LOSS OF FLORAL ORGANS IN ATELEIA (LEGUMINOSAE: PAPILIONOIDEAE: SOPHOREAE)

Ateleia herbert-smithii is unique among legumes in being a wind-pollinated tree; carpellate and staminate flowers are restricted to different trees. Development of the two floral morphs, however, is essentially the same except for smaller carpels in functionally staminate flowers and failure of pollen formation in the anthers of functionally carpellate flowers. The floral development of Ateleia herbert-smithii is highly atypical among papilionoids and the tribe Sophoreae. Order of organ initiation is: sepals, solitary petal, carpel, and lastly all stamens in erratic order. Sepal order is unidirectional from the abaxial side, the normal pattern for papilionoids. Only one petal, the vexillum or standard, is initiated. Subsequent initiation is completely different from the usual unidirectional pattern of most papilionoids. A meristem ring forms, delimiting the solitary carpel centrally. Ten stamen primordia are initiated on the meristem ring, first laterally, then adaxially, and lastly abaxially. There is a tendency for antesepalous stamens to form before the antepetalous ones. The loss of four of the five petals is thought to alter drastically the subsequent organogeny as to position of organs and their order of initiation. Carpel initiation in Ateleia is precocious, but not uniquely so among legumes.     

FLORAL DEVELOPMENT IN CAESALPINIA (LEGUMINOSAE)

Utilizing scanning electron microscopy, we studied the early floral ontogeny of three species of Caesalpinia (Leguminosae: Caesalpinioideae): C. cassioides, C. pulcherrima, and C. vesicaria. Interspecific differences among the three are minor at early and middle stages of floral development. Members of the calyx, corolla, first stamen whorl, and second stamen whorl appear in acropetal order, except that the carpel is present before appearance of the last three inner stamens. Sepals are formed in generally unidirectional succession, beginning with one on the abaxial side next to the subtending bracts, followed by the two lateral sepals and adaxial sepal, then lastly the other adaxial sepal. In one flower of C. vesicaria, sepals were helically initiated. In the calyx, the first-initiated sepal maintains a size advantage over the other four sepals and eventually becomes cucullate, enveloping the remaining parts of the flower. The cucullate abaxial sepal is found in the majority of species of the genus Caesalpinia. Petals, outer stamens, and inner stamens are formed unidirectionally in each whorl from the abaxial to the adaxial sides of the flower. Abaxial stamens are present before the last petals are visible as mounds on the adaxial side, so that the floral apex is engaged in initiation of different categories of floral organs at the same time.     

红车轴草提取物中总鹰嘴豆芽素类成分HPLC测定方法的研究

对红车轴草提取物中总鹰嘴豆芽素类成分测定方法进行了研究，建立了以鹰嘴豆芽素类Ononin,Sissotrin,Formononetin及Biochanin A为个标的HPLC测定总鹰嘴豆芽素类的方法和仅以Biochanin A为外标HPLC测定总鹰嘴豆芽素类的方法，两种方法所得结果基本一致。     

SEEDLING RECRUITMENT LIMITATION IN WHITE CLOVER (TRIFOLIUM REPENS; LEGUMINOSAE)

White clover (Trifolium repens L.) is a stoloniferous clonal legume. Dynamics of white clover populations are controlled by the recruitment of both seedlings (genets) and node production (ramets). To understand the role of seedling dynamics in natural populations of white clover, the recruitment and mortality of seedlings in a lawn and pasture clover population were followed and the roles of biotic and abiotic factors in seedling recruitment and survival were assessed. Seedling recruitment was several orders of magnitude less than vegetative recruitment in both lawn and pasture populations. However, at least several new genets are added to these populations on a yearly basis. In both locations, physical disturbances may open sites for seed germination, as disturbed sites had higher seed germination rates than undisturbed sites. However, these sites were not favorable for long-term establishment due to such factors as frost upheaval. Therefore, there are trade-offs in safe site attributes for seed germination and establishment that strongly limit recruitment in natural populations of white clover.     

POLLEN AND TAPETUM DEVELOPMENT IN DESMODIUM GLUTINOSUM AND D. ILLINOENSE (PAPILIONOIDEAE; LEGUMINOSAE)

Each of the four microsporangia has three or four wall layers, a uninucleate tapetum of various cell shapes with nuclei that remain in prophase, and 12-24 pollen mother cells (PMCs). A sterile transverse septum sometimes bisects the microsporangium. PMCs secrete callose but not uniformly, and contact among them continues through meiosis. Simultaneous cytokinesis by furrowing isolates each microspore in callose, which later disperses. The separated microspores become vacuolate, undergo mitosis to become pollen, and later become filled with food reserves. Endothecial wall thickening and tapetal dissolution occur after pollen engorgement. Calcium oxalate crystals form in tapetal cells during the sporogenous stage, reach maximum size during early meiosis, and remain prominent until tapetal dissolution.     

COMPARATIVE DEVELOPMENT OF SECRETORY CAVITIES IN THE TRIBES AMORPHEAE AND PSORALEEAE (LEGUMINOSAE: PAPILIONOIDEAE)

The tribes Amorpheae and Psoraleeae of the Leguminosae (Papilionoidae) share the characteristics of one-seeded fruits and gland-dotted foliage. Because of this, they traditionally have been considered closely related (either a single tribe or two closely related tribes). However, Barneby (1977) has suggested that the Amorpheae and Psoraleeae are not close but previously had been combined on the basis of a superficial resemblance. This paper describes the structure of the secretory cavities responsible for the gland dots. Approximately 50% of the species of each tribe were surveyed for cavity structure with leaflet clearings. Eight species were then chosen for developmental studies of their glands. Several distinct kinds of secretory cavities are present in these plants. Trabeculate cavities (found only in the Psoraleeae) are traversed by many elongated cells. This type of cavity and nontrabeculate cavities of the Psoraleeae initiate with localized dorsiventral elongation of protodermal cells to form a hemispherical protuberance on the leaf primordium surface. Development proceeds with separation of the cells of a protuberance along their lateral walls facing the protuberance center. As the leaf expands, the protuberance sinks until its apex is flush with the leaf surface. The result is a cavity lined by an epithelium of modified epidermal cells. Trabeculate cavities have more cells in the initial protodermal bump than nontrabeculate “epidermal” cavities, and the central cells of the protuberance are not involved in epithelium formation, but become separated from other cells on all lateral sides, transversing the cavity as trabeculae. Cavities of the Amorpheae are all nontrabeculate and subepidermal. They initiate with periclinal divisions of protodermal cells that result in two cell layers. The exterior layer differentiates into epidermis, while the interior layer divides to produce a small spherical group of cells (“epithelial initials”). Schizogeny occurs in the center of these cells to produce an epithelium-lined cavity. Previous studies of cavity development in the Amorpheae described lysigenous and schizo-lysigenous cavities for most species. These early reports are reviewed, and the possible role of preparation artifacts in producing images of lysigenous development in general is discussed.     

FLORAL ONTOGENY IN SOPHOREAE (LEGUMINOSAE: PAPILIONOIDEAE). I. MYROXYLON (MYROXYLON GROUP) AND CASTANOSPERMUM (ANGYLOCALYX GROUP)

Floral ontogeny is described and documented using scanning electron microscopy in Myroxylon balsamum and Castanospermum australe, representatives respectively of Polhill's Myroxylon and Angylocalyx groups (Leguminosae: Papilionoideae), groups exhibiting relatively unspecialized flower structure for the tribe Sophoreae. Both are woody tropical trees with axillary or terminal racemes or panicles. Bracteoles are present in both Myroxylon and Castanospermum. Flowers are initiated singly in bract axils, which are produced in acropetal order by the inflorescence apical meristem. The flower structure of both includes a broad calyx tube, five petals lacking any fusion (only the vexillary distinctive), ten free homogeneous stamens in two whorls, and a long-stipitate carpel. The two taxa are alike in early organogenetic stages with essentially acropetal order of initiation: sepals, petals, outer stamens plus carpel, inner stamens. Within each whorl the order is unidirectional from the abaxial side. They are alike through middevelopment with one exception. There is accelerated vexillar enlargement in Castanospermum by middevelopment, not found in Myroxylon. Both have a hypanthium, which forms late in development. In both, large flower size, exserted stamens, and hypanthium are adaptations to bird-pollination. Differences between the two that are manifested in late development include strongly zygomorphic calyx and petal color change over time (Castanospermum), stamens sagittate and apiculate with some basal filament fusion (Myroxylon), stigma form differences, and fruit form.     

TRACHEID BAR AND VESTURED PITS IN LEGUME SEEDS (LEGUMINOSAE: PAPILIONOIDEAE)

The tracheid bar, a strip composed of vertically oriented large tracheid-like cells (tracheoids), occurs only in the hilum of seeds of papilionoid legumes. An anatomical survey of the bar was made from seeds representing 232 species of 97 genera from 29 of the 31 tribes recognized by Polhill. Seeds were sectioned freehand, coated, then viewed by SEM. The tracheid bar is quite uniform in its general features throughout the subfamily, although differences in size and shape of both the bar and the tracheoids were found. Eight species from tribes considered to be among the primitive elements of the subfamily exhibited three variant forms: horizontal tracheary elements instead of the usual bar (2 species), tracheid bar with subtending but separate vascular bundle (1 species), and the tracheid bar with fused horizontal tracheary elements (5 species). Bordered pits of individual cells in the tracheid bar virtually always lacked a membrane and had smooth, warty, or variously elaborate vestures on the border. This appears to be the first report of vestured pits other than in secondary xylem. With some exceptions, bordered pits tended to be vestured in primitive tribes, warty in intermediate tribes, and smooth or only slightly warty in the most advanced tribes.     

THE INFLUENCE OF GROWTH STAGE ON THE RESPONSE OF RED CLOVER (TRIFOLIUM PRATENSE L.) TO GIBBERELLIC ACID

The response of seedling and first harvest year plants of red clover ( Trifolium pratense L.) to treatment with gibberellic acid (GA) at various growth stages is described.
Seedlings sprayed before the seventh leaf stage developed into single-stemmed plants; treatment with GA at the third- or fourth-tiller stage resulted in final stem numbers similar to those of controls. Emergence was earliest, and the number of heads per plant greatest where sprayings were delayed until the third- or fourth-tiller stage.
In first harvest year plants significant increases in the number of heads per stem were obtained with certain treatments, especially those which had two applications of 0.5 mg. GA per plant during the elongation of the first four internodes. This was related in all treatment groups to modifications of the branching pattern, and also to the increased incidence of multiple heading. Earlier emergence resulted from treatment at all growth stages, the effect being maximal where three well-spaced sprayings were applied during active stem extension.     

OVERLAPPING ORGAN INITIATION AND COMMON PRIMORDIA IN FLOWERS OF PISUM SATIVUM (LEGUMINOSAE: PAPILIONOIDEAE)

The floral ontogeny of Pisum sativum shows a vertical order of succession of sepals, petals plus carpel, antesepalous stamens, and antepetalous stamens. Within each whorl, unidirectional order is followed among the organs, beginning on the abaxial side of the flower, as in most papilionoids. Unusual features include the four common primordia which precede initiation of discrete petal and antesepalous stamen primordia, and the marked overlap of organ initiations between whorls which are usually separately initiated. The stamens arise in free condition, then become diadelphous by intercalary growth at the base of nine stamens, and finally become pseudomonadelphous by surface fusion between the vexillary stamen filament and the adjacent edges of the filament tube. The early initiation of the carpel is not unique among papilionoids, but is somewhat unusual.     

HISTOGENESIS OF LEAF VENATION IN TRIFOLIUM WORMSKIOLDII (LEGUMINOSAE)

The histogenesis of the leaf venation in Trifolium wormskioldii was followed, using whole cleared leaflets and transverse and paradermal microtome sections. Mature leaflets are about 8 mm in length. The midvein precambium appears in leaflets 100 μ in length; secondary vein procambium is first seen in 230 μ leaflets; procambium of minor veins first appears in leaflets 400-600 μ in length. Phloem is apparent before xylem, in midvein, secondary veins, and minor veins. Vein endings are initiated and they mature last. No evidence was found to support the theory of “vein breakage.” In the greenhouse 24 days elapsed from leaf initiation to exposure of leaf tips; 6 more days elapsed until full leaf expansion.     

POLLEN BRUSH OF PAPILIONOIDEAE (LEGUMINOSAE): MORPHOLOGICAL VARIATION AND SYSTEMATIC UTILITY

The pollen brush commonly is referred to as a “bearded” or “pubescent” style in taxonomic literature and traditionally is taken to be an aggregation of trichomes on the distal end of the style, and occasionally including the stigma. We present data that support the taxonomic utility of the pollen brush but define it more specifically as a dense aggregation of erect trichomes emanating from the style (not stigma or ovary) and functioning in secondary pollen presentation. We recommend avoiding such vague terminology as bearded or pubescent styles as these refer not only to the pollen brush but also to ciliate and penicillate stigmas and ciliate styles. The latter three conditions have some taxonomic use, and since their occurrence is not necessarily correlated with the presence of a pollen brush, they should be distinguished from it. We estimate that the pollen brush has arisen independently in the following eight taxa: 1) Crotalaria and Bolusia (Crotalaraieae), 2) subtribe Coluteinae (Galegeae), 3) Tephrosia subgenus Barbistyla (Millettieae), 4) Adenodolichos (Phaseoleae subtribe Cajaninae), 5) Clitoria (Phaseoleae subtribe Clitoriinae), 6) the subtribe Phaseolinae (Phaseoleae), 7) the Robinia group (Robinieae), and 8) the tribe Vicieae. Its hypothesized homology within each of these groups is supported by a cooccurrence with other taxonomic characters, both morphological and molecular.     

FLORAL DEVELOPMENT IN SAURURUS CERNUUS (SAURURACEAE): 1. FLORAL INITIATION AND STAMEN DEVELOPMENT

The inflorescence of Saururus cernuus L. produces lateral “common” primordia in acropetal succession on the flanks of the inflorescence meristem; curiously, the “subtending” bract is initiated upon the lateral primordium rather than subtending it. On the basis of mature floral structure, flowers of S. cernuus have previously been described as having spiral initiation of parts. The current ontogenetic investigation contradicts this interpretation. Stamens arise in three successive pairs; the carpels also are initiated in pairs. Floral symmetry is shown to be bilateral from the onset of organ initiation, a rare feature among primitive angiosperms. On the basis of symmetry and paired initiation of organs, the possibility of close relationships between Saururaceae and Magnolialian or Ranalian lines appears remote.     

THE GRANULAR INTERSTITIUM IN THE POLLEN OF SUBFAMILY PAPILIONOIDEAE (LEGUMINOSAE)

A wide range of transitional forms of granular interstitia from simple to complex and from random to ordered occur in the pollen of the subfamily Papilionoideae. Three main types are described: 1) large, widely spaced irregular granules (Type A); 2) densely packed groups of columellae and granules (Type B); and 3) a mass of more or less disorganized granules (Type C). In the genus Calopogonium (tribe Phaseoleae) all three types have been found in different species. Two of the types have been found in different species of the genus Psoralea (tribe Psoraleeae). Granular structures so far occur in six tribes: Desmodieae, Indigofereae, Loteae, Phaseoleae, Psoraleeae, and Vicieae. All of the tribes are regarded as being evolutionarily advanced in both macro and micro characters and many, but not all, show specialized pollen characters. It is concluded that the granular interstitium is a derived structure in papilionoid legumes.     

IDENTIFICATION OF PROCAMBIUM IN THE PRIMARY ROOT OF TRIFOLIUM PRATENSE (FABACEAE)

Histochemical and morphometric analyses were used to identify and define an early stage of procambial differentiation in 1.5–2.0-cm-long (48 hr after germination) primary roots of Trifolium pratense L. Esterase activity was used as a histochemical marker for early differentiation of procambium. Morphometric analysis of cell length and width as a function of distance from the root cap junction was performed on the same tissue using brightfield and Nomarski DIC optics. This combination of techniques allowed the identification of esterase activity in both the cell wall and cytoplasm and permitted the determination of the exact location, size, and shape of the histochemically stained cells within the apex. Esterase activity identified the proendodermis and procambial cylinder (six to seven cells in diameter) two to three cells proximal to the root cap junction. In this system, esterase activity proved to be an earlier marker for procambial differentiation than morphometric or cytological changes. It is suggested that these techniques will be useful in characterizing procambial pattern development in more complex shoot systems.     

FLORAL DEVELOPMENT AND VASCULATURE IN HYDROCLEIS NYMPHOIDES (BUTOMACEAE)

The flower of Hydrocleis nymphoides consists of three sepals which arise in spiral succession, three simultaneously arising petals, numerous stamens and staminodia which arise in centrifugal order, and six carpels. A residual apex remains at maturity. The first-formed members of the androecium are stamens and the later-formed members are staminodia which develop below the stamens and which become outwardly displaced during expansion of the receptacle. The androecium is supplied by branching vascular trunk bundles. The carpels are completely open but the ventral margins are slightly conduplicately appressed basally. A single dorsal bundle provides the stigmatic area with vascular tissue, and a network of small placental bundles supplies the numerous laminar ovules. There are no clearly defined ventral bundles. It is suggested that Hydrocleis nymphoides is neither the most primitive nor the most advanced member of the family. A pattern of phylogenetic reduction in the androecium and receptacle is suggested for the entire family.     

HYBRIDIZATION IN THE GENUS GLYCINE SUBGENUS GLYCINE WILLD. (LEGUMINOSAE,PAPILIONOIDEAE)

The genus Glycine is composed of two subgenera, Glycine and Soja. Soja includes the cultivated soybean, G. max, and its wild annual counterpart G. soja, while Glycine includes seven wild perennial species. Hybridization was carried out within and between wild perennial species of the subgenus Glycine. The success rate (pods set/flowers crossed) was 11% for intraspecific and 8% for interspecific crosses. A total of 220 F₁ hybrids was examined morphologically and cytologically where possible. Hybrids within G. canescens (2n = 40) and G. latifolia (2n = 40) were fertile as expected. Glycine clandestina (2n = 40) was morphologically separable into at least three groups, which produced fertile hybrids within each group. One cross between two groups gave vegetatively vigorous but sterile hybrids. The majority of crosses within G. tabacina (2n = 80) were fertile, except that extremely narrow-leaved forms gave sterile hybrids in combination with more usual forms. Sterility was also encountered in G. tomentella when aneuploids (2n = 78) from New South Wales, Australia, were crossed with tetraploids (2n = 80) from either Queensland, Australia, or Taiwan; crosses between the latter two populations resulted in seedling lethality. Cytological behavior of sterile hybrids followed a similar pattern, whether at the diploid or tetraploid level. The frequency of chromosome pairing was approximately half that expected if genomes showed full pairing homology. Bivalent disjunction at anaphase I was usually followed by precocious division of the majority of univalents. Telophase I and II were characterized by lagging chromosomes and micronuclei, so that resulting pollen was misshapen and sterile. Chromosome pairing data from sterile intraspecific hybrids at the tetraploid level may indicate a polyphyletic origin of tetraploids, whereby different diploid populations were involved in their formation. Similarly, chromosome pairing in sterile intraspecific diploid hybrids may indicate that the various diploid groups arose independently of one another. Both 40- and 80-chromosome forms are fully diploidized, however, and if they are of ancient origin, divergence since that time could have resulted in the chromosomal differentiation which becomes apparent when intraspecific hybridization is effected. Diploid (2n = 40) interspecific hybrids G. falcata × G. canescens, and G. falcata × G. tomentella grew poorly and did not reach flowering stage. Diploid (2n = 40) crosses between G. latifolia and G. tomentella produced inviable seedlings. Tetraploid (2n = 80) hybrids between G. tomentella and G. tabacina were vegetatively vigorous but sterile owing to low chromosome pairing at meiosis, indicating little pairing homology between the two species. Diploid hybrids between G. canescens and G. clandestina, however, showed almost complete chromosome pairing at diakinesis and partial fertility. Although morphologically distinct, these two species have not diverged sufficiently to prevent hybridization and possible gene exchange through recombination. Self compatibility, perennial growth habit, and geographic isolation have favored divergence among Glycine populations to the point that gene exchange appears no longer possible in many cases. Internal isolating mechanisms have been shown to operate at various levels of plant development from hybrid lethality at seedling stage, to failure of seed-set in sterile but vegetatively vigorous hybrids.