MITROSPERMUM VINCULUM SP. NOV., A CARDIOCARPALEAN OVULE FROM THE UPPER PENNSYLVANIAN OF OHIO

Numerous anatomically preserved ovules assignable to the genus Mitrospermum have been discovered in Upper Pennsylvanian sediments of Eastern Ohio. Although basically similar to Mitrospermum compressum, the newly discovered specimens exhibit several consistent differences. Ovules are strongly platyspermic, up to 4.2 mm long, 4.0 mm wide, and 0.6 mm thick. In the minor plane, ovules are broadest at the base and taper toward the micropyle. The integument exhibits three topographic regions: endotesta, sclerotesta, and sarcotesta. The sarcotesta is extremely broad in the major plane, where it forms two membranous wings. A single terete vascular bundle enters the base of the ovule, traverses the integument, and divides to form two integumentary bundles and a conspicuous nucellar platform. Integumentary bundles extend toward the tip of the ovule at the margin of the sarcotesta and sclerotesta. A pollen chamber with a prominent nucellar beak is delimited at the tip of the nucellus. Consistent differences in vascularization, size, nature of the seed base, features of the pollen chamber, and the Late Pennsylvanian age demonstrate that the specimens represent a distinct species. The discovery of these ovules extends the stratigraphic range of Mitrospermum to the Upper Pennsylvanian of Ohio.     

ONTOGENY OF THE PALEOZOIC OVULE,CALLOSPERMARION PUSILLUM

The ontogeny of the upper Pennsylvanian age gymnospermous ovule, Callospermarion pusillum, is described from petrifaction specimens collected at the Berryville locality in Illinois. Ovules exhibit a wide range of dimensional and structural features that indicate an extensive developmental sequence. Specimens range from ovules with indistinct zonation of the thin-walled integument to those with thick-walled cells of the sclerotesta. The apex of the fleshy nucellus in some specimens is preserved as a cellular mound, while in others a well-formed cellular pollen chamber is present; still other ovules are characterized by a papery-thin nucellus and pollen chamber wall. The megagametophyte of most specimens is represented by a hollow megaspore membrane that may be restricted to the base of the nucellus, or occupy the entire seed cavity. In a few specimens cellular gametophytes are preserved, and in one ovule archegonia with supposed eggs are also present. Variability in each of the features is compared with ontogenetic changes in comparable structures of living gymnospermous ovules and is correlated with ovule size. A developmental sequence for the fossil ovules is proposed.     

OVULE ABORTION IN ‘NONPAREIL’ ALMOND (PRUNUS DULCIS [MILL.] D. A. WEBB)

The earliest indication of ovule abortion in almond (Prunus dulcis [Mill.] D. A. Webb ‘Nonpareil‘) is the deposition of callose (as indicated by aniline blue fluorescence) 2 days after pollination which is 2 days before clear histological symptoms of ovule degeneration are evident and 6 days before fertilization of the viable ovule. Callose deposition begins in the chalazal region of the nucellus where the funicular trace enters the ovule and ramifies into the integuments. As ovule abortion progresses, callose deposition in the inner integument extends as a ring around the nucellus. Movement of the fluorescent dye disodium fluorescein (uranin) indicated that translocation from the vascular trace into abortive ovules becomes blocked at the chalazal position. The dye freely penetrates and diffuses into viable ovules but fails to penetrate abortive ovules. Lack of, or delayed and irregular, megagametophyte development was another characteristic of abortive ovules. Biochemical and histochemical analyses of abortive and viable ovules indicated that carbohydrate depletion parallels ovule abortion. These observations lead to the conclusion that ovule abortion is accompanied by blockage in metabolite supply although whether this blockage is the primary cause or a consequence of ovule abortion is uncertain.     

Resolving the systematic and phylogenetic position of isolated ovules: a case study on a new genus from the Permian of China

Isolated ovules occur in many fossil plant assemblages, where they provide important insights into seed‐plant diversity and evolution. However, in many cases, the ovules cannot be attributed to individual groups of seed plants, restricting systematic and evolutionary assessments that can be made from otherwise well‐characterized fossil taxa. In the present paper, we describe a new kind of ovule discovered in tuffaceous sediments from the Permian‐aged Xuanwei Formation in Guizhou Province, China. This ovule has 180° rotational symmetry and an integument comprising a variably thick sarcotesta, a uniformly thick sclerotesta and a uniformly thin endotesta. The nucellus is attached to the integument at least basally and contains a collapsed seed megaspore; a nucellar apex is absent. Both the integument and nucellus are vascularized by paired bundles in the major plane of the ovule; the integumentary bundles are considerably larger than the nucellar bundles and the nucellar bundles emerge from a conical vascular pad. Generation of a three‐dimensional reconstruction based on serial peels revealed the gross morphology and organization of the ovule and highlighted the presence of features consistent with cardiocarpalean‐type ovules (ovule shape, histological features of the integument) and also features more typical of lagenostomalean‐ and trigonocarpalean‐type ovules (large integumentary bundles, presence of nucellar bundles). To assess the affinity and evolutionary significance of the ovule, it has been included in a cladistic matrix of cardiocarpalean‐, lagenostomalean‐ and trigonocarpalean‐type ovules. Results place the ovule within the cardiocarpalean group of ovules known to have been produced by several plant groups, including cordaitean coniferophytes, pteridosperms and Palaeozoic conifers. The cladistic topology supports generic level distinction of the present species, requiring the establishment of Muricosperma guizhouensis Seyfullah & J.Hilton gen. & sp. nov . Lagenostomalean ovules produced by hydrasperman pteridosperms form a basal paraphyletic grade, whereas trigonocarpalean ovules produced by medullosan pteridosperms form a monophyletic group in which Stephanospermum is paraphyletic with respect to Rhynchosperma and Pachytesta. The results also place the Mississippian ovule Mitrospermum bulbosum apart from all of the Pennsylvanian species of Mitrospermum that form a strongly supported clade. Consequently, M. bulbosum is transferred to the new genus Whitaddera Seyfullah & J.Hilton as W. bulbosa (Long) Seyfullah & J.Hilton. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 84–108.     

The Ovule Structure and Development of Male and Female Gametophytes in Araentotaxus

The ovules of Amentotaxus are 2–2.5 cm in length and about 1.3 cm in width. Aril, which contains about 14—17 secretory canals in transection, is com- posed of parenchyma. The vascular bundles are absent in aril. The integument is com- posed of ceils of outer, middle, and inner regions. The middle region of the integument is about 10 cells in thickness. The Integuments are basally parenchymatous before pollination and then the cells become elongating and thickenning in middle region after pollination. Finally the micropylar canal is forced to be closed. In addition, there are 8–14 vascular bundles with centripetal xylem arranged in a ring in integument. One of the most remarkable feature of the ovule is that there is a pollen chamber formed at the top of nucellus before pollination. At this time because epidermis of nucellus does not disintegrate, the pollen chamber looks like conical in longitudinal section. After pollination the pollen chamber was becoming closed due to elongation and thickenning of cells in the middle region of integument. At the base of ovule there are 4–5 pairs of bracts which contain a larger secretory cavity and a centripetal xylem in a vascular bundle. It is interesting that wax layer of 30–40 μm thick is covered on the surface of integument, aril and bract. Usually 3–4 microsporangia which are hypodermal in origin, occur in abaxial side of a microsporophyll. In some cases the tapetum is partly enlarging and extruding into the developing tetrahedral tetrads. The mature pollen comprises an antheridial initial and a tube cell. About 20 pollen grains may germinate in the same ovule. The megaspore divide successively 8 times to produce 256 nuclei and then cell wall formation takes place. The female gametophyte is about 830–908 μm in length and 500 μm in width. The archegonia are single, terminal, and 6–7 in number. The mature archgonium, with ventral canal nucleus, is about 430 μm in length and 80–108 μm in width. The female gametophyte is often growing against the upper part of the nuceilus and makes the cells of the latter gradually to be disintegrated. The ovule construction of Amentotaxus is in some degree similar to that of Ginkgo in having a comparatively well developed pollen chamber. The mature pollen of Amentotaxus, which is similar to that of Cephalotaxus is composed of 2-cells. In sum, Amentotaxus perhaps is the most primitive genus in Taxaceae and it is closely related to Cephalotaxus.     

STRUCTURALLY PRESERVED FOSSIL PLANTS FROM ANTARCTICA. IV. TRIASSIC OVULES

Small, anatomically preserved ovules are described from specimens collected at Fremouw Peak in the central Transantarctic Mountains. The ovules occur within a silicified peat in the upper part of the Fremouw Formation, which is considered to be Early to Middle Triassic. Ovules are radially symmetrical and ovoid, with an integument that consists of a narrow endotesta and a complex sclerotesta. The bilayered nucellus has a characteristic scalloped appearance and is attached to the integument only at the base. Of 43 ovules examined, 50 to 60% contain cellular megagametophyte tissue. One specimen contains a possible archegonium with embryo. Although the ovules have not been found attached, their possible affinities are discussed in relation to the known flora from this locality and other, comparable-aged floras from Gondwana.     

Permineralized Cardiocarpalean Ovules in Wetland Vegetation from Early Permian Volcaniclastic Sediments of China

In this paper we report the first occurrence of permineralized plant fossils in volcaniclastic lithologies from China and in doing so describe a new species of cardiocarpalean ovule permineralized within tuffaceous sediments from a recently discovered locality near Pingquan, Hebei Province, North China. The tuff is part of the Early Permian Taiyuan Formation which contains a diverse fossil plant assemblage that includes cordaitean spermatophytes, lepidodendralean lycophytes, equisetophytes, and filicalean ferns, all of which were typical of Permian floras of northern China at this time. Specimens of Cardiocarpusdabiziae sp. nov. have the characteristic platyspermic shape and vascularization of cardiocarpalean ovules, and display prominent protuberances on the exterior of the integument. The integument of the ovule is composed of three layers; a thick sarcotesta, a thin and comparatively dense sclerotesta, and a single layer of large endotesta cells. The pollen chamber produces a slender nucellar beak. In several specimens the megaspore membrane contains well-developed tissue of the megagametophyte. The ovules have prominent external integumentary protuberances which suggest that well-developed systems of plant/animal interactions were operative in Early Permian wetland biotas of the Cathaysian realm.     

ASPECTS OF CONE AND OVULE ONTOGENY IN CRYPTOMERIA (TAXODIACEAE)

The axillary complex of female cones of Cryptomeria is initiated as a tangentially extended triangular structure with a rounded apex. It is bilaterally symmetrical. Structures interpreted as prophylls are differentiated first, but they become insignificant in later development. They are succeeded by two successive pairs of lobes, each lobe being the common primordium for an adaxial ovule and a tooth. The ovule initially much exceeds the tooth. The apex of the complex has a diversity of fates and may differentiate as an ovule-tooth pair. A one-to-one relation between teeth and ovules may be lost by abortion of ovules. The initial relation between teeth and ovules is obscured in later development due to extension of tissues at the base of the complex associated with considerable enlargement of the teeth. Histogenesis of the various parts is described, together with the vascular system. There is a vascular supply to the tooth but not the ovule. The results support a direct comparison with the extinct transition conifers Pseudovoltzia and Aethophyllum but do not fully support Florin's generalized model for the arrangement of parts in the axillary complex of conifers.     

Latisemenia longshania,gen. et sp. nov., a new Late Devonian seed plant from China

The earliest known ovules in the Late Devonian (Famennian) are borne terminally on fertile branches and are typically enclosed in a cupule. Among these ovules are some that have terete integumentary lobes with little or no fusion. Here, we report a new taxon, Latisemenia longshania, from the Famennian of South China, which bears cupulate ovules that are terminal as well as opposite on the fertile axis. Each ovule has four broad integumentary lobes, which are extensively fused to each other and also to the nucellus. The cupule is uniovulate, and the five flattened cupule segments of each terminal ovule are elongate cuneate and shorter than the ovule. Associated but not attached pinnules are laminate and Sphenopteris-like, with an entire or lobate margin. Latisemenia is the earliest known plant with ovules borne on the side of the fertile axis and may foreshadow the diverse ovule arrangements found among younger seed plant lineages that emerge in the Carboniferous. Following the telome theory, Latisemenia demonstrates derived features in both ovules and cupules, and the shape and fusion of integumentary lobes suggest effective pollination and protection to the nucellus. Along with other recent discoveries from China, Latisemenia extends the palaeogeographic range of the earliest seed plants.     

Anatomically preserved Cycadeoidea (Cycadeoidaceae), with a reevaluation of systematic characters for the seed cones of Bennettitales

Four anatomically preserved ovulate cycadeoid cones have been recovered from three localities in Upper Cretaceous (Turonian/Coniacian-Late Campanian) sediments of Vancouver and Hornby Islands, British Columbia, Canada. All of the specimens are preserved by calcareous cellular permineralization and are quite similar to seed cones described as several species of Cycadeoidea and Bennettites. These cones, described as Cycadeoidea maccafferyi sp. nov., consist of tightly packed interseminal scales and ovulate sporophylls with terminal ovules. Two specimens also preserve remains of a small receptacle. Interseminal scales and ovulate sporophylls are oriented parallel to one another. Ovules are distinctly stellate at the base of the micropylar tube, and the sarcotesta consists of both longitudinally oriented tubular cells and large radially elongated cells attached to the sclerotesta. The vascular strand below each ovule is highly contorted in a pattern that is characteristic of contractile tissue in the roots of living plants. These specimens are the most recent anatomically preserved cycadeoid cones yet discovered, revealing details of the reproductive biology shortly before extinction of the clade. Superb preservation of the British Columbia cones confirms that Bennettitales lack a cupule, have radial seeds, and have a vascularized nucellus (but no integumentary tracheids), and that no pollen chamber is produced. Together with a new species of Williamsonia preserved at one of the same localities, these specimens reveal a clear set of contrasting systematic characters for differentiating between isolated seed cones of Williamsoniaceae and Cycadeoidaceae.     

Translocation of Uranin within the Living Ovules of Vanilla

In the ovules of Vanilla (Vanilla planifolia Andr.) before fertilization, outer integument surrounded the lower part of ovule. Uranin got into ovule through funiculus, forming, the first center of fluorescence at the chalaza zone of ovule. Then uranin was transported to micropyle end along inner integument, forming the second center of fluorescence at micropyle end of inner integument. Soon, fluorescence appeared in the egg apparatua. After fertilization, the outer integument ovule extended upward, forming micropyle ogerber with inner integument. After getting into ovule through funiculus, uranin spreads to- ward several directions: l. transported to outer integument at the entrance of micropyle; 2. transported downward to chalaza zone along outer integument at the side of funiculus; 3. extended from chalaza zone to the inside and to the outer integument at the side far from funiculus The ovules of Vanilla had no vascular bundles. On transporting in inner integument, however, the cells in inner layer next to the embryo sac appeared to be the major passage. In mature embryo sac, there was cuticle between inner integument and embryo sac at the half of micropyle end. But between embryo sac at the half of chalaza end and nucellus, cuticle was absent. Nutrient could get into embryo sac from chalaza end undoubtedly. As egg apparatus showed the fluorescence after formation of fluorescence center of inner integument at micropylar end, the possibility that nutrient got into embryo sac from micropyle could not be excluded.     

北美香柏雌球果的发育

用扫描电镜(SEM)观察了北美香柏 Thuja occidentalis 雌球果的发育过程。在北京,北美香柏的雌球果是在八月初由营养芽转变而来,雌球果一般有4~6对苞片,中间2~3对可育,每一苞片腋部着生两枚胚珠,在可育苞片腋部最先观察到一扁平的隆起,并在其上分化出两个胚珠原基,接着分化出珠被和珠心,最后形成扁平而两侧对称的胚珠。在北美香柏雌球果发育过程中,约一半的雌球果在2~3对可育苞片中位于下面的1~2对的腋部产生3个胚珠原基,中间一个较小,并在以后的发育中逐渐退化。由此推测北美香柏的雌球果可能是由祖先类群中每一苞片具多于2个胚珠的雌球果演化而来。在光镜下对雌球果维管系统的观察发现,传粉前幼小雌球果的苞片内仅有一束维管束,传粉后随着苞片基部的居间生长,有4—8束维管束在苞片内形成,但是新发育的维管束木质部和韧皮部相对位置与正常叶性器官一致,这与在以往报道的柏科植物成熟雌球果的苞片中均有反向维管束的发育不同。北美香柏雌球果早期发育和维管束分析结果支持傅德志和杨亲二提出的解释裸子植物生殖器官形态演化的“苞鳞-种鳞复合体”理论。关键词北美香柏;雌球果的发育;胚珠分化;SEM     

CONE AND OVULE ONTOGENY IN TAXODIUM AND GLYPTOSTROBUS (TAXODIACEAE – CONIFERALES)

Seed cones in Taxodium distichum and Glyptostrobus pensilis occupy the position of permanent shoots and are initiated in the summer preceding spring pollination. Morphological features are similar in the two genera, reflecting their close taxonomic relationship. Ovule complexes originate as two (rarely more) ovule primordia in the axil of each fertile bract but without any indication of a preceding discrete ovuliferous scale. When the nucellus, integument, and micropyle are well developed, a series of up to ll abaxial lobes forms at the base of each ovule pair. They become fused by basal growth. After pollination the common basal meristem of lobes and bract extends by intercalary growth to form the conspicuous “ovuliferous scale” of the mature cone; the lobes enlarge and exceed the ovules. Despite the topographic similarity in the cones of both genera, there are differences in vasculature such that the vascular traces to the axillary complex originate directly from the axial cylinder in Glyptostrobus but from the bract trace in Taxodium. The complex vasculature of the mature cone develops late and primarily as an expression of intercalary growth.     

The Cultivation of Hybrid Cotton in Vitro to Overcome the Incompatibility of Hybridization between the Species

2—3 days after cross-pollination the hybrid ovules of cotton (Gossypium hirsutum×G. arboreum) were cultivated in vitro. The hybrid zygot in excised ovule started to divide, frequently passing through several stages to develop into cotyledonous embryos. The embryos were inducted into seedlings. The hybrid plants in blossom were obtained as a result. The histological study was done for ovules of self- pollination in vivo and of cross-pollination in vitro. The early abortion of endosperm in hybrid ovules was observed. In the prophase the nucellus was disintegrated slowly, and in the middle and terminal phase it disintegrated more rapidly. These were identified in hybrid F1. The size of corolla is intermadiate form between female and male. The violet-red spots at the base of petals are similar to male, while there are no such spots in female. None of the pollen can fertilize. Hybrid F1 is not fertile after self-pollination or backcross by the pollen of parent. The chromosome number in root tip Cell of hybrid F1 is 2n=3x=39.     

Influence of intraovular reserves on ovule fate in apricot (Prunus armeniaca L.)

 In many plant species with multiovulate ovaries, a considerable reduction in the number of ovules takes place. However, the underlying physiological causes are not clear. In Prunus spp., although flowers present two ovules, usually only one seed is produced. We have followed the development and degeneration of the two ovules in apricot (Prunus armeniaca L.) and examined the extent to which carbohydrates within the ovule might be involved in determining the fate of the ovule. While the primary ovule grows in the days following anthesis, growth of the secondary ovule is arrested. Starch distribution along the different ovular tissues exhibits several changes that are different in the two ovules. Primary ovule growth is inversely related to starch content and this growth takes place independently of pollination since it occurs in the same way in pollinated and unpollinated flowers. In the secondary ovule, starch disappears simultaneously from all ovular structures and callose is layered at the chalazal end of the nucellus. The size of the secondary ovule does not change significantly from anthesis to degeneration, and callose starts to accumulate 5 days after anthesis. Likewise, this process occurs independently of pollination. These results are discussed in terms of the implications of the starch content of ovules in fertilization success and ovule fate. Received: 26 August 1997 / Revision accepted: 17 December 1997     

Anatomically preserved cardiocarpalean ovules in the late Palaeozoic Angara floral province of Russia

A new species of Cardiocarpus from the Sim River Basin in the southern Urals of Russia is the first anatomically preserved cardiocarpalean ovule from the Permian of the Angara floral province. Specimens are preserved in a soft organic limestone with large numbers of juvenile bactritoid protoconchs. Ovules are roughly ellipsoidal in the major plane, up to 6.0–7.5 mm in length and 5.3 mm in width, with a diminutive wing and a rounded chalaza. The outer surface of the ovules is smooth. In longitudinal sections, two vascular tissue channels traverse the sclerotesta at the chalaza, and appear to extend to near the apex within each wing. There is a simple, dome-shaped pollen chamber at the apex, which lies below a narrow micropyle. The sclerotesta consists of one or two layers of longitudinally orientated cells at the interior, and sclereids that are radially elongated towards the periphery. The sarcotesta is incompletely preserved, but displays an outer layer of cells that are somewhat elongated towards the long axis of the ovule. Numerical cladistic analysis reveals little correlation between the structure of cardiocarpalean morphotaxa and the phylogenetic relationships of the plants that produced them, where affinities are known. Cardiocarpus angarensis sp. nov. enriches our understanding of Permian spermatophytes in temperate climates of the Northern Hemisphere, and reveals that ovules similar to those of Euramerican and Cathaysian provinces characterized an Angaran spermatophyte.  Journal compilation © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 297–305. No claim to original US government works.     

Studies of carboniferous pteridosperm seeds: The relationship of the integument and nucellus in Pachytesta gigantea

Specimens of the medullosan ovule Pachytesta gigantea possess internal ribs which extend from the chalaza up to the level of the pollen chamber. These internal ribs are present adjacent to the primary ribs on the inner surface of the integument and appear histologically similar to the sclerotesta. The cuticle of the endotesta and the nucellus closely conform to the internal ribs in shape and the shrinkage pattern exhibited by the cuticular membranes suggests that the cuticle of the nucellus and cuticle of the endotesta may have been fused to one another along the internal ribs at some stage of development. The implications of this possible nucellar—integumentary fusion are discussed relative to the classification and phylogeny of Paleozoic seeds.     

Seed parasitism redirects ovule development in Douglas fir

Many parasitic species of insects complete their entire development in seeds. They feed off storage reserves within the ovule. These reserves only normally accumulate in fertilized ovules. Consequently, female insects that oviposit their eggs directly into the plant ovule need to be able to select correctly, as unfertilized ovules of conifers normally become so-called empty seed. We provide clear evidence that in conifers, seed-parasitizing insects do not need to discriminate between fertilized and unfertilized plant ovules when ovipositing their eggs. A host-specific insect, the chalcid Megastigmus spermotrophus Wachtl (Hymenoptera: Torymidae), lays its eggs in ovules of Douglas fir (Pseudotsuga menziesii (Mirbel) Franco) before fertilization has taken place in the plant. Oviposition not only prevents the expected degeneration and death of unfertilized ovules, but it induces energy reserve accumulation. Ovules that would otherwise develop as empty seed are redirected in their development by the insect to provide food for the developing larvae. Instead of the insect exploiting normal events during seed development, the insect manipulates seed development for its own reproductive advantage.     

Anatomical diversity of funicles in Leguminosae

To clarify the diversity in funicular internal structures in Leguminosae, 59 legume species (classified into 46 genera, 20 tribes, and 3 subfamilies) were examined by a paraffin-sectioning method. The vascular bundles of legume funicles were clarified as collateral, amphicribral, or amphivasal. In species in which the funicular vascular bundle was collateral throughout the funicle, the xylem is positioned at the pericarpial side in the basal part of the funicle, and the xylem was always positioned at the micropylar side of the phloem in the apical end of the funicles. Whenever the seed direction (from hilum to the micropyle) faces the stylar side, the funicular vascular bundle appears to twist between the basal and the apical part of the funicle. This twist would involve a rotation of the seeds (ovules) during seed (ovule) development. This also may mean that the direction (from hilum to the micropyle) of legume seeds originally faces the pericarp.