The embryology ofStegnospermataceae,with a discussion on its status,affinities and systematic position

The embryology ofStegnosperma halimifolium andS. watsonii has been studied in detail. The tapetum is of the secretory type and its cells become multinucleate. Simultaneous cytokinesis in the pollen mother cells follows meiosis. The ripe pollen grains are 3-celled. The ovule is crassinucellate, bitegmic and amphitropous, with the micropyle formed by the inner integument alone. The female archesporium is one celled, and the parietal tissue 3–5 layered. The embryo sac development conforms to thePolygonum type. A central strand, 6 or 7 cells thick, differentiates inside the nucellus and extends from the base of the embryo sac to the chalazal region. The endosperm is nuclear. The embryogeny conforms to the Caryophyllad type. The seed coat is formed by the outer epidermis of the outer integument and the inner epidermis of the inner integument. Based on this evidence and other data, the status of the genus as an independent family,Stegnospermataceae (Stegnospermaceae) is confirmed. Apparently, it forms a connecting link betweenPhytolaccaceae andCaryophyllaceae.     

<Emphasis Type="Italic">Gypsy embryo</Emphasis> specifies ovule curvature by regulating ovule/integument development in rice

The embryo position in a seed is stable in most plant species, indicating the existence of a strict regulatory mechanism that specifies the embryo position in the seed. To elucidate this mechanism, we analyzed the gypsy embryo (gym) mutant of rice, in which the position of the mature embryo in the seed is altered at a low frequency. Analyses of early embryogenesis and ovule development showed that the ectopic embryo was derived from an ill-positioned egg cell, which resulted from the incomplete curvature of the ovule. Although the development of both the inner and outer integuments was impaired, the ovule curvature was associated closely with the extent of inner integument growth. Therefore, inner integument development controls ovule curvature in rice. The expression patterns of OSH1 and OsMADS13 indicated that, in gym, a small number of indeterminate cells are maintained on the style side of the ovule and then in the integument primordium at a low frequency. The prolonged survival of these indeterminate cells disturbs normal integument development. The gym fon2 double mutant suggests that GYM and FON2 are involved redundantly in floral meristem determinacy. Possible functions of the GYM gene and the ovule developmental mechanism are discussed.     

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.     

Embryology of theaceae — Anther and ovule development ofAdinandra,Cleyera andEurya

Anther and ovule development of the theaceous Ternstroemioideae is reported for the first time on the basis of eight specles of three generaAdinandra, Cleyera andEurya. Anthers of these three genera are similar and can be characterized by the following traits: tapetum of glandular type, anther dehiscing latrorse-introrse, both connective and anther epidermis heavily tanniniferous, and connective and even anther wall layers having abundant druses. Their ovules are also very similar in being bitegmic and tenuinucellate, and in having a micropyle formed by the inner integument only, three cell-layered integuments, a raphe bundle terminating at chalaza, usually amphitropous or less often campylotropous ovule, embryo sac formation of Polygonum type, ephemeral antipodal cells, and tanniniferous ovule epidermis. Such ovules are readily distinguishable from those of Camellioideae and all other families. It is suggested that the three genera studied are closely related, and that the degree of embryological specialization is highest inAdinandra and lowest inEurya. On the basis of the significant embryological discrepacies, the Ternstroemioideae seem to have diverged rather distantly from the other core-subfamily Camellioideae of the Theaceae.     

Development and structure of the comose seeds ofHillia (Rubiaceae)

The hairs at the apical end of the seeds ofHillia are pluriseriate, multicellular structures. The cells making up a hair are elongated exotesta cells and, consequently, also have secondary thickenings identical (H. parasitica) or similar (H. costanensis) to those found on the exotesta cells on the main body of the seeds. Hair formation already starts in bud stage: at and around the chalazal region of an ovule, integument epidermis cells are grouped together to form ± elongated packets of 4–7 cells. The cells of each packet undergo further elongation and anticlinal division so that a hair on a mature seed may be up to c. 30 mm long. Basally, the seeds have a tail- to ± wing-like appendage, made up of only two cell layers, the exotesta of the ab- and adaxial side of the seed. This basal appendage shows the same anatomical structure as the wings of various anemochorous rubiaceous seeds. Although seed hairs of this kind are unique in theRubiaceae and — from the point of development and structure — not homologous to exotesta wings, the presence of a basal wing-like appendage suggests thatHillia, previously often placed into a tribe of its own (Hillieae), can be accommodated in theCinchoneae, a tribe in which winged, anemochorous seeds predominate. The tufts of hairs of the comose seeds ofHillia look superficially similar to those of certainAsclepiadaceae andApocynaceae (like theRubiaceae belonging to the orderGentianales). Comparisons based on literature data, however, reveal that there are striking differences in the position, development and structure of the hairs (produced at the micropylar end, initiated after fertilization, hairs unicellular, etc.).     

Studies on systematic embryology inScilla (Hyacinthaceae)

Scilla persica and 5 species of the so-calledS. hohenackeri group, namely,S. furseorum, S. puschkinioides, S. vvedenskyi, S. hohenackeri, andS. greilhuberi, have been investigated embryologically with special reference to embryo sac and endosperm development.Polygonum-type embryo sac development was stated inS. puschkinioides andS. greilhuberi. 8-nucleate, normally structured embryo sacs, which could not be specified further due to sparse availability of the material, were stated inS. furseorum, S. vvedenskyi, andS. hohenackeri. InS. persica the embryo sac develops according to the bisporicAllium-type. In most species endosperm development was stated to be nuclear, exceptS. hohenackeri, where the type could not be specified. Other traits of possible taxonomic significance are the number of layers in the outer integument, which is mostly 4, or 5–6 inS. furseorum, and the occurrence of polyploid versus haploid and early degenerating antipodal nuclei, the latter occurring only inS. persica andS. furseorum. These embryological characters may be useful for assessing taxonomic relationship of the present species with other allied groups withinScilla, in particular, theS. siberica alliance,S. messeniaca, and theS. bifolia alliance. TheAllium-type embryo sac, which occurs inS. persica, is also characteristic for theS. siberica alliance, and may be a common derived character. Lack of antipodal polyploidization, as characteristic forS. persica andS. furseorum, occurs also in theS. siberica alliance, and is perhaps another common derived trait indicating phylogenetic relationship. Nuclear endosperm development is more frequent in spring-flowering squills than helobial development, which has previously been stated inS. messeniaca, some species of theS. siberica alliance, and inS. litardierei. While helobial endosperm may be primitive forHyacinthaceae in general, it may, by reversal, also occur as a derived character, at least in some species of theS. siberica alliance.     

Embryogenesis and seed development in Sinomanglietia glauca (Magnoliaceae)

The development of the floral bud, especially the ovule and seed coat, of Sinomanglietia glauca was observed. Floral buds were covered by eight to nine hypsophyll pieces. The hypsophyll nearest the tepal was closed completely and characterized by two arrays of densely stained cells with dense cytoplasm, which split longitudinally at flowering. The perianth consisted of 16 tepals arranged in three whorls. The gynoecium was composed of numerous apocarpous carpels; the ovule was anatropous with two integuments. Embryogenesis was of the Polygonum type, and the endosperm was nuclear. The inner integument degenerated during seed development. The seed of S. glauca had an endotestal seed coat comprised of a sclerotic layer derived from the inner adaxial epidermis of the outer integument and a sarcotesta derived mainly from the middle cells between the inner and outer epidermis of the outer integument. The embryo developed normally, so embryogenesis is not the cause of difficult regeneration.     

The embryology and relationships ofOliniaceae

Two of the five species ofOliniaceae (Olinia emarginata andO. ventosa), a monotypic and problematic family of theMyrtales, were investigated embryologically.Oliniaceae clearly agree with otherMyrtales in their basic embryological characteristics, and are characterized further by having an ephemeral endothecium, a campylotropous ovule, and a thick, three-five-layered, outer integument. A combination of these three characteristic features is unknown elsewhere inMyrtales, so that embryological features do not support a close relationship with any other member of the order. Shared distinctive anther characteristics (i.e. ephemeral endothecium) suggest thatOliniaceae are derived from the common ancestor ofCrypteroniaceae s. str.,Rhynchocalycaceae, Alzateaceae, andPenaeaceae.     

Studies on the embryology and gynoecium structures inDrimys winteri (Winteraceae) and someAnnonaceae

Drimys winteri (Winteraceae) and 11 species ofAnnonaceae, namelyAnnona montana, Artabotrys hexapetalus, Bocagea sp.,Papualthia sp.,Polyalthia nitidissima, Tetrameranthus umbellatus, T. duckei, Uvaria sp.,Xylopia malayana, X. aromatica, andX. emarginata, were investigated embryologically with special reference to development of ovule and embryo sac. The ovules are anatropous, crassinucellate and in most taxa bitegmic. The inner integument is of epidermal origin. TheAnnonaceae investigated have a multi-layered, later vascularized outer integument with most probably subepidermal initiation. In contrast,Drimys winteri has a three-layered, non-vascularized outer integument of epidermal origin. The annonaceous genusTetrameranthus (T. umbellatus andT. duckei) possesses a middle integument between the inner and the outer one, stated here for the first time in a neotropic representative ofAnnonaceae. Within the angiosperms this feature occurs inAnnonaceae only. The embryological characters are rather homogeneous. Differences between the species investigated are found in, e.g. the number of cell layers in the inner integument, which is commonly two inAnnonaceae as compared to three inDrimys winteri, the presence or absence of a hypostase, the number of layers in the nucellar epidermis, great differences in size of ovules, and the species-specific pattern of tannin deposition in the ovules. In the species so far investigated the embryo sacs develop according to thePolygonum-type. InXylopia malayana andBocagea sp. in addition the carpels were investigated. They are conduplicate. InXylopia malayana the free carpels are united by an extragynoecial compitum, inBocagea sp. each stigma produces its isolated mucilage cap. The results obtained from the investigated taxa are discussed and compared with published data on embryology and gynoecium structure in other annonaceous and winteraceous taxa.     

Interspecific crosses inHordeum (Poaceae)

A crossing programme including 30 species and 40 cytotypes within the genusHordeum was undertaken. Viable hybrids were obtained in 302 combinations, 15 of which were intraspecific. Differences in seed set and in germination were observed in crosses between different groups of species. Obtaining crosses between different taxonomic groups was generally more difficult when diploid material was used. Some species, e.g.,H. lechleri, H. jubatum, andH. brachyantherum showed a higher crossability than others. The chromosome numbers of the hybrids were usually those expected from the parental numbers but aneuploid series around the expected numbers were rather frequent. Three cases of unreduced gametes were found. Selective chromosome elimination was restricted to combinations including eitherH. vulgare orH. bulbosum.—Despite a very diverse morphology, all South American diploid species together with the two North American diploidsH. intercedens andH. pusillum appear to be closely related. The hexaploid American speciesH. procerum, H. lechleri, andH. arizonicum are also related. The two North American tetraploid speciesH. jubatum andH. brachyantherum sometimes form semifertile hybrids. The Asiatic speciesH. roshevitzii appears to be related to both North and South American taxa.     

Ant pollination of the palaeoendemic dioeciousBorderea pyrenaica (Dioscoreaceae)

The importance of ants for pollination in the dioeciousBorderea pyrenaica (Dioscoreaceae), a Tertiary palaeoendemic plant of the Pyrenees (NE Iberian Peninsula) was studied. The frequency of different visitors (ants, lady beetles, andDiptera) to staminate and pistillate flowers was quantified, and their effectiveness as pollinators was examined by means of fruit and seed set in selective experimental exclusions. Although ants were less abundant on flowers than other visitors, they were the most effective pollinators. Some qualitative factors of this mutualistic ant-plant interaction may account for their effectiveness: the small size of the ants, their high visitation rate to pistillate flowers, and the lack of reduction in viability of the pollen transported on the integument. In addition, the sedentary nature of ants assures their presence during the flowering period. The most abundant floral visitors ofB. pyrenaica were therefore not the most effective pollinators.     

Insertion sites of the transposable elementmariner are fixed in the genome ofDrosophila sechellia

Summary The abundance of the transposable elementmariner differs dramatically in the genomes of the closely related speciesDrosophila simulans, D. mauritiana, D. sechellia, andD. melanogaster. Natural populations ofD. simulans andD. mauritiana have 1–10 and 20–30 copies per diploid genome, respectively, and the insertion sites are polymorphic. The element has not been found inD. melanogaster. In this paper we show thatD. sechellia, a species endemic to the Seychelles Islands, contains only twomariner elements that are at fixed sites in the genome. One element, inserted in chromosome 2R at 51A1–2, contains three deletions and is missing much of the 3 end. The other element, inserted in chromosome 3L at 64A10–11, is the full length of 1286 bp. Although the sequence of the full-length element is polymorphic in populations ofD. sechellia, at least some of the sequences are closely related to elements fromD. simulans andD. mauritiana that are known to be active. However, judging from the progeny of crosses betweenD. sechellia andD. simulans, the biological activity of the full-lengthD. sechellia element appears to be low, either because of the nucleotide sequence of the element or because of its position in the genome, or both.     

濒危植物马蹄参的大孢子发生和雌配子体发育

为探讨马蹄参(Diplopanax stachyanthus Hand.-Mazz.)濒危机制与雌性生殖发育的关系,采用石蜡切片法观察马蹄参大孢子形成和雌配子体的发育过程。结果表明,马蹄参雌蕊单心皮,子房下位,1室,1枚胚珠。胚珠横生于短片状胎座上,具单珠被,厚珠心。单孢原细胞自珠心1~2层表皮细胞处分化;大孢子四分体为直线形。成熟胚囊中,2个极核在受精前融合为次生核,3个反足细胞不发达,较早退化;二核胚囊时期出现二核分裂不均且较小核退化消失的异常发育现象。因此,马蹄参雌配子体发育过程中出现异常现象是造成其结实率低的主要原因。     

Adaptation of exotic Azolla to tropical environment of Thailand

There are seven known species of Azolla, two of which have been used in cultivated systems, the tropical speciesA. pinnata, and the temperate speciesA. filiculoides. OnlyA. pinnata is indigenous in Thailand. In this study the two exotic species,A. caroliniana andA. microphylla, were evaluated under the various tropical field conditions in Thailand. When compared with seven selected strains ofA. pinnata under three field conditions,A. caroliniana andA. microphylla were similar to the indigenous species, in terms of growth performance, N₂ fixation, and yield. This study suggests thatA. caroliniana andA. microphylla can be successfully cultivated as a nitrogen fixing green manure for rice production in the tropics.     

Embryology ofCrocus thomasii (Iridaceae)

The embryology ofCrocus thomasii is described. Male meiosis is of simultaneous type, and gives rise to starchy microspores which develop into lipoid pollen grains; these are two-celled and show a spinulate acolpate, abaculate exine lacking apertures. The tapetum is glandular and its cells become bi- or sometimes multinucleate. The ovule is anatropous and bitegmic; the inner integument forms the micropyle. Megasporogenesis is heteropolar with starch accumulation in the functional chalazal megaspore. Embryo sac development conforms to thePolygonum type. The endosperm development is nuclear. The embryo develops according to the Caryophyllad type. In the ripe seed it is differentiated and enveloped by a starchy cellular endosperm. The embryological characters observed strongly favour a close relation betweenC. thomasii andC. sativus.     

THE FEMALE GAMETOPHYTE OF THREE SPECIES OF GRINDELIA AND OF PRIONOPSIS CILIATA (COMPOSITAE)

In the four species studied, Grindelia stricta ssp. blakei, G. arizonica, G. lanceolata, and Prionopsis ciliata (Haplopappus ciliatus), the female gametophyte develops according to the Polygonum (normal) type from the chalazal megaspore of a row of four. In most cases only two antipodal cells are formed, the micropylar one originally containing two nuclei. The number of nuclei increases in both cells; one or both antipodal cells typically grow laterally into the integument, often extending to near the surface of the ovule. This resembles the condition previously reported in Grindelia squarrosa. Since the four species of Grindelia have similar antipodal outgrowths, such outgrowths may be considered typical for this genus. The fact that Prionopsis resembles Grindelia in regard to outgrowths from the antipodal cells and differs in this respect from typical Haplopappus helps to justify its separation from Haplopappus, and perhaps its suggested merger with Grindelia.     

Diversity of seed storage protein patterns in wild peanut (Arachis,Fabaceae) species

55 accessions of wild peanuts (Arachis spp.) introduced from South America were analyzed for seed storage protein composition using SDS-PAGE electrophoresis. The objectives of the study were to evaluate variability within sect.Arachis and to classify taxa based on protein composition. 25 different band positions were resolved. Individual accessions had 11 to 18 bands which included the conarachin region (MW > 50 kD), two to five bands in the acidic arachin region (MW 38–49.9 kD), three to seven in the intermediate MW region (23 to 37.9 kD), two to five bands in the basic arachin region (18–22.9 kD), and one to three bands in the low MW protein region (14–17.9 kD). These data were utilized in a principal coordinate analysis based on the matrix of genetic distances between all pairs of the 55 accessions. Several groups of accessions conformed to expected species classification includingA. batizocoi, A. stenosperma, andA. monticola; whileA. duranensis, A. cardenasii, A. helodes, andA. correntina did not form good groups. The study showed that great diversity exists for protein profiles and seed storage proteins have potential for aiding species classification and for serving as markers for interspecific hybridization studies.     

Embryology of tribeDrypeteae, an enigmatic taxon ofEuphorbiaceae

The tribeDrypeteae, whose traditional assignment inPhyllanthoideae ofEuphorbiaceae is now doubtful, is studied embryologically on the basis of a literature survey and examination of six additional species in two of the four constituent genera.Drypeteae are characterized by having several embryological features that are unknown in otherPhyllanthoideae, such as a two- or three-celled ovule archesporium; a thin, two cell-layered parietal layer in the nucellus; no nucellar beak or cap; an early disintegrating nucellar tissue; thick, multiplicative, inner and outer integuments; an endothelium; a few discrete vascular bundles in the outer integument; and a fibrous exotegmen (or its derived state). EmbryologicallyDrypeteae do not fit within thePhyllanthoideae and, as available nucleotide sequence data from therbcL gene suggest, are rather placed nearErythroxylaceae, Rhizophoraceae, Chrysobalanaceae, andLinaceae. Drypeteae share with those families a combination of the fibrous exotegmen, the endothelium, and the thick, multiplicative inner integument.     

Agar quality of commercial agarophytes from different geographical origins: 1. Physical and rheological properties

Six economically important species ofGracilaria, from a number of commercial sources around the world, andGracilariopsis lemaneiformis, collected from two Japanese localities, were used as the sources of raw material for the evaluation of agar quality. Agar-agar was extracted by pretreatment with various concentratrions of NaOH (0%, 3%, 5%, 7%, 10%) incubated at 80 °C for 2 h. Agar yield, viscosity, dynamic gelling and melting temperature and gel texture were determined for 1.5% agar gels. The highest agar yield was obtained fromG. gracilis from Argentina (39.5%), while the lowest was from BrazilianG. gracilis (13.37%). Dynamic gelling temperature was highest in the agar fromG. gracilis from Turkey (59 °C) and lowest in the non-alkali treated agar isolated fromG. edulis from Indonesia (46 °C). Melting temperature ranged from 96 °C in the agars from the JapaneseGracilariopsis andG. chilensis from Chile to 69 °C in the non-alkali treated agar fromG. edulis from Indonesia. In general, all species produced an agar with high gel strength after treatment with 5% NaOH, except forG. chilensis and the twoGracilariopsis species, which produced an agar with high gel strength after treatment with 3, 7 and 10% NaOH. The highest gel strength (2056 ± 13.6 cm^–2) and hardest gel (261 ± 19.89 g mm^–2) were obtained fromG. lemaneiformis from Japan (Oita Prefecture) after treatment with 7 and 10% NaOH respectively. The lowest gel strength (351 ± 93 cm^–2) was obtained fromG. gracilis from Brazil after treatment with 3% NaOH. The softest gel (66.31 ± 9.63 g mm^–2) was isolated fromG. tenuistipitata from China, after treatment with 3% NaOH. The most flexible gel (11.62 ± 0.31 g mm^–2 × 10²) was obtained fromG. chilensis from Chile after treatment with 3% NaOH.Author for correspondence     

Comparison between the soil seed banks of a burnt and an unburnt Quercus pyrenaica Willd. forest

Soil samples from burnt and unburnt sites dominated by Quercus pyrenaica Willd. forest in León province (NW Spain) were collected, separated into an upper layer (0–2 cm depth) from a lower layer (2–5 cm), put in a greenhouse, and seeds allowed to germinate.A total of 670 identified seedlings comprising 56 species and belonging to 24 families were recorded. Most seedling were herbaceous perennials (hemicryptophytes: 59%), followed by annuals (therophytes: 23%). Many species that germinated from buried seeds were not observed as plants at any site, and came from exogenous communities. The principal means of seed dissemination were anemochory (45%) and autochory (23%).The number of species as well as seedlings was not significantly different between soils from the burnt and unburnt sites, but seedlings were more numerous in samples from the unburnt sites, as the number of seedlings was always highest in the upper soil layer. Germination behaviour of four particular species was characterized.Seedlings from the seed bank temporarily were found in the early stages of recovery of the burnt Quercus pyrenaica forests.