Physiological integration in Cassia fasciculata Michx.: inflorescence removal and defoliation experiments

Summary In the annual herb Cassia fasciculata virtually every leaf subtends an axillary inflorescence. We examined the degree to which these leaf-inflorescence units (reproductive nodes) were physiologically independent of each other in the production of flowers, fruits, and seeds. Removal of up to 4 of every 5 inflorescences resulted in substantial increases in fruit and seed production by remaining, intact reproductive nodes. These increases nearly compensated for and manipulated reproductive nodes were associated with different vascular strands. When 2 of every 3 leaves were removed, fruit and seed production were reduced at both intact and defoliated reproductive nodes. Taken together, these results suggest that neighboring reproductive nodes in C. fasciculata are not physiologically independent of one another, and that competition among fruits and seeds for parental resources occurs over several reproductive nodes.Scientific contribution no. 1595 from the New Hampshire Agricultural Experiment Station     

Bee-pollination inHibbertia fasciculata (Dilleniaceae)

In direct contrast to mostHibbertia spp., the flowers ofH. fasciculata R. Br. ex D. C. bear only a single whorl of stamens and these stamens are arranged separately (not in typical bundles). The short filaments are appressed to the three carpels so that the inflated, porose and introrsive anthers form a centralized cluster obscuring the three ovaries. The three slender styles emerge at right angles from between the filaments. These styles curve upward and the stigmas form the three points of a triangle; each stigma is approximately one millimeter outside the centralized cluster of anthers. The flowers are nectarless and bear a bright yellow corolla. A pungent and unpleasant fragrance appears to be concentrated within the pollenkitt. When native bees attempt to forage for the pollen, within the cluster of anthers, the ventrally deposited loads of pollen, on the bees' abdomens, contact the outer triangle of stigmas. The major pollinators ofH. fasciculata are female bees in the polylectic genera,Lasioglossum (subgenusChilalictus, Halictidae) andLeioproctus (Colletidae). These bees carry an average of more than two pollen taxa when they are caught foraging onH. fasciculata. 78% of the 47 bees, captured onH. fasciculata carried the pollen from at least one sympatric taxon bearing nectariferous flowers (e.g., genera in theMyrtaceae, Compositae, andEpacridaceae). The pollination biology ofH. fasciculata is assessed in relation to the known radiation of bee-pollinated flowers in the genusHibbertia, and within theDilleniaceae s. l.     

Effect of 2,4-dichlorophenoxyacetic acid on the dark- and light-induced pulvinar movements in Cassia fasciculata Michx

2,4-dichlorophenoxyacetic (2,4-D) applied to excised leaves of Cassia fasciculata modified the dark-induced (scotonasty) and light-induced (photonasty) leaflet movements, showing that this compound acts on rapid turgor variation and the concomitant ion migrations, in particular K⁺. 2,4-D inhibited the scotonastic closure in a dose-dependent manner from 10^–8 M to 10^–5 M and promoted the photonastic opening in the same range of concentrations. The compound acted rapidly since a treatment as short as 5 min gave an obvious effect on the motile reaction; however, a lag period of 45–60 min was needed to observe its effect. Although 2,4-D is a weak acid, its greatest physiological efficiency was obtained with pH values close to neutrality. The physiological results are discussed in relation to the chemical properties and the characteristics of transport of the molecule.Abbreviations ABA abscisic acid - 6-BAP 6-benzylaminopurine - 2,4-D 2,4 dichlorophenoxyacetic acid - GA₃ gibberellic acid - HEPES N-[2-hydroxyethyl] piperazine-N-[2-ethanesulphonic acid] - IAA indole-3-acetic acid - NAA 1-naphthaleneacetic acid - MES 2-(N-morpholino)-ethanesulphonic acid     

Structural aspects of ovule and seed development and nonrandom abortion inMelilotus officinalis (Fabaceae)

Summary Only one ovule matures into a seed inMelilotus officinalis. Although eight ovules form within an ovary, only the basal ovule develops into a mature seed, whereas the other ovules degenerate. The investigation of ovule and seed structure at different developmental stages and a comparison of quantitative characters of differently fated ovules within an ovary were undertaken by light, phase contrast, and fluorescence microscopy. In this species, campylotropous ovules develop simultaneously on marginal placentae in an apocarpous unilocular gynoecium. Megasporo- and megagametogenesis proceed normally and are completed in bud. The maturation of the Polygonum type embryo sac takes place after the flower opens. Shortly before fertilization, synergids show signs of degeneration in all ovules. At this stage, neither the structure nor the sizes of ovules within one ovary differ significantly. In spite of this, only the basal ovule develops into a seed. Rarely, one of the upper-situated ovules or the basal and another ovule mature into seeds. Seed enlargement is insignificant until the stage when globular embryo and nuclear endosperm are formed. At the seed-filling stage, other ovules have collapsed and the seed gradually comes to occupy the total volume of the pod. The fruit-to-seed length ratio decreases considerably during seed ripening. At fertilization, ovary length is four times greater than ovule length. In the mature state, the fruit and seed lengths are approximately equal. Seed size and weight diminish with an increase in seed number within a pod, although pod size remains constant. It is assumed that nonrandom abortion of young seeds inM. officinalis is under maternal control and is not related to structural abnormalities in ovule development or with limitation in pollen. We suppose that evolution of this species may have proceeded in the direction of a decrease in seed number and an increase in its sizes, which may play an important role in seed dispersal and seedling establishment.     

Insect and wind pollination inUrginea maritima (Liliaceae)

Three co-existing pollination mechanisms are found inUrginea maritima: insect-, wind-, and self-pollination. The flowers exhibit a typical insect-pollination syndrome; they offer abundant exposed nectar as well as pollen. Out of the many different visitors only a few could be regarded as pollinators:Apis mellifera, Polistes gallicus, andVespa orientalis. Wind pollination also occurs and generally is responsible for self-pollination.It is argued that the development of extra wind-pollination accompanied by partial self-incompatibility is an adaptation to increase pollination in an unfavourable season (August–September), when insects are scarce.     

Factors affecting intrafruit pattern of ovule abortion and seed production in Hormathophylla spinosa (Cruciferae)

We investigated the pattern of intrafruit seed production in Hormathophylla spinosa (Cruciferae) over a 7-yr period. H. spinosa fruit have two chambers, each chamber containing two ovules, and usually situate perpendicular to the infructescence axis with one chamber above the other. The percentage of ovules ripening to seeds is usually lower than 50%. In addition, we found a consistent position-dependence in seed ripening probability, since more than 90% of the ripe seeds are arranged in the lower chamber of the fruit. This pattern of seed production is not produced by the direct or indirect effect of seed predators, by pollen limitation or by nonrandom fertilizations. By contrast, fruit removal experiments showed that sibling rivalry can partially explain the seed production pattern, be the cause of the reduction in the seed number from the uppermost threshold of 50%. Moreover, experimental manipulations of fruit orientation suggest that the position-dependence in seed production is the cause but not the effect of seed ripening. We presume that some architectural effect is producing a significant increase in the ripening probability of the ovules arranged in the lower chamber, causing thus the observed pattern in the intrafruit seed production in H. spinosa and severely constraining the maximum number of seeds per fruit to two.     

Desmosomes and hemidesmosomes in the flagellate Crithidia fasciculata

Summary The flagellum of the trypanosomatid flagellate Crithidia fasciculata expands asymmetrically as it emerges from the reservoir. Where the flagellar memhrane approaches the membrane lining the reservoir, desmosomes are found. These structures are arranged in several slightly curved lines and have many features in common with vertebrate desmosomes.In cultures, the flagellates stick to each other by their flagella and form rosettes. In these bundles of cells, probable sites of adhesion between flagella, or between flagella and pieces of debris, are marked by a dense filamentous tract which passes posteriorly along the flagellum and by a thick band lying just below the flagellar membrane. It is suggested that similar adhesions are found in the insect host where the flagellate attaches itself to the gut wall.     

Effect of parthenocarpy on ovule development in cucumber (Cucumis sativus L.) after pollination with normal and irradiated pollen

In nonparthenocarpic cucumber, analysis of the interaction between season and female receptivity demonstrated two different modes of fruit development and seed set. Spring climatic conditions induced significantly longer ovaries at anthesis with an autonomous fruit set as in natural parthenocarpy. Autumn climatic conditions induced smaller ovaries with fruit set influenced by the pollination stage of the female flower. These differences involve, during fruit development, a regulation of the empty seed number especially at both ends of female receptivity period. Indeed, seasons have no effect on ovule receptivity and fertility measured by full seed number. It is supposed that priority is given to seeds with an embryo in the competition for limited maternal resources in spring as well as in autumn. In spring, the use of irradiated pollen during the whole female receptivity period did not modify the seed set profile. Moreover, a cytological study of 6-day-old ovules pollinated by irradiated pollen has revealed a similar precocious embryogenesis compared to the control. In cucumber, pollination with irradiated pollen involves stenospermocarpy rather than induced parthenocarpy. However, we have not been able to determine whether the early abortive embryogenesis induced by irradiated pollen is gynogenetic or abnormal zygotic.     

The pollination biology ofHibbertia stricta (Dilleniaceae)

The exines of pollen grains ofHibbertia stricta (DC.)R. Br. exF. Muell. (Sect.Pleurandra) wear an oily, yellow pollen coat that stains positively for lipids. The pollen is collected by asocial bees, exclusively. The most common floral foragers are members of the genusLasioglossum (subgenusChilalictus;Halictidae) and they harvest pollen via thoracic vibration. As these bees cling to the inflated anthers their pollen smeared bodies come in contact with either of the two wet, nonpapillate stigmas. The stigmas respond positively to cytochemical tests for the presence of esterase immediately following expansion of the corolla, indicating the effective pollination period. The foraging patterns of the bees are narrowly to broadly polylectic. AsH. stricta flowers are nectarless, it is not surprising that bees bearing mixed pollen loads always carry the pollen of at least one nectariferous, coblooming plant. The pollination biology ofH. stricta is compared with otherHibbertia spp. and with pollen flowers in general.     

Population variation in plant traits associated with ant attraction and herbivory in Chamaecrista fasciculata (Fabaceae)

The benefits of ant–plant–herbivore interactions for the plant depend on the abundance of ants and herbivores and the selective pressures these arthropods exert. In plants bearing extrafloral nectaries (EFN), different mean trait values may be selected for by different populations in response to local herbivore pressure, ultimately resulting in the evolution of differences in plant traits that attract ants as defensive agents against herbivory. To determine if variation in traits that mediate ant–plant interactions reflect herbivore selective pressures, we quantified intra- and inter-population variation in plant traits for eight populations of the EFN-bearing annual Chamaecrista fasciculata (Michx.) (Fabaceae). Censuses in rural and urban areas of Missouri and Illinois (USA) showed population differences in ant attendance and herbivore pressure. Seeds were collected from each population, and plants were grown in a common greenhouse environment to measure sugar production, nectar volume and composition, EFN size and time of emergence, leaf pubescence, and leaf quality throughout plant development. Populations varied mainly in terms of nectary size, sugar production, and nectar volume, but to a lesser degree in leaf pubescence. Populations of C. fasciculata within urban areas (low in insect abundance) had small nectaries and the lowest nectar production. There was a positive correlation across populations between herbivore density and leaf damage by those herbivores on the one hand and sugar production and nectar volume on the other. These results, in conjunction with lack of evidence for maternally based environmental effects, suggest that population differences in herbivore damage have promoted differential evolution of EFN-related traits among populations. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Bird pollination in South African species ofSatyrium (Orchidaceae)

Field observations showed that three South African orchid species,Satyrium carneum, S. coriifolium andS. princeps, are pollinated by sunbirds. Foraging sunbirds insert their bills into the labellum chamber of the flowers and suck nectar from the labellum spurs with their tongues. The column overarches the entrance to the labellum and pollinaria become affixed to the upper mandible of the bill. Birds often attempt to remove pollinaria by rubbing their bills against a branch, but are mostly unsuccessful due to the large plate-like viscidia which attach the pollinaria very firmly to the bill. Other modifications for bird pollination in theseSatyrium species may include the red, carmine or orange colour of the flowers, sturdy inflorescence stems used for perching and large amounts of dilute nectar in the spurs. EachSatyrium species was pollinated by several species of sunbird, and despite some differences in flowering time, occasional hybrids occur at a site whereS. carneum andS. coriifolium share the same sunbird pollinators.     

Evolution,pollination, and systematics of the tribeNeottieae (Orchidaceae)

The various classifications of the orchid tribeNeottieae are reviewed and a new classification is proposed that divides the tribe into three subtribes,Neottiinae, Limodorinae, andCephalantherinae, based primarily on characters of the column (gynostemium). A cladistic analysis illustrates that these three subtribes are more closely related to one another than either is to any other group in subfam.Neottioideae, although there are very few apomorphic characters for the tribe. Pollination biology is also discussed showing links between breeding systems and distribution. There is also a possible role between column and labellum morphology and the emergence of a deceptive pollination syndrome from one of reward.     

Insect pollination and floral mechanisms in South African species ofSatyrium (Orchidaceae)

The morphologically diverse flowers in the genusSatyrium reflect adaptations to a wide range of pollinators. Several recently discovered pollination systems inSatyrium are described and illustrated here; these include pollination by solitary bees, carrion flies, butterflies and moths. Two basic types of floral mechanism are recognised inSatyrium: (1) Species pollinated by lepidoptera and birds have long floral spurs and plate-like viscidia seated in lateral rostellum notches; these viscidia become attached to the proboscis or bill of the pollinator. (2) Species pollinated by flies and bees have relatively short floral spurs and globose viscidia seated in terminal rostellum notches; these viscidia become attached to the face, thorax or eyes of the pollinator.     

Divergent pollination systems in the cleistogamous species,Collomia grandiflora (Polemoniaceae)

Summary A structural study of pollination in the dimorphic flowers ofCollomia grandiflora, a cleistogamous species, reveals significant differences in stigma behavior during pollination, stylar structure, the timing of generative cell division, and pollen tube growth rate patterns. The cleistogamous flower shows a loss of protandry and the stigma is receptive only after reflexing and closing of its lobes. In contrast, the chasmogamous stigma is receptive when reflexed and closes when pollen has been deposited on the lobes. Pollen tube penetration of the dry stigma papillae and entry into the style is similar in the two morphs. The chasmogamous style is solid and the cleistogamous style partly hollow. The matrix of secretion produced by the transmitting tract cells is mainly carbohydrate with a trace of lipids. It is fibrillar in nature and appears to be partly comprised of wall material from the transmitting tract cells. In the chasmogamous pollen, the generative cell enters the tube before division, which occurs between 30 and 60 min after pollination. This division correlates with an increased growth rate for the pollen tube. In the cleistogamous pollen, contact with the stigma triggers generative cell division inside the hydrated pollen grain before germination. The two resulting sperm cells exit the grain 15–30 min after pollination when the pollen tube is in the stigma lobes. The cleistogamous pollen tube shows only one phase of growth which occurs at a rate similar to that of the slow, first phase of the chasmogamous pollen.Abbreviations CH chasmogamous - CL cleistogamous - DAPI 4, 6-diamidino-2-phenylindole     

The significance of pollination time and frequency and of purity of pollen loads for seed set in Rhinanthus angustifolius (Scrophulariaceae) and Viscaria vulgaris (Caryophyllaceae)

Summary The effects of time of the day and frequency of application and of purity of artificially deposited pollen loads on fruiting and seed set were studied in Rhinanthus angustifolius (diurnally visited by bumblebees) and Viscaria vulgaris (diurnally visited by bumblebees, nocturnally by sphingid moths).Time of the day did not influence pollination success in either species. Increase of pollination frequency increased fruiting and seed set in Rhinanthus but had no affect on Viscaria. Five successive artificial pollinations were needed to achieve seed set equal to that observed naturally in Rhinanthus while a single artificial pollination was sufficient in Viscaria. Mixing Lupinus and Viscaria pollen did not reduce fruiting and seed set in Viscaria. The results are discussed in relation to observed seed sets in early and late flowers, and small and large patches of Viscaria vulgaris and among Rhinanthus flowers in populations of different densities.     

Correlations between 2C DNA values and habit inCassia (Leguminosae:Caesalpinioideae)

Ten species of the genusCassia show a range of 2C DNA amounts from 1.30 to 2.54 pg at the same ploidy level. Remarkably, a distinct 2-fold increase is depicted by an arboreal speciesC. excelsa while the rest comprising of herbs, trees and shrubs have a range from 1.30 to 1.47 pg. These form a natural grouping with respect to mean DNA amounts which differ by 0.05 pg in the herbs, trees and shrubs respectively.     

The fine structure of Crithidia fasciculata with special reference to the organelles involved in the ingestion and digestion of protein

Summary As in other trypanosomatids, the cell membrane of Crithidia fasciculata overlies a single layer of microtubules. Each microtubule possesses a large number of periodically arranged drumstick-like appendages and adjacent microtubules are joined by fibrillar connectives. Anteriorly, the microtubules gradually taper to terminate just before or just after entering the reservoir. An attempt is made to correlate microtubule tapering with maintenance of form of the truncated anterior end of the cell. Smooth and coated vesicles are proliferated from the Golgi saccules and the prominent contractile vacuole lies nearby. The single mitochondrion is extensive and expanded at one point to form a capsule for the kinetoplast. The cristae are predominantly plate-like but other configurations do occur. The cytostome, a shallow invagination of the reservoir membrane, is found between two constrictions in the reservoir wall. Supporting the cytostome are several microtubules which penetrate deeply into the cytoplasm. Ingestion of ferritin occurs by pinocytosis from the cytostome and by coated vesicle formation from the reservoir membrane. Digestion probably occurs in multivesicular bodies which contain acid phosphatase activity.     

Oriole pollination ofErythrina breviflora (Leguminosae): Evidence for a polytypic view of ornithophily

Erythrina breviflora is visited by large numbers of passerine birds of which orioles (Icterus: Icteridae) are the primary pollinators. The flowers produce large quantities of nectar but they are rarely visited by hummingbirds. Inflorescence and floral morphology, and low levels of sucrose in the nectar probably explain the rarity of foraging hummingbirds. A comparison of Old WorldErythrina and their pollinators with New World species pollinated by orioles and hummingbirds suggests that parallel evolution has occurred. When the comparison is expanded to include other species pollinated by orioles, it is clear that various New WorldIcteridae, Thraupidae, etc. are ecological equivalents of Old WorldOriolidae, Pycnonotidae, Sturnidae, etc. and that flowers pollinated by these birds have similar characteristics.     

Notes on the pollination mechanisms ofMoraea inclinata andM. brevistyla (Iridaceae)

Individual flowers ofMoraea inclinata are nectariferous and last about six hours. They appear to be pollinated largely by bees in the familyHalictidae (Lasioglossum spp.,Nomia spp.,Zonalictus) and to a lesser extent by bees in the familyAnthophoridae (Amegilla). The mechanism of bee-pollination inM. inclinata is the Iris type; i.e., each flower consists of three pollination units (an outer tepal, a partly exserted anther, and the opposed style branch which terminates in a pair of petal-like crests). Bees rarely visit more than one pollination unit per flower. Transferral of pollen to the bee is passive and nototribic although all bees collected on the flowers were female and 55% of the bees carried pollen loads with 2–5 pollen taxa in their scopae.Moraea brevistyla flowers are nectariferous but lack scent and last two days. They are visited infrequently by bees and only one femaleLasioglossum spec. carried the pollen ofM. brevistyla. Unlike flowers ofM. inclinata those ofM. brevistyla deposit pollen only on the head and thorax. Bee-mediated autogamy in both species is avoided due to the erratic foraging patterns of the bees and the flexibility of each stigma lobe as the bee backs out of the flower. Approximately 2–4 flowers in the inflorescences of both species (6–8 flowers/infloresence) develop into capsules.