branches occurs within 48 hours after decapitation. Axillary buds of H. bolanderi grow continuously, and both the meristem and vascular system of the buds are derived directly from the apical meristem of the shoots.     

THE SPORANGIUM OF HORNEOPHYTON LIGNIERI (RHYNIOPHYTINA)

Numerous sporangia of Horneophyton lignieri from the Rhynie Chert locality in Scotland have been studied. The sporangia are branched, with two to four columellate lobes of varying length, and a continuous sporogenous zone or cavity occurs among the lobes. Unbranched sporangia, generally thought to be the typical form for the plant have not been found, and their presence is not established. Although not definitely proven, evidence suggests that the sporangia opened by means of a small apical pore or stoma. An area of thick-walled cells at the apex of each sporangial lobe probably played some role in this opening. Radial, trilete, azonate spores ranging from 39–49 μm in diam, with curvaturae perfectae are produced most commonly in tetrahedral tetrads and occasionally in isobilateral tetrads. Matters of spore preservation and possible ornamentation are discussed. The branched sporangia of this genus are unique among bryophytes and vascular plants and provide some evidence that certain synangia may have arisen from a single sporangium rather than from multiple sporangia borne singly at the tips of ultimate branches.     

A marattialean fern,Scolecopteris libera n. sp., from the Asselian (Permian) of Inner Mongolia,China

Scolecopteris libera n. sp. is established on partial three-dimensionally preserved materials containing both sterile and fertile fronds collected from the volcanic tuff at the top of the Taiyuan Formation at the Wuda Coalfield, Inner Mongolia, China. It is characterized by tripinnate fronds with rachises over 200 mm in diameter; and long lanceolate or falciform pinnules with thick veins. Pinnules are of the pecopterid-type, with those at the base of the ultimate pinnae usually divided into small lobes. Eight to twelve circular synangia are arranged in two rows along the midvein. Synangia are borne on a short pedicel attached to the middle of lateral veins. A synangium is composed of 7–10 fusiform exannulate sporangia with pointed apices. The sporangia are free among each other beyond the base of synangia. In situ microspores of the Cyclogranisporites leopoldii type differ from all other in situ Paleozoic marattialean spores.     

Cell division patterns in the apices of subterranean axis and aerial shoot of Psilotum nudum (Psilotaceae): morphological and phylogenetic implications for the subterranean axis

The cell division pattern in the apical meristem of Psilotum nudum was examined using epi-illumination microscopy and a paraffin method. In the subterranean axis, about half of the derivative cells of the apical cell produce tetrahedral daughter apical cells by the first three or more oblique divisions. Roughly half of these apical cells give rise to the apical meristems of axes, whereas the other half do not. Various relative activities of the mother and daughter apical cells give rise to disordered branching patterns. In the ill-organized apical meristem as well as the leafless and capless structure, the Psilotum subterranean axis differs from the basic organs of vascular plants such as stem and root and seems to be an independent organ. The cell division pattern characteristic of the subterranean axis persists in the young unbranched aerial shoots, although fewer daughter apical cells are produced. Dichotomous branching of the aerial shoots, as in a variety of organs of pteridophytes, involves loss of the mother apical cell followed by appearance of two daughter apical cells.     

Morphogenesis of the reproductive shoots of Welwitschia mirabilis and Ephedra distachya (Gnetales), and its evolutionary implications

For decades, Gnetales appeared to be closely related to angiosperms, the two groups together forming the anthophyte clade. At present, molecular studies negate such a relationship and give strong support for a systematic position of Gnetales within or near conifers. However, previous interpretations of the male sporangiophores of Gnetales as pinnate with terminal synangia conflict with a close relationship between Gnetales and conifers. Therefore, we investigated the morphogenesis of the male reproductive structures of Welwitschia mirabilis and Ephedra distachya by SEM and light microscopy. The occurrence of reduced apices to both halves of the antherophores of W. mirabilis gives strong support for the assumption that the male ‘flowers’ of W. mirabilis represent reduced compound cones. We assume that each half of the antherophore represents a lateral male cone that has lost its subtending bract. Although both halves of the antherophores of Ephedra distachya lack apical meristems, the histological pattern of the developing antherophores supports interpreting them as reduced lateral male cones as well. Therefore, the male sporangiophores of Gnetales represent simple organs with terminal synangia. Although extant conifers do not exhibit terminal synangia, similar sporangiophores are reported for some Cordaitales, the hypothetical sister group of conifers. Moreover, several Paleozoic conifers exhibit male cones with terminal sporangia or synangia. Therefore, we propose that conifers, Cordaitales and Gnetales originated from a common ancestor that displayed simple sporangiophores with a terminal cluster of sporangia.     

Floral development in bolting garlic

Garlic (Allium sativum L.) is a completely sterile plant, propagated only vegetatively. The aim of this research was to study the sequence of morphological processes occurring during floral initiation and development of a number of bolting garlic accessions from the Allium gene bank in Israel by using SEM. The garlic inflorescence is an umbel-like flower arrangement, the branches (flower clusters) of which arise from a common meristem. The numerous flowers have a distinct morphology typical of the genus Allium. Flower-stalk elongation precedes the swelling of the apical meristem and its subdivision into several centers of floral development. Within clusters, floral primordia develop unevenly. Differentiation of topsets begins after floral differentiation on the peripheral part of the apical surface, and their size, number and rate of development vary among genotypes. At least four morphological types differing in flower/topset ratio were distinguished among the 12 clones studied in this investigation. For further studies of flowering physiology and fertility restoration, only clones which can differentiate the greatest proportion of normal flowers and the least of topsets in the apical meristem should be selected. Received: 28 June 2000 / Revision accepted: 6 November 2000     

Establishment of polarity in angiosperm lateral organs

In seed plants, lateral organs such as leaves and floral organs are formed from the flanks of apical meristems. Therefore, they have an inherent positional relationship: organ primordia have an adaxial side next to the meristem, and an abaxial one away from the meristem. Surgical and genetic studies suggest that a morphogenetic gradient, which originates in the meristem, converts the inherent polarity into a functional one. Once an adaxial-abaxial axis of polarity is established within organ primordia, it provides cues for proper lamina growth and asymmetrical development. Several key participants in this process have been identified, and analyses of these genes support and refine our views of axis formation in plants. The complex relationships between and within various members of these plant-specific gene families (class III HD-ZIPs, YABBYs and KANADIs) might account for a substantial part of the morphological variation in lateral organs of seed plants.     

Herbivory could unlock mutations sequestered in stratified shoot apices of genetic mosaics

Many higher plants have shoot apical meristems that possess discrete cell layers, only one of which normally gives rise to gametes following the transition from vegetative meristem to floral meristem. Consequently, when mutations occur in the meristems of sexually reproducing plants, they may or may not have an evolutionary impact, depending on the apical layer in which they reside. In order to determine whether developmentally sequestered mutations could be released by herbivory (i.e., meristem destruction), a characterized genetic mosaic was subjected to simulated herbivory. Many plants develop two shoot meristems in the leaf axils of some nodes, here referred to as the primary and secondary axillary meristems. Destruction of the terminal and primary axillary meristems led to the outgrowth of secondary axillary meristems. Seed derived from secondary axillary meristems was not always descended from the second apical cell layer of the terminal shoot meristem as is expected for terminal and primary shoot meristems. Vegetative and reproductive analysis indicated that secondary meristems did not maintain the same order of cell layers present in the terminal shoot meristem. In secondary meristems reproductively sequestered cell layers possessing mutant cells can be repositioned into gamete-forming cell layers, thereby adding mutant genes into the gene pool. Herbivores feeding on shoot tips may influence plant evolution by causing the outgrowth of secondary axillary meristems.     

FLORAL STRUCTURE AND ORGANOGENESIS IN PODOPHYLLUM PELTATUM (BERBERIDACEAE)

The life cycle of Podophyllum can be divided into two phases, a subterranean phase during which a conspicuous winter mixed terminal bud forms at the end of a rhizome, and an aerial phase, during which the primordia of the structures within the winter bud give rise the next spring to an aerial shoot composed of a stem, 2 leaves, and a single flower. The transition from a vegetative to a floral apex occurs at the end of July, when the apical meristem becomes a globoid structure. During the first and second weeks of August, the floral organs are laid down along the sides of an elongated floral apex. The order of initiation of the floral organs is sepals, petals, stamens, gynoecium, and stamens. Petal primordia are initiated in early August, but growth ceases after they attain a height of about 2 mm. This inhibition persists until the middle of May in the next growing season, when the petals grow to 12 mm within 2 weeks. At anthesis the petals have enlarged to a length of 2 cm or more. The gynoecium is usually composed of a single terminal carpel. The ovules are chiefly supplied by branches from a ventral bundle complex, but that is supplemented by medullary bundles that are formed in the base of the gynoecium, below the loculus. It could be argued that these medullary bundles are surviving remnants of the vascular supply to a second carpel, no longer extant. A transmitting tract extends from the stigma about half the distance to the loculus. The tract is lined with unicellular glandular cells and is open from the stigma to the loculus.     

STUDIES OF PALEOZOIC MARATTIALEANS: AN EARLY PENNSYLVANIAN SPECIES OF THE FERTILE FERN SCOLECOPTERIS

Remains of the fossil Marattiales are very rare in Lower Pennsylvanian sediments. The present report describes a new species of the fertile fern foliage Scolecopteris from the Lewis Creek, Kentucky locality (Lower or lower Middle Pennsylvanian). Scolecopteris conicaulis n. sp. has radial synangia composed of a ring of 4–7 elongate, exannulate sporangia. Most features of the synangia of S. conicaulis were previously hypothesized to be primitive in Scolecopteris based on geologically younger species. Supposed primitive characters include the large synangium pedicel with fiber core, an outer-facing sporangial wall lacking differentiation or zonation, and large spores. The anatomy of the sporangium walls, pinnule morphology, and general spore type support an association with the Minor group of Scolecopteris. The new species is similar in several important features to Scolecopteris (Cyathotrachus) altus, the only other anatomically preserved fertile marattialean known from this early time, and indicates a considerably earlier origin for fertile foliage of this type.     

DETERMINATE (det) MUTANT OF PISUM SATIVUM (LEGUMINOSAE: PAPILIONOIDEAE) EXHIBITS AN INDETERMINATE GROWTH PATTERN

Inflorescence ontogeny and morphology of the det mutant of Pisum sativum L. were investigated using scanning electron microscopy. This mutation causes the production of a limited number of axillary flowers followed by the formation of an apparent terminal flower slightly offset from the vertical. Our study indicates that the apparent terminal flower arises from an axillary meristem. The terminal meristem senesces and differentiates hairs, forming a rudimentary stub in the same manner as axillary meristems of conventional (Det) and det plants. Thus the dramatic effect of the det gene on inflorescence architecture results from early apical arrest rather than conversion of the terminal meristem to a flower as implied by the symbol det. This mutant will be valuable in elucidating regulation of apical arrest.     

ONTOGENY OF THE INFLORESCENCE OF SAURURUS CERNUUS (SAURURACEAE)

The spicate inflorescence of Saururus cernuus L. (Saururaceae) results from the activity of an inflorescence apical meristem which produces 200–300 primordia in acropetal succession. The inflorescence apex arises by conversion of the terminal vegetative apex. During transition the apical meristem increases greatly in height and width and changes its cellular configuration from one of tunica-corpus to one of mantle (with two tunica layers) and core. Primordia are initiated by periclinal divisions in the subsurface layer. These are “common” primordia, each of which subsequently divides to produce a floral apex above and a bract primordium below. The bract later elongates so that the flower appears borne on the bract. All common primordia are formed by the time the inflorescence is about 4.4 mm long; the apical meristem ceases activity at this stage. As cessation approaches, cell divisions become rare in the apical meristem, and height and width of the meristem above the primordia diminish, as primordia continue to be initiated on the flanks. Cell differentiation proceeds acropetally into the apical meristem and reaches the summital tunica layers last of all. Solitary bracts are initiated just before apical cessation, but no imperfect or ebracteate flowers are produced in Saururus. The final event of meristem activity is hair formation by individual cells of the tunica at the summit, a feature not previously reported for apical meristems.     

Aulacotheca collicola n.sp. (medullosaceae) from the early Pennsylvanian of the Illinois basin

Compressed synangia referable to the medullosan pollen organ genus Aulacotheca Halle have been discovered in Early Pennsylvanian sediments from Rock Island County, Illinois, U.S.A. Specimens were recovered from localized shales containing floral elements suggesting an upland environment. Based on synangial size, morphology, sporangial number, and prepollen type, two distinct forms, A. collicola n.sp. and Aulacotheca sp., are recognized. Synangia of A. collicola are 11–17 × 2.5 – 4.0 mm and have four to six sporangia. Free sporangial tips are acutely pointed and dehiscence is through a longitudinal slit to the inside of the synangium. Prepollen averages 181 × 129 μm and has a monolete suture with median deflection. Sexine on proximal and distal surfaces shows narrow, closely spaced, anastomosing inpockets; distal and lateral walls are separated by a deep, encircling fold. Synangia of Aulacotheca sp. are 21–33 × 3.8 – 5.5 mm and have 4–6 sporangia. Distally, free sporangial tips have a narrow extension, giving a mucronate appearance. Features of these species suggest that greater morphological diversity in synangial and pollen structure occurs in Aulacotheca and the medullosan pteridosperms than previously recognized.     

FASCIATION AND DICHOTOMOUS BRANCHING IN ECHINOCEREUS (CACTACEAE)

In Echinocereus reichenbachii dichotomous branching and fasciation (cresting) are rare events. Both were found together in only a few of many populations investigated and are interpreted as variants of a single phenomenon. They may occur at any stage of shoot development, but crest meristems arise most commonly on young branches among clusters of normal shoots. Sometimes they appear on unbranched young plants or seedlings, very rarely on older shoots. Dichotomy results from the division of an apical meristem into equal parts each of which functions independently, producing a forked shoot. Fasciation involves the extension of a single meristem into an apical ridge. The product is a flabellate shoot that becomes undulate if growth along the summit continues. In longisection linear meristems appear similar to radial sections of normal shoots; in median sagittal section they have a much extended central mother cell zone within which the cell pattern resembles a rib meristem. Although crest meristems become sluggish or even inactive with age, localized renewed growth may occur spontaneously or be induced by injury. In this species the random production of normal shoots from crest meristems (defasciation) was not observed, but if much or all of such a meristem is removed, branches may arise from lateral areoles, and these are always normal. It seems, therefore, that whatever induces fasciation in E. reichenbachii originates in and is restricted to the apical meristem and its immediate vicinity.     

Isolation and characterization of a rice homebox gene, OSH15

In many eukaryotic organisms including plants, homeobox genes are thought to be master regulators that establish the cellular or regional identities and specify the fundamental body plan. We isolated and characterized a cDNA designated OSH15 (Oryza sativa homeobox 15) that encodes a KNOTTED-type homeodomain protein. Transgenic tobacco plants overexpressing the OSH15 cDNA showed a dramatically altered morphological phenotype caused by disturbance of specific aspects of tobacco development, thereby indicating the involvement of OSH15 in plant development. We analyzed the in situ mRNA localization of OSH15 through the whole plant life cycle, comparing the expression pattern with that of another rice homeobox gene, OSH1. In early embryogenesis, both genes were expressed as the same pattern at a region where the shoot apical meristem would develop later. In late embryogenesis, the expression pattern of the two genes became different. Whereas the expression of OSH1 continued within the shoot apical meristem, OSH15 expression within the shoot apical meristem ceased but became observable in a ring shaped pattern at the boundaries of some embryonic organs. This pattern of expression was similar to that observed around vegetative or reproductive shoots, or the floral meristem in mature plants. RNA in situ localization data suggest that OSH15 may play roles in the shoot organization during early embryogenesis and thereafter, OSH15 may be involved in morphogenetic events around the shoot apical meristem.