Embryology of Gomortegaceae (Laurales): characteristics and character evolution

The embryological characteristics of Gomortegaceae, which are poorly understood, were investigated on the basis of Gomortega nitida, the only species of the family, to understand better the evolution of this group within Laurales. Comparisons with other Laurales and Magnoliales (a sister group of Laurales) show that Gomortega has many embryological features in common with the other lauralean families. Notably, Gomortega shares a testa without or with at best only a poorly developed mesotesta as a synapomorphy with all other Laurales. The genus further shares anthers dehisced by valves as a synapomorphy with the other Laurales (except for Calycanthaceae and Monimiaceae), and a non-multiplicative testa and bisporangiate anther as synapomorphies with Atherospermataceae and Siparunaceae (although the non-multiplicative testa occurs as a homoplasy in Monimiaceae, and the bisporangiate anther in Monimiaceae pro parte, Lauraceae pro parte and Hernandiaceae, respectively). Gomortega shows simultaneous cytokinesis to form pollen grains, a one-celled ovule archesporium and non-specialized chalaza, all or part of which may be synapomorphies shared with Atherospermataceae. Gomortega appears to have no embryological autapomorphies, but further comparison with Atherospermataceae is required.Kweon Heo and Yukitoshi Kimoto contributed equally to this work.     

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.     

Embryology of Megacodon stylophorus and Veratrilla baillonii (Gentianaceae): descriptions and systematic implications

The embryological characters of Megacodon stylophorus (C. B. Clark) H. Smith and Veratrilla baillonii Franch. are described for the first time and the systematic relationships of Megacodon and Veratrilla are discussed. Both species share the following embryological features. The anthers are tetrasporangiate. The formation of anther walls is of the Dicotyledonous Type. The tapetum is of glandular type with uninucleate cells which have a dual origin. The tapetal cells on the connective side show radial elongation or periclinal division and intrude into the anther locule to form 'placentoids'. Cytokinesis in the microsporocyte meiosis is of the simultaneous type and the microspore tetrads are nearly always tetrahedral, rarely decussate. The ovary is bicarpellate and unilocular. The ovule is unitegmic and tenuinucellar. The formation of the embryo sac is of the Polygonum Type. Before fertilization, the two polar nuclei fuse into a secondary nucleus. Fertilization is porogamous. The development of the endosperm is of the Nuclear Type. However, the two species show variation in the following features: the number of cell layers which form the anther locule wall; construction of the wall of the mature anther; cell number in mature pollen grains; ovule number in cross sections of placentae; degree of ovule curvature; character of the hypostase and seed shape. In a comparison with the other taxa in the tribe Gentianeae using embryological features, Megacodon is referred to as an independent genus and should be treated as a member of the subtribe Swertiinae; Veratrilla is better separated from Swertia s . l . as an independent genus. Veratrilla is more derived than Swertia s . s. and shows a close relationship with S. tetraptera . © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 317–331.     

中国特有属藤山柳属(猕猴桃科)的花药和胚珠发育及其分类学意义

首次报道了刚毛藤山柳（Clematoclethra scandens subsp.scandens）的花药和早期胚珠发育的胚胎学特征：花药4室;发育完整的花药壁6层,为基本型花药壁,由外至内分别为表皮、药室内壁、两层中层和两层绒毡层;腺质型绒毡层为2核或3核,纤维性加厚发生于药室内壁;小孢子母细胞减数分裂为同时型胞质分裂,小孢子四分体多为四面体型;成熟花粉粒为2-细胞型,3孔沟或稀4孔沟。早期胚珠为近倒生,单珠被,薄珠心,珠孔由单珠被构成。比较胚胎学研究认为藤山柳属和猕猴桃科的其它两个属都具有单珠被胚珠及其他相似的花药发育特征,这些特征和形态分类系统范畴的第伦桃科和山茶科有明显区别。因此,本文支持花粉形态学、细胞学和分子系统学的观点,认为藤山柳属与水东哥属和猕猴桃属构成一个较好的单系,一并放在猕猴桃科。     

Embryology of Koeberlinia (Koeberliniaceae): Evidence for core-Brassicalean affinities

Koeberlinia, comprising a single xerophytic species K. spinosa, had previously been placed in various families, mainly Capparaceae. Current molecular evidence now places it in its own family Koeberliniaceae, thought to be related to the Bataceae/Salvadoraceae among the 17 other families of the Brassicales. We investigated 55 embryological characters of the genus, most of which are not understood yet, and thereby assessed its systematic relationships. Koeberlinia has many embryological features in common with the Capparaceae and seven other core-Brassicalean families (i.e., Brassicaceae, Cleomaceae, Emblingiaceae, Gyrostemonaceae, Pentadiplandraceae, Resedaceae, and Tovariaceae), specifically by possessing a campylotropous ovule with a nonmultiplicative (two-cell-layered) outer integument, reniform seeds with a curved embryo, and a fibrous exotegmen in the mature seed coat. However, Koeberlinia is clearly distinguished from them by a tenuinucellate rather than crassinucellate ovule as previously reported, markedly enlarged apical nucellar epidermal cells, and an "exotestal" seed coat. Embryologically, Koeberlinia resembles neither the Bataceae nor the Salvadoraceae, although only limited embryological data are available for these two families. Embryological evidence thus favors its joining the core Brassicales, but additional molecular analyses and embryological studies on the missing data of the Bataceae and Salvadoraceae are needed for final confirmation of its phylogenetic position.     

Embryology of Swertia (Gentianaceae) relative to taxonomy

The embryological features of three species of Swertia ( s.l.) – S. erythrosticta , S. franchetiana , and S. tetraptera – were characterized, and the observations were used, together with previously gathered data on other species, to evaluate a recently proposed polyphyly, based on molecular data, of Swertia s.l. Comparisons of species within the genus showed that they have diversified embryologically, and there are significant between-species differences. Notable features that vary between species include the number of cell layers that form the anther locule wall, the construction of the wall of the mature anther, tapetum origin, the cell number in mature pollen grains, the structure of the fused margins of the two carpels, the ovule numbers in placental cross-sections, the shape of the mature embryo sac, the degree of ovule curvature, antipodal variation and the presence of a hypostase, and seed appendages. They share characters that are widely distributed in the tribe Gentianeae, such as a dicotyledonous type of anther wall formation, a glandular tapetum with uninucleate cells, simultaneous cytokinesis following the meiosis of the microsporocytes, tetrahedral microspore tetrads, superior, bicarpellary and unilocular ovaries, unitegmic and tenuinucellar ovules, Polygonum -type megagametophytes, progamous fertilization, nuclear endosperm, and Solanad-type embryogeny. The presence of variation in embryological characters amongst the species of Swertia s.l. strongly supports the view that Swertia s.l. is not a monophyletic group. Frasera is better separated from Swertia s.l. as an independent genus, and is only distantly related to Swertia s.s. judging from the numerous differences in embryology. Swertia tetraptera is very closely related to Halenia , as they show identical embryology.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 383–400.     

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.     

The diversity of stamen structures and dehiscence patterns among Magnoliidae

ENDRESS, P. K. & HUFFORD, L. D., 1989. The diversity of stamen structures and dehiscence patterns among Magnoliidae . Structure of stamens, particularly the patterns of anther dehiscence were studied over a wide range of families of the Magnoliidae with emphasis on the Magnoliales and Laurales as the most conservative orders of the angiosperms. Valvate dehiscence by proximal and distal stomial bifurcation was found (in addition to the already known Sarcandra and Polyalthia) for the first time in Degeneriaceae, Himantandraceae, Eupomatiaceae, in some additional Annonaceae, and in Peumus of the Monimioideae sensu lata. At least proximal bifurcations of the stomia occur in some Magnoliaceae and Ranunculaceae. An endothecial-like connective hypodermis was found (in addition to the already known Chloranthaceae and Magnoliaceae) in some Annonaceae, in Pseudowintera (Winteraceae), and in Thalictrum (Ranunculaceae). In the Annonaceae an endothecial-like connective hypodermis is partly correlated with valvate dehiscence by stomial bifurcations (as in many Hamamelididae). However, in many Magnoliidae stamens with this valvate pattern the anther is massive, especially in ‘laminar’ stamens, and the counterforce to the opening valves is therefore provided on the morphological and not on the histological level. Concomitant with valvate dehiscence by circular or elliptic flaps in the Laurales is often structural and functional dissocation of the two pollen sacs of a thcca, which is expressed by: (1) independent opening of each pollen sac, (2) lack of disruption of the interlocular zone of a theca, (3) frequent occurrence of asymmetry of the two pollen sacs of the theca, (4) frequent loss of one pollen sac per theca. In Berberidaceae with similar flaps asymmetry of the two pollen sacs of a theca is also common. These finds, together with the detection by paleobotanists of valvate anthers from the Lower Cretaceous, point to the probability that valvate anthers were more common in primitive angiosperms than previously thought.     

The structure of phenotypic variation in gender and floral traits within and among populations of Spergularia marina (Caryophyllaceae)

We sampled four wild populations of the highly autogamous Spergularia marina (Caryophyllaceae) in California to detect and to measure the magnitude of within- and among-population sources of phenotypic variation in gender and floral traits. From flowers and fruits collected from field and greenhouse-raised plants, we measured ovule number, seed number, mean seed mass, pollen production (greenhouse families only), mean pollen grain volume (greenhouse families only), anther number, anther/ovule ratio, pollen/ovule ratio (estimated using different flowers for pollen than for ovules; greenhouse families only), petal number, and petal size. Using greenhouse-raised genotypes, variation among maternal families nested within populations was evaluated for each trait to determine whether populations differ in the degree of maternally transmitted phenotypic variation. For each population, we used 15 greenhouse-raised maternal families to estimate the broad-sense heritability and genetic coefficient of variation of each floral trait. The magnitude and statistical significance of broad-sense heritability estimates were trait- and population-specific. Each population was characterized by a different combination of floral traits that expressed significant maternally transmitted (presumably genetic) variation under greenhouse conditions. Flowers representing two populations expressed low levels of maternally transmitted variation (three or fewer of nine measured traits exhibited significant maternal family effects on phenotype), while flowers representing the other two populations exhibited significant maternal family effects on phenotype for five or more traits. Our ability to detect statistically significant differences among the four populations depended upon the conditions under which plants were grown (field vs. greenhouse) and on the floral trait observed. Field-collected flowers exhibited significant differences among population means for all traits except anther number. Flowers sampled from greenhouse-raised maternal families differed among populations for all traits except ovule number, seed number, and petal size. We detected negligible evidence that genetic correlations constrain selection on floral traits in Spergularia marina.     

Development of Ovules and Embryo sacs in Ostrya virginiana (Betulaceae) and Its Systematic Significance

The development of ovules and embryo sacs in Ostrya virginiana was studied for the first time. Most ovaries had two ovules which were anatropous, unitegmic and crassinucellate. The ovule usually possessed several archesporial ceils which divided periclinally into the upper parietal cell and the lower sporogenous cell. The sporogenous cell functioned directly as megaspore mother cell. The tetrad of megaspores was linear in arrangement, and every megaspore might be functional. One ovule often contained 2- 6 embryo sacs and the embryo sac belongs to Polygonum type. It can be concluded from the present data that all ovules among the genera of the Betulaceae are unitegrnic. There are more groups with the phenomenon of multiple embryo sacs in anemophic plants such as Betulaceae, Casuarinaceae, Graminae, Jnglandaceae, Myricaceae, Simaroubaceae, Ulmaceae, than in entomophilous plants. Multiple embryo sacs also occur among some parasitic plants and saprophytes, e.g. Orobanchaceae, Cassytha in Lauraceae, Cuscuta in Convolvulaceae and Utricularia in Lentibulariaceae. It may be inferred that the characteristic of multiple embryo sacs be an evolutionary adaptation of those plants with lower pollination rate to increase the rate of fertilization. Finally, a comparison of embryological characters among the genera of the Betulaceae shows that the family is of a number of common embryological characters, such as multicellular archesporium, multiple embryo sacs in one ovule, and a long interval between pollination and fertilization. The diversity and systematic significance of several embryological characters among the “higher” hamamelid families are also discussed. It is still hard to explain the phy-logenetic relationships among those families clearly only with.     

Sporogenesis and gametogenesis in Sladenia and their systematic implication

Sporogenesis and gametogenesis in the monotypic genus Sladenia is reported for the first time. The anther wall is of the monocotyledonous type of formation and consists of four-layers, namely a glandular tapetum, one middle layer, an endothecium, and an epidermis. Fibrous thickenings are developed in the epidermis at or near the stage of anther maturity rather than in the endothecium. Successive cytokinesis in microsporogenesis results in tetrahedral or rarely isobilateral tetrads of microspores. Pollen grains are two-celled when shed. The ovule is bitegmic, tenuinucellate, and anatropous. The nucellus degenerates during meiosis of the megaspore mother cell. The micropyle is formed by only the inner integument. Development of the female gametophyte conforms to the Adoxa type. Sladenia has been treated either as a member of Actinidiaceae, Dilleniaceae or Theaceae s.l. , or as belonging to an independent family, Sladeniaceae. Recent molecular studies have suggested that it belongs to Ternstroemiaceae within the Ericales s.l . Several noteworthy embryological characters are distinct from those of the other putatively related families and support the recognition of a separate family, Sladeniaceae. Additional embryological evidence confirms this conclusion.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 305–314.     

峨眉双蝴蝶的胚胎学研究

首次报道了峨眉双蝴蝶Ｔｒｉｐｔｅｒｏｓｐｅｒｍｕｍ ｃｏｒｄａｔｕｍ（Ｍａｒｑ．）Ｈ．Ｓｍｉｔｈ的胚胎学特征,研究结果用以讨论双蝴蝶属的系统演化关系。主要研究结果如下：花药四室;药壁发育为双子叶型;绒毡层属单型起源,细胞具单核,属腺质型绒毡层,药隔处的绒毡层细胞形成类胎座,其余部位的绒毡层细胞仍为一层细胞;在花药成熟时,花药的药室内壁纤维状如厚且柱状伸长,表皮细胞减缩退化,纤维状加厚不明显。成熟花     

条纹龙胆的胚胎学研究

首次报道了条纹龙胆（Ｇｅｎｔｉａｎａ ｔｓｒｉａｔａ Ｍａｘｉｍ．）的胚胎学特征,研究结果用以讨论龙胆属狭蕊组（Ｇｅｎｔｉａｎａ Ｓｅｃｔ．Ｓｔｅｎｏｇｙｎｅ）的系统演化关系。主要研究结果如下：花药四室;药壁发育为双子叶型;绒毡层细胞仅来源于初生壁细胞,故绒毡层起源属单型起源,细胞单核,原位退化,属腺质型绒毡层,药隔处的绒毡层细胞经多次平周分裂形成２层至多层的绒毡层细胞,其余部位的绒毡层细胞仍为１     

Embryology of <Emphasis Type="Italic">Japonolirion</Emphasis> (Petrosaviaceae,Petrosaviales): a comparison with other monocots

Japonolirion osense, the sole species of the genus, endemic to Japan, which is placed together with Petrosavia in the Petrosaviaceae and the order Petrosaviales, is still poorly known with respect to systematic characters. Here I present an embryological study of the anther, ovule, and seed of J. osense. Japonolirion is characterized by a glandular anther tapetum, simultaneous cytokinesis in the microspore mother cell, two-celled mature pollen grains, anatropous and crassinucellate ovules, a two-cell-layered nucellar cap formed early in ovule development, antipodal cells hypertrophied in post-fertilization stages, the ab initio cellular mode of endosperm formation, and exotegmic seeds. Comparisons with the basal monocots Acorus (Acorales) and Araceae (Alismatales), and with the more derived monocots Nartheciaceae (Dioscoreales) and Velloziaceae/Triuridaceae (Pandanales), showed that Japonolirion is clearly distinct from those basal and more derived monocots, supporting a distinct position for Petrosaviaceae or Petrosaviales within the monocots. Extensive comparisons further suggest that the two-cell-layered nucellar cap, whose cells are rich in cytoplasm at the time of fertilization in Japonolirion and thus obviously function as the obturator, is likely to be a common characteristic of the basal monocots and may even be a link with the magnoliids.     

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.     

Embryology of Hortonioideae and Monimioideae (Monimiaceae,Laurales): characteristics of the ‘lower’ monimioids

We investigated the embryology of the ‘lower’ monimioids, i.e. Monimioideae (Monimia, Palmeria and Peumus) and Hortonioideae (Hortonia), which are poorly described embryologically. Our results show that, contrary to what has been reported in the literature, ‘lower’ monimioids show very little variation in their embryological characters. Comparisons with Mollinedioideae (a large derived subfamily in Monimiaceae) and other families in Laurales show that the ‘lower’ monimioids are relatively consistent in sharing predominantly isobilateral tetrads of microspores and megaspores, a non‐specialized chalaza, and a mesotestal–endotestal seed coat (with tracheoidal cells of the meso‐ and endotesta). It is likely that, while the shared successive cytokinesis during meiosis of microspore mother cells supports the Monimiaceae–Hernandiaceae–Lauraceae clade obtained by molecular evidence, no synapomorphies exist to support a sister‐group relationship of Monimiaceae with Hernandiaceae or Lauraceae. Instead, the lack of hypostase in ovules and/or young seeds, the lack of endosperm in mature seeds and the amoeboid tapetum in the anther are likely synapomorphies of Hernandiaceae and Lauraceae. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 158 , 228–241.     

Anther wall development,placentation, sporogenesis and gametogenesis in Smilax davidiana A.DC: A contribution to the embryology of Smilax

Embryological characters can be used to address taxonomic relationships and complement molecular phylogenetics and are of special value at the genus level. However, embryological information is fragmentary in Smilax and completely unknown in Smilax davidiana, a Chinese species. Anther wall development, placentation, sporogenesis and gametogenesis of S. davidiana are characterized here. The anther is bisporangiate, anther wall formation is of the Dicotyledonous type, both epidermis and endothecium develop fibrous thickenings, and the tapetum is secretory and of dual origin. Cytokinesis in the microsporocyte meiosis is successive, the microspore tetrad is tetragonal, and mature pollen is two-celled. The ovary is mostly trilocular with an axile placentation (a small fraction of the ovaries are unilocular with parietal placentation), the ovule is anatropous, bitegmic and crassinucellate, with embryo sac development of the Polygonum type. This study documents for the first time the embryological characters of S. davidiana in detail and contributes much to the embryology of Smilax.     

ENDOTHECIAL THICKENINGS IN ANTHERS OF PORATE MONOCOTYLEDONS

A survey of the presence of endothecial thickenings in 20 genera from 11 families of porate monocotyledons tested the long-standing hypothesis that porate anther dehiscence is correlated with the absence of thickenings. Although this hypothesis is widely repeated in texts, it has been supported by a limited, biased data base from studies of Ericaceae and Melastomataceae, two large families of dicotyledons. The results of our broad study of monocotyledons show that endothecial thickenings are present in most porate monocotyledons. Thickenings are absent in only two of the 11 porate families surveyed, Mayacaceae and Commelinaceae, both in the Commelinales. Thickenings are also absent in Xyris (Xyridaceae), a nonporate genus. These findings indicate that in monocotyledons porate anther dehiscence and the absence of thickenings are not as closely related as previously supposed.