Development and structure of the seed-coat of Lannea discolor (Sonder) Engl. (Anacardiaceae)

VON TEICHMAN, I., 1988. The development and structure of the seed-coat of Lannea discolor (Sonder) Engl. (Anacardiaceae). The bitegmic, anatropous ovule contains a group of nucellar cells with slightly thickened and intensively staining cell walls. Besides this hypostase sensu stricto, the nucellus cells in the chalaza become tanniniferous. This tanniniferous chalazal-nucellar tissue is intially plate-like. It is referred to as the hypostase sensu lato. The latter and the chalaza enlarge significantly. The raphe, extensive chalaza and well-developed cup-like hypostase sensu lato play an important role in the development of the seed-coat. The inner, tanniniferous epidermis of the inner integument persists in parts of the mature seed-coat. The outer, distinctly tanniniferous epidermis of the outer integument shows in the mature seed-coat a degree of secondary wall thickening. This undifferentiated type of seed-coat of L. discolor (tribe Spondieae) is remarkably similar to that of Camnosperma minor (tribe Rhoideae), both also showing tendency towards the exotestal type. In the Rhoideae the endotestal, i.e. differentiated type, of seed-coat is also present. The exalbuminous seed of L. discolor represents a derived and advanced type.     

The generic position of Lithraea brasiliensis Marchand (Anacardiaceae): evidence from fruit and seed structure

In Lithraea brasiliensis Marchand the exocarp is characterized by brachysclereids and the parenchymatous mesocarp by large secretory ducts; inner sclerenchymatous ridges are absent in die mesocarp. The stratified endocarp s. s. comprises a crystal layer, palisade-like brachysclereids, osteosclereids and macrosclereids. The osteosclereids are characterized by a distinct light line or linea lucida , which has hitherto also been recorded in a species of Rhus. In the partially pachychalazal seed, a typical Anacardiaceae-like hypostase typifies the chalazal part of the seed coat, while the integumentary seed coat reveals a well preserved outer epidermis, a compressed endotegmen and well developed inner cuticular layer. Our comparison of die characters of the ovule, fruit and seed of L. brasiliensis with those of various species of Rhus and other genera of the tribe Rhoeae (some closely related) presents evidence that L. brasiliensis could be most closely associated with the genus Rhus.     

Taxonomic significance of pericarp and seed structure in Heeria argentea (Thunb.) Meisn. (Anacardiaceae) including reference to pachychalazy and recalcitrance

Heeria argentea (tribe Rhoeae), a monotypic, dioecious tree, is endemic to the core area of the Cape Floristic Region. The mature exocarp consists of a uniseriate layer of palisade-like epidermal cells, interspersed with modified stomata. The mature endocarp sensu stricto develops solely from the inner epidermis. It is essentially two-layered and resembles the state in Protorhus longifolia. This endocarp is here proposed as a distinct fourth endocarpal subtype under the so-called Anacardium -type. The large, pachychalazal, recalcitrant seed develops from the single, anatropous, bitegmic, crassinucellate ovule. This ovule is characterized by an extensive chalaza, vascularization and Anacardiaceae-type hypostase. The pachychalazal seed coat contains abundant vascular bundles and a tanniniferous hypostase. The inner epidermis of the inner integument differentiates into an endotegmen. The contribution of the integuments towards seed coat development is negligible. Concerning characters of the disc in the female flower, the meso- and endocarp, as well as seed size, degree of pachychalazy, nutrient reserves (starch) in the chlorophyllous cotyledons and hypogeal germination, Heeria shows a very close phylogenetic relationship to Protorhus longifolia. However, fruit and seed structure clearly supports the taxonomic separation of Heeria from Ozoroa. Data also support the view that Heeria is a tropical relict, and the hypothesis that pachychalazy, greater seed size, as well as recalcitrant seed viability behaviour constitute ancestral seed character states. Pachychalazy is regarded as a functional adaptation for more efficient transfer of nutrients.     

Notes on the ontogeny and structure of the seed-coat of Sclerocarya birrea (Richard) Hochst. subsp. caffra (Sonder) Kokwaro (Anacardiaceae)

The pendulous, bitegmic, anatropous ovulr with dorsal raphe is suspended at the tip of a massive funicle. A group of nurellar cells with intensively staining cell walls, the hypostase sensu stricto , is present. The initially plate-like tanniniferous chalazal-nucellar tissue, with suberin and lignin impregnated cell walls represents a hypostase sensu lato . The mature seed-coat is formed by the raphe, extensive chalaza, adjacent, well-developed, cup-like hypostase sensu lato , remnants of the two integuments and a cuticular layer. The exalbuminous seed of Sclerocarya birrea suhsp. caffra (the Marula), is regarded to he a derived and phylogenetically advanced type. The undifferentiated seed-roat is very similar to that found in Lannea discolor which, like the marula, belongs to the tribe Spondieae. The similarities in the structure of the seed-coat and seed of the marula and L. discolor confirm their proposed close phylogenetir relationship.     

Ovules and seeds in Acalyphoideae (Euphorbiaceae): structure and systematic implications

Acalyphoideae, the largest subfamily of Euphorbiaceae, are investigated with respect to ovule and seed structure on the basis of 172 species of 80 genera in all 20 tribes of Acalyphoideae sensu Webster. All species of Acalyphoideae examined have bitegmic ovules with a non-vascularized inner integument. However, noticeable differences exist among and sometimes within the genera in the thickness of the inner and outer integument, the presence or absence of vascular bundles in the outer integument, whether ovules are pachychalazal or not, the presence or absence of an aril, seed coat structure (in terms of the best-developed mechanical cell-layer), and the shape of cells constituting the exotegmen. For the latter two characters, two different types of seed coat (i.e., "exotegmic" and "exotestal") and three different types of exotegmic cell (i.e., palisadal, tracheoidal and ribbon-like) were distinguished. Comparisons showed that three tribes Clutieae, Chaetocarpeae and Pereae are distinct from the other Acalyphoideae as well as from the other Euphorbiaceae in having an exotestal seed coat with a tracheoidal exotegmen. The tribe Dicoelieae is also distinct from the other Acalyphoideae in having an exotegmic seed that is composed of ribbon-like cells of exotegmen (i.e., cells both longitudinally and radially elongated, sclerotic and pitted). The tribe Galearieae, which should be treated as a distinct family Pandaceae, is also distinct from the other Acalyphoideae in having an exotegmic seed with a tracheoidal exotegmen (i.e., cells longitudinally elongated, sclerotic and pitted). The remaining genera of Acalyphoideae always have an exotegmic seed with a palisadal exotegmen (i.e., cells radially elongated, sclerotic and pitted). The shared palisadal exotegmen supports the close affinity of Acalyphoideae (excluding five tribes) with Crotonoideae and Euphorbioideae. Within the remaining genera of Acalyphoideae, a significant diversity is found in ovule and seed morphology with respect to the thickness of the inner and outer integument, the size of chalaza, vascularization of an outer integument and an aril.     

Development and structure of the seed of Ozoroa paniculosa (Anacardiaceae) and taxonomic notes

The anatropous, bitegmic and crassinucellar ovule has a nuclear endosperm development. It is further characterized by a hypostase sensu lato. This hypostase being an integral part of the chalaza undergoes a secondary extension with it. At maturity the exalbuminous seed is partially pachychalazal and therefore two anatomically distinct larger parts can be distinguished in the mature seed coat. An endotegmen typifies the integumentary seed coat, while a saddle-shaped hypostase characterizes the chalazal seed coat. This seed coat shows several characteristics of the typical anacardiaceous pachychalazal seed. The cotyledons store lipids and protein as nutrient reserveS. A well-developed cuticle, cuticular layer, cutin and callose in the hypostase cell walls, as well as tannin-like deposits in the seed coat, protect the physiologically ripe seed against dehydration.     

Anatomy and ontogeny of Pterodon emarginatus (Fabaceae: Faboideae) seed.

The aim of this study was to describe the anatomy and ontogeny of Pterodon emarginatus seed using the usual techniques. The ovules are campilotropous, crassinucelate, and bitegmic. The following processes occur during integument development: anticlinal divisions and phenolic compound accumulations in the exotesta, whose cells become palisade; predominantly periclinal divisions and cell expansion in the mesotesta, where the rapheal bundle differentiates; differentiation of the hourglass-cell layer adjacent to the palisade; fusion of outer and inner integuments, which remain individualized structures only at the micropylar end; and intense pectin impregnation in the mesotesta thicker walls with lignification restricted to the xylem. At the hilar pole, the Faboideae seed characteristic structure develops, with double palisade layer, subhilar parenchyma, and tracheid bar. The younger nucellus shows thicker pectic cell walls and is consumed during seed formation. The endosperm is nuclear and, after cellularization, shows peripheral cells with dense lipid content; the seeds are albuminous. The axial embryo shows fleshy cotyledons, which accumulate lipid and protein reserves; starch is rare. Although the seed structure is characteristic of the Fabaceae, the inner integument coalesces into the outer integument without being reabsorbed.     

The ontogeny and structure of the pericarp and seed-coat of Harpephyllum caffrum Bernh. ex Krauss (Anacardiaceae)

The exocarp sensu lato , which develops from the outer epidermis and adjacent parenchyma of the ovary wall, consists of collenchyma cells with a stomatous epidermis. The fleshy, parenchymatous mesocarp or sarcocarp develops after endocarp differentiation. The endocarp is partly spongy and partly woody. The spongy endocarp contains most of the vascular tissue and fills the cavities and grooves of the intricately sculptured outer woody endocarp. The inner woody endocarp and adjacent woody, endocarpal operculum develop from the inner epidermis and subepidermal parenchyma of the ovary wall. The bitegmic, anatropous ovule develops into a derived, exalbuminous seed with an undifferentiated seed-coat. An extensive chalaza, extensive hypostase sensu lato and the raphe are important in the development of the seed-coat. The pericarp and seed-coat of H. caffrum is compared with those of Sclerocarya birrea subsp. caffra and Lannea discolor . The close phylogenetic relationship of these three species of the Spondieae is reaffirmed. The marked similarities in pericarp and seed structure between H. caffrum and species of the genus Spondias are noted.     

Systematic and phylogenetic significance of the seed coat in the shrubby African Iridaceae, Nivenia, Klattia and Witsenia

MANNINGJ. C. & GOLDBLATT, P., 1991. Systematic and phylogenetic significance of the seed coat in the shrubby African Iridaceae, Nivenia, Klattia and Witsenia. The seeds of Nivenia, Klattia and Witsenia, a natural alliance within Nivenioideae, are among their most distinctive features, and seed number and shape constitute at least two synapomorphies for this monophyletic lineage. In this paper we survey the structure of the seed surface of five species of Nivenia and one each of Klattia and Witsenia by scanning electron microsopy (SEM), and study the development of the seed coat in a representative species of each of the three genera. Outgroups for comparison were selected from genera inside and outside the subfamily. The six genera of Nivenioideae are united in having a 3 (– 2) seriate outer integument but the shrubby genera are unique in the family in their tangentially flattened ovules which mature into depressed, flat, scutiform seeds. They display a range of variation in seed coat characters but are specialized in the subfamily and family in having a transparent outer integument. Other genera have pigmented deposits in the outer epidermis of the outer integument. Nivenia and Witsenia are further specialized in having a partially exfoliating outer integument. Klattia is unique among the shrubby genera in having deposits in the second layer of the tegmen, in retaining an intact outer epidermis of the tegmen, and in the fusiform shape of the epidermal cells and striate surface of the seed. Our results are incorporated in a cladogram that expresses our current understanding of the phylogenetic relationships of Nivenioideae.     

Structure and development of the ovule and seed in Aristolochiaceae, with particular reference to Saruma

All members of Aristolochiaceae have anatropous, bitegmic, crassinucellate ovules, which are endostomic except in Saruma and Asarum arifolium where ovules are amphistomic. The outer integument is two cell-layered and the inner integument is three cell-layered. The chalazal megaspore is the functional one. All these conditions appear to be plesiomorphic for the order Piperales, which consists of five families, Aristolochiaceae, Hydnoraceae, Lactoridaceae, Piperaceae and Saururaceae. The embryo sac in Aristolochiaceae is eight-nucleate and corresponds to the Polygonum type; a hypostase is frequently present in this family. The seed coat of Aristolochia s.l., Asarum, Saruma and some Thottea species consists primarily of a two cell-layered testa, and a three cell-layered tegmen. In some species the cells of the outer epidermis become radially elongated, forming reticulate wall thickenings. Cells of the inner layer of the testa have crystals and thickened inner walls. The three layers of the tegmen are tangentially elongated, and become cross fibres at maturity, as fibres of the outer and inner layers are parallel to the seed axis, whereas those of the middle layer are perpendicular to it. This type of seed coat anatomy is synapomorphic for Aristolochiaceae. In addition, the gross morphology of the seed and elaiosome histology are remarkably similar in Asarum and Saruma, thus supporting a sister-group relationship between them. Embryological and seed characters do not supply any synapomorphy that support a close relationship between Aristolochiaceae, Hydnoraceae and Lactoridaceae. Instead, some seed features such as the absence of seed appendages and the collapsed cells of endotesta may indicate a close relationship of Lactoris with Piperaceae plus Saururaceae, although this is the subject of further analysis.     

长豇豆种皮和种脐的发育

长豇豆的胚珠具内外两层珠被,内珠被在种子发育早期退化消失,种皮仅由外珠被发育而成。外珠被的外表皮细胞径向伸长,外壁和经向壁增厚,形成约占成熟种皮厚度一半的栅栏层;亚表皮细胞发育为骨状石细胞层。第三层细胞类似于亚表皮层但细胞壁增厚不明显,其内方的多层薄壁细胞形成海绵组织。种脐具两层栅栏细胞,外栅栏层及其以外部分由珠柄组织发育而成管胞群。本文还对脐缝和管胞群的作用以及豆科种子的吸水机制进行了讨论。     

Development of ovule and seed in Rapateaceae

VENTURELLI, M. & BOUMAN, F., 1988. Development of ovule and seed in Rapateaceae. The structure of the ovules and/or seeds of twelve species of Rapateaceae were studied, some additional embryological characters also being recorded. The ovules are always anatropous, bitegmic and crassinucellate, but they differ in the shape, size and in thickness of the outer integument. In Rapateaceae the outer integument is initiated subdermally. The seed coat of the Rapateaceae shows two mechanical layers: an endotesta with silica present as bodies or as incrustations in cell walls, in conjunction with an exotegmen with a jigsaw cell pattern complicated by a labyrinth-like sculpturing of the outer cell walls. The innermost layer of the inner integument is tanniniferous. Large hilar scars with tracheidal plates on the corresponding fruit wall and a persistent obturator are recorded in Rapateaceae. On the basis of embryological characters the family fits well into the Commelinales. Testa structure most closely resembles that of the Commelinaceae. The differences in ovule and seed structure agree with the currently accepted tribal classification.     

Embryology of Siparunaceae (Laurales): characteristics and character evolution

Embryological characters of Siparunaceae, which are poorly understood, were studied on the basis of two constituent genera, an African Glossocalyx and a South American Siparuna, to better understand their evolution within Laurales. These two genera have many embryological characteristics in common with the other lauralean families. Noticeably, they share the multi-celled ovule archesporium (uncertain in Glossocalyx) as a synapomorphy with all the other lauralean families except Lauraceae, the anthers dehisced by valves as a synspomorphy with all the other lauralean families except Calycanthaceae and Monimiaceae, and the bisporangiate anther as a synapomorphy with Gomortegaceae and Atherospermataceae. Siparunaceae are, however, distinct from all other laularean families in having unitegmic ovules that were derived from bitegmic ovules, probably due to an elimination of the outer integument. Likewise, the lack of the testa (i.e., developed outer integument), the "endotegmic" seed coat, and the perichalazal seed at maturity are also characteristics of Siparunaceae. Within the family, Siparuna differs from Glossocalyx in having plural tetrads of megaspores and plural, starchy-rich, one-nucleate, tubular embryo sacs (autapomorphies). On the other hand, Glossocalyx is characterized by having bilaterally flattened seeds (autapomorphy). Although functional aspects of those autapomorphies are uncertain, both Glossocalyx and Siparuna show evolution in different embryological characters.     

SYSTEMATIC SIGNIFICANCE OF SEED-SURFACE FEATURES IN ORTHOCARPUS (SCROPHULARIACEAE—SUBTRIBE CASTILLEJINAE)

Scanning electron microscope and light microscope examination of seed-coat features of 26 species of Orthocarpus have allowed recognition of many species-level differences (summarized in a key) and of three seed-coat types that parallel taxonomic subgroups but support realignments at generic and infrageneric levels. Type 1 seeds (subg. Orthocarpus, sect. Orthocarpus) have a lateral hilum, sculptured inner tangential seed-coat walls, and a tightly fitting outer seed coat. They are very similar to seeds of Cordylanthus. Seeds of Types 2 and 3 have a terminal hilum and membranous inner tangential cell walls. Type 2 seeds (subg. Orthocarpus, sects. Castillejoides and Cordylanthoides, with one exception) have a net-like, loosely fitting outer seed coat that shows close relationship to seeds of Castilleja. Inner tangential walls of Type 2 seeds normally rupture. Type 3 seeds (subg. Triphysaria, with two exceptions) have a tightly fitting outer seed coat and inner tangential walls are always retained. Seed features support evidence from floral morphology and chromosome numbers that Orthocarpus as currently recognized is not a monophyletic lineage.     

Embryology of the Theaceae - anther and ovule development of Camellia, Franklinia, and Schima

The anther and ovule development of Camellia, Franklinia, and Schima (Theaceae, Camellioideae) were observed. The three genera share the following embryological traits: anther wall formation of basic type, tapetum of glandular type, walls of endothecial cells with secondary thickening, and production of pseudopollen grains in connective, which are dispersed into pollen sacs at anthesis, ovule bitegmic-tenuinucellate, micropyle formed by inner integument alone, hypostase present, and both integuments generally five-to-seven cell layered. One autapomorphy of the Camellioideae found in the present study is the production of pseudopollen. The three genera surveyed differ with respect to the number of middle layers in the anther, the presence or absence of stomata on connective epidermis, morphology of pseudopollen, type of embryo sac formation, form of ovule, ovular vasculature, and the proliferation of ovular epidermis, etc. Among the three genera, Franklinia and Schima are presumed to be closer embryologically, and Schima possesses more numerous specialized features.     

Gone and ovule ontogeny in Phyllocladus (Podocarpaceae)

TOMLINSON, P. B., TAKASO, T. & RATTENBURY, J. A., 1989. Cone and ovule ontogeny in Phyllocladus (Podocarpaceae). Cones are borne directly on phylloclades, usually in the position of basal segments or as segment appendages. Each cone consists of a series of spirally arranged bracts, of which the middle bracts each subtend a single, sessile ovule. There is no ovuliferous scale. Ovules arise as ovoid outgrowths; integument development involves periclinal divisions of hypodermal cells with the integument becoming bilobed and extended laterally. The mature ovule is flask-shaped. The integument includes an extensive middle region bounded by an inner and outer epidermis; the outer hypodermis is differentiated as two contrasted cell layers. An aril differentiates late by periclinal divisions of the outer hypodermal cells at the base of the ovule. The three outermost layers of the integument become differentiated in the mature seed as an epidermis, with thick, cutinized outer tangential walls, an outer hypodermal tanniniferous layer and a sclerotic inner layer. Each ovule is vascularized by two strands that diverge from the axial bundles delimiting the gap left by the departing bract trace.     

Ovules and seeds in Euphorbioideae (Euphorbiaceae): structure and systematic implications

Ovule and seed structure in Euphorbioideae, one of the five euphorbiaceous subfamilies, is surveyed to evaluate its systematic implications on the basis of 79 species representing four of five tribes. All Euphorbioideae, like two other "uniovulate" subfamilies Acalyphoideae and Crotonoideae, but unlike most of two "biovulate" subfamilies Oldfieldioideae and Phyllanthoideae, consistently have a persistent and palisadal exotegmen composed of radially elongate, sclerotic, and pitted cells. Within Euphorbioideae, the tribe Stomatocalyceae (also with the palisadal exotegmen) is unusual in having vascular bundles in outer integument and clearly distinct from the remaining Euphorbioideae and the other "uniovulate" subfamilies. With the exclusion of Stomatocalyceae, Euphorbioideae are not anatomically divided into major groups such as a pseudanthial and a non-pseudanthial clade, but instead have some remarkable diversity within a tribe, a subtribe, and even a genus in the three ovule and seed characters: (1) the thickness of the inner integument, (2) the thickness of the outer integument, and (3) the presence or absence of an aril. Groups of genera and species wrapped by different combinations of their characteristics, however, are not necessarily harmonized with tribal or subtribal classifications available. Anatomical similarities and dissimilarities presented in this paper, as well as relationships among taxa presented in the classifications available, will be critically evaluated in the light of results of ongoing molecular phylogenetic analyses. Electronic Publication     

Embryo sac, endosperm, and seed of Nemophila (Boraginaceae) relative to taxonomy, with a remark on embryogeny in Pholistoma

Studies on embryology and seed morphology are complementary to molecular phylogenetics and of special value at the genus level. This paper discusses the delimitation and evolutionary relationships of genera within the tribe Hydrophylleae of the Boraginaceae. The seven Nemophila species characterized by a conspicuous seed appendage are similar in embryology and seed structure. The ovule is tenuinucellate and unitegmic with a meristematic tapetum. The embryo sac penetrating the nucellar apex is of the Polygonum type, has short-lived antipodal cells, and an embryo sac haustorium. The endosperm is cellular, producing two terminal endosperm haustoria, of which the chalazal has a lateral branch. Embryogeny is of the Chenopodiad type (as in Pholistoma). The seed coat is formed from the small-celled inner epidermis of the integument. The large-celled outer epidermis of the integument disintegrates into scattered cells. Seed pits evolve from irregularly placed inner epidermal cells of the integument. The chalazal part of the ovule produces a cucullus, that functions as an ant-attracting elaiosome. Those species of Nemophila with a conspicuous cucullus form a natural genus. Nemophila is most closely related to Pholistoma. The integumentary seed pits of Nemophila might have evolved from ovular seed pits similar to those in Pholistoma.     

Early Development of the Seed Coat of Soybean (Glycine max)

Although the development of the soybean ovule has been fairlywell studied, knowledge of the sequence of events in the seedcoat during the first 3 weeks after flowering is incomplete.The goal of the present study was to document, using light microscopy,the early development of the soybean seed coat with respectto changes in structure and histochemistry. At anthesis, theseed coat consists of an outer layer of cuboidal epidermal cellssurrounding several layers of undifferentiated parenchyma (whichtogether constitute the outer integument), and an inner layerof cuboidal endothelial cells (the inner integument). At 3 dpost anthesis (dpa), the inner integument has expanded to includethree to five layers of relatively large cells with thick, heavily-stainingcell walls immediately adjacent to the endothelium. By 18 dpa,the outer integument has developed into a complex of tissuescomprised of an inner layer of thick-walled parenchyma, an outerlayer of thin-walled parenchyma containing vascular tissue whichhas grown down from the lateral vascular bundles in the hilumregion, a hypodermis of hourglass cells, and palisade layer(epidermis). The thick-walled parenchyma of the inner integumenthas become completely stretched and compressed, leaving a single,deeply staining wall layer directly above the endothelium. At21 dpa, the outermost cells of the endosperm have begun to compressthe endothelium. At 45 dpa (physiological maturity) the seedcoat retains only the palisade layer, hourglass cells, and afew layers of thin-walled parenchyma. The innermost layer ofthe endosperm, the aleurone layer, adheres to the inside ofthe seed coat. This knowledge will be invaluable in future studiesof manipulation of gene expression in the seed coat to modifyseed or seed coat characteristics. Copyright 1999 Annals ofBotany Company Soybean, Glycine max, seed coat, development, aleurone.