Heterosis in hybridization between the monoecious and dioecious subspecies of Ecballium elaterium (L.) Rich

Summary Statistical analysis was performed on a set of samples cultivated in balanced, incomplete blocks corresponding, respectively, to filial generations (pure, hybrid or backcross) distinguishable in the hybridization between the monoecious and dioecious subspecies of Ecballium elaterium (L.) Rich. We thus analyzed the possible existence of polygenes that condition the size of the plants, in particular the most observable characteristic effect of the polygenes, i.e., heterosis or hybrid vigor. The experimental results show that heterosis exists with respect to size of the individual plant. The occurrence of this phenomenon provides experimental evidence for the existence of a system of polygenes that affect this property. With respect to each of these polygenes, the genotype of each subspecies contains different alleles. This system of polygenes very likely plays a role in the phenotypic modification which, regarding the sex of the plant itself, has been recognized and confirmed previously.     

Genetic variation in Ecballium elaterium (Cucurbitaceae): Breeding system and geographic distribution

Dioecy, the separation of sexes, has arisen independently many times in the course of angiosperm evolution. Avoidance of inbreeding is clearly involved in the evolution of dioecy, and as a consequence we predict that dioecious populations should maintain higher levels of genetic variation than closely related nondioecious populations. We tested that prediction by comparing allozymic variation in two closely related taxa, the monoecious and dioecious subspecies of the Mediterranean cucurbit, Ecballium elaterium. Thirteen polymorphic loci were screened for seeds sampled from 10 monoecious and 13 dioecious populations spanning the geographic ranges of the subspecies in Spain. The dioecious subspecies showed strikingly greater allelic diversity and heterozygosity than the monoecious subspecies. A hierarchical F-statistic analysis clearly demonstrated considerable genetic variation within populations for the dioecious populations, whereas for the monoecious populations almost all genetic variation resulted from differences among populations. The general pattern of homozygosity within monoecious populations suggests that they are highly inbred. In order to assess historical influences on current patterns of genetic variation, we conducted a genetic-distance analysis. The observed relationship between genetic distance and geographic distance between populations supports the hypothesis that the subspecies' current allopatric distributions on the Iberian Peninsula are the result of separate waves of colonization from the north (monoecious) and south (dioecious).     

System of pleiotropic polygenes in the Cucurbitacea Ecballium elaterium (L.) Rich. related to durability and size of plant

Summary The statistical association between the characters life durability of plant and size of plant observed in a set of samples — cultivated forming a system of randomly balanced incomplete blocks — and corresponding to filial generations (pure, hybrid or backcross) from hybridization between the monoecious subspecies and the dioecious subspecies of the Cucurbitacea Ecballium elaterium (L.) Rich, respectively, is in concordance with the hypothesis that the system of polygenes governing the life durability of the plant in this species and the system of polygenes governing the size of the plants belong to the same system of polygenes. This system of pleiotropic polygenes governs the size and life durability of the plants at the same time.     

Impacts of floral gender and whole-plant gender on floral evolution in Ecballium elaterium (Cucurbitaceae)

Investigation of gender specialization in plants has led to several theories on the evolution of sexual dimorphism: reproductive compensation, based on enhanced reproductive efficiency with gender specialization (flowers should be larger on dioecious plants); Bateman's Principle, based on sex-specific selection (display for pollinator attraction in males and seed set in females); and intersexual floral mimicry, based on mimicry of a reward-providing gender by a non-reward providing gender (reduced dimorphism in dioecious plants due to increased spatial separation of male and female flowers). These theories were evaluated in Ecballium elaterium, which contains two subspecies, elaterium (monoecious) and dioicum (dioecious). Our results show that flowers of the dioecious subspecies are larger and allocate more to reproductive organs than do flowers of the monoecious subspecies. Both subspecies are sexually dimorphic (male flowers larger than female flowers). Variance in flower size among populations is greater in the dioecious subspecies. Finally, there is sufficient genetic variation to enable ongoing response to selection; genetic correlation constraints on independent response of female and male flowers may be stronger in the monoecious subspecies. Our findings provide support for aspects of all three theories, suggesting that the evolution of floral dimorphism is based on a complex interplay of factors.     

Foraging behavior of three bee species in a natural mimicry system: female flowers which mimic male flowers in Ecballium elaterium (Cucurbitaceae)

Summary The behavior of Apis mellifera and two species of solitary bees which forage in the flowers of monoecious Ecballium elaterium (L.) A. Rich (Cucurbitaceae) were compared. The female flowers of E. elaterium resemble male flowers visually but are nectarless, and their number is relatively smaller. Apis mellifera was found to discriminate between the two genders and to pay relatively fewer visits to female flowers (mean of 30% relative to male flowers) from the beginning of their activity in the morning. The time spent by honeybees in female flowers is very short compared to that spent in male flowers. It is surmised that the bees remember the differences between the flowers where they foraged on the previous days. In contrast, the two species of solitary bees Lasioglossum politum (Morawitz) (Halictidae) and Ceratina mandibularis Fiese (Anthophoridae) visit the female flowers with nearly equal frequencies at the beginning of each foraging day and stay longer in these flowers. Over the day there is a decline in the relative frequency of visits to female flowers and also in the mean time spent in them. The study shows that bees can collect rewards at high efficiency from the flowers of Ecballium elaterium because of their partial discrimination ability and the scarcity of the mimic flowers. It is suggested that the memory pattern of some solitary bees may be different from that of Apis mellifera. It seems that the limited memory and discrimination ability of bees can lead to a high frequency of visits to the mimic flowers during a long flowering season.     

Nectary structure and nectar secretion in Maxillaria coccinea (Jacq.) L.O. Williams ex Hodge (Orchidaceae)

BACKGROUND AND AIMS: It had previously been assumed that Maxillaria spp. produce no nectar. However, nectar has recently been observed in Maxillaria coccinea (Jacq.) L.O. Williams ex Hodge amongst other species. Furthermore, it is speculated that M. coccinea may be pollinated by hummingbirds. The aim of this paper is to investigate these claims further. METHODS: Light microscopy, histochemistry, scanning and transmission electron microscopy. KEY RESULTS: This is the first detailed account of nectar secretion in Maxillaria Ruiz & Pav. A 'faucet and sink' arrangement occurs in M. coccinea. Here, the nectary is represented by a small protuberance upon the ventral surface of the column and nectar collects in a semi-saccate reservoir formed by the fusion of the labellum and the base of the column-foot. The nectary comprises a single-layered epidermis and three or four layers of small subepidermal cells. Beneath these occur several layers of larger parenchyma cells. Epidermal cells lack ectodesmata and have a thin, permeable, reticulate cuticle with associated swellings that coincide with the middle lamella between adjoining epidermal cells. Nectar is thought to pass both along the apoplast and symplast and eventually through the stretched and distended cuticle. The secretory cells are collenchymatous, nucleated and have numerous pits with plasmodesmata, mitochondria, rough ER and plastids with many plastoglobuli but few lamellae. Subsecretory cells have fewer plastids than secretory cells. Nectary cells also contain large intravacuolar protein bodies. The floral morphology of M. coccinea is considered in relation to ornithophily and its nectary compared with a similar protuberance found in the entomophilous species M. parviflora (Poepp. & Endl.) Garay. CONCLUSIONS: Flowers of M. coccinea produce copious amounts of nectar and, despite the absence of field data, their morphology and the exact configuration of their parts argue strongly in favour of ornithophily.     

Leaf structure and histochemistry of Ecballium elaterium (L.) A. Rich. (squirting cucumber)

Light, scanning and transmission electron microscopy as well as histochemical reactions were employed to study the leaf structure and secretory activity of Ecballium elaterium, a hairy pharmaceutical perennial common in the Mediterranean region. The amphistomatic leaf has a peculiar structure due to special cells supporting the conductive bundles, a remarkable shortage of mechanical tissue, and the existence of pectin strands between mesophyll cells. The secreting activity is limited mostly to secretary hairs. These attributes of the Ecballium leaf fine structures do not resemble the common structure of leaves from Mediterranean plants and point to a peculiar strategy of this species coping with stress conditions of its habitat.     

Nectary and gender‐biased nectar production in dichogamous Chamaenerion angustifolium (L.) Scop. (Onagraceae)

In dichogamous plants, nectar characteristics (i.e. nectar amount and its composition) can differ between sexual phases. In the present study, we investigated the structural organization of the floral nectary, nectar production and carbohydrate composition in the protandrous Chamaenerion angustifolium (L.) Scop. (Onagraceae). The receptacular nectary consisted of an epidermis with numerous nectarostomata, several layers of photosynthetic secretory parenchyma, and subsecretory parenchyma. Nectariferous tissue was not directly vascularized and starch grains were rarely observed in the secretory cells, occurring exclusively in the guard cells of modified stomata. The nectar was released via nectarostomata. The floral nectar was hexose rich (32.8/39.1/28.1% glucose/fructose/sucrose) and the total concentration was constant throughout the anthesis (47% on average). However, contrasting patterns in nectar amount and carbohydrate composition between the floral sexual phases were observed. On average, female‐phased flowers produced 1.4‐fold more nectar than male‐phased flowers, and although the nectar was sucrose rich during the male phase, it was hexose rich during the female phase, suggesting sucrose hydrolysis.     

Environmental Effects on Components of Pollen Performance in Faramea occidentalis (L.) A. Rich. (Rubiaceae)1

Studies focusing on gametophytic competition have focused on differences in pollen performance (e.g., pollen germination rate and pollen tube growth rate) among donors in order to examine genetic sources of variation in siring success among them. Donors that produce the fastest growing pollen tubes are expected to fertilize more ovules relative to donors with slow growing pollen tubes under conditions of gametophytic competition. However, the performance of pollen in the field is known to be influenced by environmental conditions in addition to the genotype of donor plants. This field study of Faramea occidentalis was conducted to: (1) determine the effect of environmental conditions during pollination on pollen performance; (2) measure differences among pollen donors in pollen performance; (3) determine if the pollen of different donors responds differently to a variety of environmental conditions surrounding pollinated flowers; and (4) measure differences among pollen recipients in pollen performance. Single-donor crosses were made between four pollen donors and four pollen recipients under a variety of environmental conditions. Pollen performance was then quantified as the growth rate of the fastest pollen tube, the mean pollen tube growth rate, and by a pollen germination index. Pollination environment (the environment surrounding a pollinated flower) and recipient significantly affected all three measures of pollen performance. Pollen donors did not differ overall in pollen performance. However, there was significant among-donor variation in two of the five pollination environment conditions. Future studies of variation in relative siring success may benefit by considering pollination environment in addition to donor and recipient identity.     

A new species of Sicyos L. (Cucurbitaceae) from Oaxaca, Mexico

Sicyos mcvaughii, a new species from Oaxaca, is described and illustrated. It is similar to Sicyos sertuliferus Cogn., and grows in secondary tropical deciduous forest, between 20 and 630 meters.     

Ultrastructure of microsporogenesis and microgametogenesis inArabidopsis thaliana (L.) Heynh. ecotype Wassilewskija (Brassicaceae)

Summary The process of microsporogenesis and microgametogenesis was studied at the ultrastructural level in wild-typeArabidopsis thaliana ecotype Wassilewskija to provide a basis for comparison with nuclear male-sterile mutants of the same ecotype. From the earliest stage studied to mature pollen just prior to anther dehiscence, microsporocyte/microspore/pollen development follows the general pattern seen in most angiosperms. The tapetum is of the secretory type with loss of the tapetal cell walls beginning at about the time of microsporocyte meiosis. Wall loss exhibits polarity with the tapetal protoplasts becoming located at a distance from the inner tangential walls first, followed by an increase in distance from the radial walls beginning at the interior edge and progressing outward. The inner tangential and radial tapetal walls are completely degenerated by the microspore tetrad stage. Unlike other members of the Brassicaceae that have been studied, the tapetal cells ofA. thaliana Wassilewskija also lose their outer tangential walls, and secretion occurs from all sides of the cells. Exine wall precursors are secreted from the tapetal cells in a process that appears to involve dilation of individual endoplasmic reticulum cisternae that fuse with the tapetal cell membrane and release their contents into the locule. Following completion of the exine, the tapetal cell plastids develop membranebound inclusions with osmiophilic and electron-transparent regions. The plastids undergo ultrastructural changes that suggest breakdown of the inclusion membranes followed by release of their contents into the locule prior to the complete degeneration of the tapetal cells.     

Ultrastructural Aspects of the Nectary Spur of Limodorum abortivum (L) Sw. (Orchidaceae)

The ultrastructure of the nectary spur of Limodorum abortivum(L) Sw. was examined before and after anthesis. In cross sectionthe nectary spur shows an internal epidermal layer of thin-walledcells bordering the secretory cavity and 10–12 layersof parenchyma cells. The ultrastructure of the secretory cellssuggests the involvement of ER, Golgi and plastids in nectarsecretion. The nectar accumulated in the sub-cuticular spaceis released into the nectariferous cavity by rupture of theouter layer of the cuticle. Limodorum abortivum (L) Sw., Orchidaceae, nectary spur, nectar secretion, ultrastructure, anthesis, endoplasmic reticulum, dictyosomes, plastids     

Ultrastructure and histochemistry of Citrus limon (L.) stigma

Citrus limon has a wet stigma which can be divided in two zones: a glandular superficial one formed by papillae, and a non-glandular one formed by parenchymatic cells. The stigmatic exudate is produced by the papillae after the latter have reached their ultimate size. The papillae of the mature pistil are of varying size and composition. Both the unicellular and multicellular ones are present. The cells at the base of the papillae are rich in cytoplasm, whereas the tip cells are vacuolated. Histochemical analysis has shown that the exudate of Citrus is composed of lipids, polysaccharides, and proteins. Our results indicate that the lipidic component is produced and secreted first, followed by production and secretion of the polysaccharidic component. The lipidic component of the exudate is produced in the basal papillae cells and accumulates as droplets in dilated parts of the smooth endoplasmic reticulum (SER). Subsequently the lipid droplets are transported to the plasma membrane, and transferred by the latter into the cell walls. Then the exudate component is accumulated in the intercellular spaces and in the middle lamellar regions of the walls. Subsequently, the polysaccharidic component of the exudate is produced and secreted by the tip cells of the papillae.Abbreviations RER rough endoplasmic reticulum - SER smooth endoplasmic reticulum     

Factors Affecting Shedding of Flowers in Soybean (Glycine max (L.) Merrill)

Flower shedding in soybean, Glycine max (L.) Merrill, was studiedusing cultivar ‘Clark’, isoline E₁t, which has relativelylong racemes for convenient identification and observation ofindividual flowers. On each raceme studied, pod set was greatestat the proximal (basal) positions, whereas shedding was greatestat the most distal positions. When proximal flowers were removedas they reached anthesis, pod set increased at the more distalpositions. Pod set was increased in some instances by applicationof water directly to the ovaries as a drop in the calyx cup.Peroxidase activity changed in parallel with ovary development,increasing rapidly in growing pods but not in shedding flowers.Increases in flower peroxidase was mainly in ovary walls. Flowerstaken at or near anthesis from positions with high percent podset could be grown in vitro with especially good ovary enlargement,whereas ovaries in flowers taken from positions of low pod setdid not enlarge in culture. Unidentified substances were extracted from young pods which,when incorporated into lanolin and tested in an in situ bioassay,could mimic the effect of proximal flowers in inducing sheddingof distal flowers. Indole-3-acetic acid resembled the extractedmaterials in inducing shedding, but differed by eliciting side-effectsthat extracts did not. The growth substances abscisic acid,gibberellic acid, and benzyladenine did not promote sheddingin the in situ test. The evidence was taken to indicate that soybean flower sheddingis induced in distal flowers by substances from the more proximal,fertilized ovaries, and that this is possibly due to interferencewith some of the intense metabolic changes that follow pollinationand fertilization.     

Processes of isolation and introgressive interplay between Platanthera bifolia (L.) Rich and P. chlorantha (Custer) Reichb. (Orchidaceae)

NILSSON, L. A., 1983. Processes of isolation and introgressive interplay between Platanthera bifolia L. Rich, and P. chlorantha (Custer; Reichb. (Orchidaceae). Platanthera bifolia and P. chlorantha are highly specialized for pollination by crepuscular and nocturnal Lepidoptera. The plants are largely sympatric, have overlapping flowering-times, and are completely interfertile. Principal differences exist in floral fragrance and column morphology. Isolation is effected by a set of integrated morphological, ethological, and seasonal mechanisms. Odour-based constancy by moths, differences in column structure, attachment of pollinaria, spurlength, nectar availability, and flowering-peaks reduce interspecific pollen transfer. The plants form a functional species pair in relation to their pollination environment and locally share the same main pollinator species.
Hybrids, which infrequently occur in areas of sympatry, emit a blend of the species-specific fragrance compounds and have intermediate column structure. They are frequently visited by-moths but suffer a serious reduction in seed production due to imperfect interaction vis-à-vis the pollen vectors. Hybrid viscidia, in an intermediate position on the column, are touched by moths' scaly and hairy palps and get insufficient attachment, evidently causing hybrids to act solely as female recipient. First backcross individuals largely join the functional range of either species which promotes rapid introgression. The two gene-pools interplay, although always at a low level. Patterns of introgressive hybridization were exemplified.     

Effects of X-rays and Colchicine on Pollen of Trichosanthes anguina L. (Cucurbitaceae)

Pollen grains of M, and C, generation plants of Trichosanthes anguina L. were studied after treatment of seeds with X-rays and colchicine respectively. Pollen sterility increased with increase in X-ray dose and colchicine concentration. The polar axis of X-irradiated populations was shorter than that of controls. But colchicine treatment resulted in larger pollen grains than the controls, although the exine did not increase as much. Ex-perimentally-produced tetraploids had larger pollen grains than diploids and triploids. The triploids were characterised by changes in pore morphology. Of 293 treated plants examined, 11 aberrant plants were isolated in the M, and C, generations. These plants had large and small fertile pollen grains, and also showed treatment effects in external morphology and chromosomal aberration during meiosis. The changes in polar diameter were not associated with variation in chromosomal material.     

Construction of an SSR and RAD-Marker Based Molecular Linkage Map of Vigna vexillata (L.) A. Rich

Vigna vexillata (L.) A. Rich. (tuber cowpea) is an underutilized crop for consuming its tuber and mature seeds. Wild form of V. vexillata is a pan-tropical perennial herbaceous plant which has been used by local people as a food. Wild V. vexillata has also been considered as useful gene(s) source for V. unguiculata (cowpea), since it was reported to have various resistance gene(s) for insects and diseases of cowpea. To exploit the potential of V. vexillata, an SSR-based linkage map of V. vexillata was developed. A total of 874 SSR markers successfully amplified single DNA fragment in V. vexillata among 1,336 SSR markers developed from Vigna angularis (azuki bean), V. unguiculata and Phaseolus vulgaris (common bean). An F₂ population of 300 plants derived from a cross between salt resistant (V1) and susceptible (V5) accessions was used for mapping. A genetic linkage map was constructed using 82 polymorphic SSR markers loci, which could be assigned to 11 linkage groups spanning 511.5 cM in length with a mean distance of 7.2 cM between adjacent markers. To develop higher density molecular linkage map and to confirm SSR markers position in a linkage map, RAD markers were developed and a combined SSR and RAD markers linkage map of V. vexillata was constructed. A total of 559 (84 SSR and 475 RAD) markers loci could be assigned to 11 linkage groups spanning 973.9 cM in length with a mean distance of 1.8 cM between adjacent markers. Linkage and genetic position of all SSR markers in an SSR linkage map were confirmed. When an SSR genetic linkage map of V. vexillata was compared with those of V. radiata and V. unguiculata, it was suggested that the structure of V. vexillata chromosome was considerably differentiated. This map is the first SSR and RAD marker-based V. vexillata linkage map which can be used for the mapping of useful traits.     

Seed Morphology, Anatomy and Ultrastructure ofPhyllocladusL. C. & A. Rich. ex Mirb. (Phyllocladaceae (Pilg.) Bessey) in Connection with the Generic System and Phylogeny

To analyse the status and phylogeny of the genusPhyllocladus,seedsof all seven species of the genus were studied. The complexof features of thePhyllocladusreproductive system point to theisolated position of the genus within the conifers. Relationswith Podocarpaceae s. l., Taxaceae s. l. and Cephalotaxaceaeappeared to be remote because a complex of features clearlydistinguishesPhyllocladusfrom the afore-mentioned taxa. We findit advisable to circumscribe the family Phyllocladaceae as Bessey(1907) did, and place it into the order Taxales Knobl. in Warming(1890). From investigations of the seeds it appears the genusPhyllocladusconsistsof seven species, forming five groups. There is a significanttendency for transformation of the female reproductive structureswithin the generic boundaries ofPhyllocladus—seeds, originallysolitary, tending to aggregate in various kinds of compact clusters.Copyright1999 Annals of Botany Company PhyllocladusL. C. & A. Rich, ex Mirb., seed anatomy, seed morphology, systematics, phylogenetic relationships, Phyllocladaceae, Podocarpaceae s. str., Acmopyleaceae, Nageiaceae, Austrotaxaceae, Amentotaxaceae, Torreyaceae, Taxaceae, s. str., Cephalotaxaceae, Taxales.     

Ultrastructure and function of the labial nephridia and the rectum of Orchesella cincta (L.) (Collembola)

Summary The collembolan Orchesella cincta possesses a well-developed coelomoduct kidney. The presence of podocytes in the wall of the sacculus and the fact that the epithelium of the nephridial tubule has the ultrastructural characteristics of resorbing cells, indicate that this is an ultrafiltration-reabsorption kidney.Apparently also the rectum is lined by a reabsorptive epithelium; the cells possess an extensive system of apical and basal infoldings. This view is sustained by the fact that the stereology of the apical channel system varies in animals kept under different moisture conditions. During the intermoult period, both organs are subject to strong morphological changes, which are obviously related to the feeding rhythm.The authors wish to thank Dr. T. Sminia for his stimulating interest during the investigations, Dr. J.C. Jager for statistical advice and Mr. G.W.H. van den Berg for drawing the figures