Evidence of Dichogamy in Santalum album L.

Flowering, fruit set, embryological development, and pollination trials were Investigated In Santslum album L. Each ovary may have three to four ovules. Mlcrosporogenesls and megasporogenesls In the same flower were synchronized at the earlier stages of flower development. However, at anthesls, when pollen was mature, the magaspore had developed only to the stage of a one- to two-nucleus embryo sac. As the eight-nucleus embryo sac developed, some mamelon cells began to undergo programmed cell death, forming holes Into which the eight-nucleus embryo sacs extended, becoming ＂N＂ or ＂S＂ shaped. The development from a two-nucleus embryo sac to a matured eight-nucleus embryo sac lasted up to 10 d. Fruit-set from open pollination was less than 2%. The endosperm develops prior to division of the zygotic embryo and one to three embryos and endosperms were formed In the same fruit. A mature seed usually germinates to produce one seedling; however, two and three seedlings from one seed were also observed, albeit at a low frequency. Pollination trials showed that no seed sets when Inflorescences were covered with s bag; however, artificial pollination could Improve fruit set. Our pollinaUon trials and embryological studies proved that the flower of S. album Is dlchogamous and fruit set has high heterozygosity.     

Incompatibility in angiosperms

 Since Darwinian times considerable knowledge has accumulated on the distribution, physiology and genetics of self-incompatibility (SI) in higher plants. In the second half of this century the first attempts were made to identify the biochemical bases of SI. These included thediscovery that cutinase enables pollen tube penetration at the surface* of the stigma in Cruciferae, sorting of segregation pollen S-phenotypes by serological techniques, a lock-and-key model of the SI reaction, the first detection and characterisation of SI proteins and the discovery of the role of the tapetum in the determination of pollen phenotypes in homomorphic sporophytic SI. This pioneering work was followed by a worldwide effort to identify and understand the cellular and molecular processes which lead to the recognition and rejection of SI pollen. The present review article summarizes briefly the current state of knowledge in areas essential for the understanding and exploitation of SI and outlines new information that has become available during recent years. Received: 14 March 1997 / Revision accepted: 10 June 1997     

Stomatal patterning in angiosperms

My thesis is that understanding stomatal patterning requires a holistic perspective. Since stomata are structures critical to the survival of terrestrial plants, they need to be viewed in relation to their function and their interface with other structural components. With this outlook, I begin by discussing pattern types, means of measuring them, advantages of each type of measurement, and then present patterning from evolutionary, physiological, ecological, and organ views. I suggest areas where I believe profitable studies might enable us to better understand stomatal patterning. The final sections of the paper review stomatal patterning on angiosperm leaves and present a theory of patterning. With the abundance of molecular information, and coming genomic sequences and new tools, an opportunity exists to dissect the process of how cells are selected to become different from their neighbors and assume a fate critical to plant survival. Understanding this biological process at the molecular level requires comprehending the broad base on which stomatal patterning rests.     

Late-acting self-incompatibility in angiosperms

In most self-incompatible (SI) plants, pollen tube growth in self-pollinated flowers is inhibited on the stigma or in the style. SI systems that operate in the ovary have been assumed to be extremely rare. Evidence from many plant species is presented to show that the SI barriers in the ovary, described here as late-acting SI systems, are quite common. The late-acting SI systems are divided into four categories: (1) ovarian inhibition of incompatible pollen tubes before the ovule is reached; (2) prefertilization inhibition in the ovule; (3) post-zygotic rejection of the embryo, and (4) ovular inhibition for which the cytological details have not been established. Whether or not post-zygotic incompatibility systems can be distinguished from inbreeding depression depends upon the assumptions underlying the genetic models of self-incompatibility. However, four approaches are outlined that could distinguish between active uniform rejections that are presumably evolved responses to inbreeding depression and the passive, variable failures that are commonly understood to be expressions of typical inbreeding depression. Possible advantages of late-acting SI include an extended period of time over which pollen genotypes may be evaluated by the maternal parent and greater flexibility in the choice of male parents. Due to a paucity of data regarding the genetics and physiology of lateacting SI systems, little can be said at this time about the possible diversity of such systems of their evolutionary relationships with classical gametophytic and sporophytic SI. An hypothesis for the operation of post-zygotic SI is described whereby maternal resources to developing embryos are terminated if the embryo (and/or endosperm) fall below a threshold level of heterosis. This hypothesis is a modification of one first proposed by Westoby and Rice in 1982 to explain variable maternal resource allocation to developing embryos.     

Anther plastids in angiosperms

In the anther of angiosperms, all types of plastids are found in the course of pollen development. They are located in the different cell layers of the microsporangium and have various functions that contribute to the formation of the functional male gametophyte. This includes photosynthesis, stomata opening, sugar storage and/or mobilization, lipid synthesis and secretion for pollenkitt formation, as well as serving as a physiological buffer under stress conditions. They are also involved in plastid inheritance, but to different extents, according to the species. The plastid is a semi-autonomous organelle. Plastid division in the anther is synchronous with cell division, except in the vegetative cell during pollen maturation. Furthermore, recent data seem to show that plastids are affected by programmed cell death and DNA degradation, which occur in the whole anther throughout pollen development. However, the timing of plastid disappearance fluctuates in the different cell layers and also depending on species. In vitro, following androgenesis, plastids that originate in the microspore are responsible for the occurrence of albino plantlets in Poaceae. This trait reflects the relative independence of the plastid genome when compared with that of the nucleus. In this family, microspore plastids may become so involved in programmed cell death that they are unable to follow the alternative sporopohytic program. The different pathways of plastid differentiation in neighboring anther cell layers require an accurate regulation of cell development that remains widely unknown in the anther.     

Dichogamy,gender variation and bet-hedging inPseudowintera colorata

Summary Temporal patterns of variability in the longevity of the male and female phases of individual flowers and in the gender expression of plants of a dichogamous New Zealand tree,Pseudowintera colorata (Winteraceae), were documented in field studies. Two measures for the duration of phases in a dichogamous flower are distinguished; the nominal phases based on morphological features of the flower, and the effective phases reflecting the duration of their functions. Flower and phase longevity and phenotypic gender varied considerably throughout the season and among individuals. Temporal variability in phenotypic gender was loosely synchronized among the 12 plants sampled. Three effects of an environmental factor (temperature) were noted. First, increased temperatures shortened the duration of the female phase but had no effect on the duration of the male phase. Second, pollination frequency was positively correlated with temperature. These results indirectly suggest that increased pollination may shorten the duration of the female phase. Third, average population maleness, measured as the proportion of open flowers in the population on a given day which were in the male phase, was positively correlated with temperature. It is postulated that temperature indirectly influences temporal patterns of gender expression in the population through its differential effects on the longevity of the male and female phases in individual flowers. A theoretical model of bet-hedging shows that, if the direction of an environmental effect on the proportions of the sexual phases is irreversible, selection favours asynchronous dichogamy and reduces the temporal variability as much as possible. If the direction of the response is reversible, heterodichogamy is favoured.     

Dichogamy and sexual dimorphism in floral traits in the andromonoecious Euphorbia boetica

BACKGROUND AND AIMS: Euphorbia boetica (Euphorbiaceae) is a functional andromonoecious species that shows both intra- and interfloral dichogamy, hermaphrodite cyathia being protogynous. Sexual dimorphism of the cyathia of E. boetica is examined according to their gender and arrangement on the inflorescence. METHODS: Data were obtained from two natural populations, where the distribution of male and hermaphrodite cyathia in the inflorescence was recorded. The size, pollen production and viability, and nectar secretion were measured in both types of cyathia. KEY RESULTS: Most cyathia were male at the first levels of the inflorescence, then hermaphrodite cyathia predominated at the successive levels, although at the last levels the proportion of male cyathia increased. Male cyathia at basal positions lack ovaries, whereas those at distal positions showed vestigial ovaries. The size of the cyathia varied significantly depending on the level of the inflorescence where they were produced: those of the last levels were usually smaller. The hermaphrodites were significant bigger than males; however, these differences were due to the differential distribution of each cyathium type in the inflorescence. Male cyathia produced significantly more pollen and nectar than hermaphrodites. CONCLUSIONS: In Euphorbia boetica, basal male cyathia could be explained by the presence of protogyny, and apical male cyathia seem to respond to a preemption of resources. A true dimorphism affecting primary sexual characters and related to gender function appears at lower levels of the inflorescence, whereas an apparent size dimorphism due to positional effects occurs at upper positions. Longevity and distribution of cyathia, and their pattern of nectar production, could improve both male and female fitness.     

Half tetrad analysis in angiosperms

A biochemical technique involving analysis of endosperm is proposed for detecting meiotic crossovers and for gene mapping in angiosperms with bisporic embryo sacs. In bisporic embryo sac development two spores resulting from meiosis-II division of the same meiosis-I daughter cell contribute two thirds of the total genetic information to the triploid endosperm nucleus, the other third coming from a sperm in the fertilizing pollen grain. In controlled crosses where the marker gene codes for allozymes with phenotypes sensitive to gene dosage, the maternal meiotic contribution to the endosperm nucleus may be determined, thereby allowing crossovers between the marker locus and the centromere to be detected.     

Cold resistance in Antarctic angiosperms

Deschampsia antarctica Desv. (Poaceae) and Colobanthus quitensis (Kunth) Bartl. (Cariophyllaceae) are the only two vascular plants that have colonized the Maritime Antarctic. The primary purpose of the present work was to determine cold resistance mechanisms in these two Antarctic plants. This was achieved by comparing thermal properties of leaves and the lethal freezing temperature to 50% of the tissue (LT50). The grass D. antarctica was able to tolerate freezing to a lower temperature than C. quitensis. The main freezing resistance mechanism for C. quitensis is supercooling. Thus, the grass is mainly a freezing‐tolerant species, while C. quitensis avoids freezing. D. antarctica cold acclimated; thus, reducing its LT50. C. quitensis showed little cold‐acclimation capacity. Because day length is highly variable in the Antarctic, the effect of day length on freezing tolerance, growth, various soluble carbohydrates, starch, and proline contents in leaves of D. antarctica growing in the laboratory under cold‐acclimation conditions was studied. During the cold‐acclimation treatment, the LT50 was lowered more effectively under long day (21/3 h light/dark) and medium day (16/8) light periods than under a short day period (8/16). The longer the day length treatment, the faster the growth rate for both acclimated and non‐acclimated plants. Similarly, the longer the day treatment during cold acclimation, the higher the sucrose content (up to 7‐fold with respect to non‐acclimated control values). Oligo and polyfructans accumulated significantly during cold acclimation only with the medium day length treatment. Oligofructans accounted for more than 80% of total fructans. The degrees of polymerization were mostly between 3 and 10. C. quitensis under cold acclimation accumulated a similar amount of sucrose than D. antarctica, but no fructans were detected. The suggestion that survival of Antarctic plants in the Antarctic could be at least partially explained by accumulation of these substances is discussed.     

Aperture pattern ontogeny in angiosperms

Pollen grains display a wide range of variation in aperture number and arrangement (pattern) in angiosperms. Apertures are well-defined areas of the pollen wall surface that permit pollen tube germination. For low aperture numbers, aperture patterns are characteristic of the major taxonomic divisions of angiosperms. This paper presents a developmental model that explains most of the aperture patterns that are recorded in angiosperms. It is based on the analysis of the different events that occur during meiosis and lead to microspore differentiation. It demonstrates that variation occurring during meiosis in angiosperms is sufficient to produce the core morphological set of the most commonly observed pollen morphologies.     

Root-nodule formation in non-leguminous angiosperms

Summary The world-wide survey under the IBP of root-nodule formation in non-leguminous Angiosperms is progressing reasonably satisfactorily, and it is anticipated that when all the results have been collated a useful body of new data will be yielded. In recent studies, also forming part of the IBP, in the author's laboratory, the nodules of further species in the genera Alnus, Myrica, Ceanothus, Coriaria and Dryas have been examined for nitrogen-fixing properties, with positive results. Also the extent to which the nodule endophytes from species of Alnus and Myrica respectively are able to symbiose satisfactorily with other host species in the same genus has been investigated, and the conclusion reached that especially in Myrica there is very considerable specialisation among the endophytes. A marked diurnal variation in the rate of fixation of nitrogen in the nodules of non-legumes growing in a glasshouse lit by daylight has been found, with maximal rates being attained around midday. The implication is that this is the period of the maximal availability of carbohydrates in the nodules, but actual analyses have so far failed to reveal this. Analyses of the amino acid composition of the nodules in several genera have shown that except in Alnus, where citrulline is prominent, asparagine is in most cases the dominant amino acid.     

Asymmetrical crossing barriers in angiosperms

Patterns of reproductive isolation between species may provide insight into the mechanisms and evolution of barriers to interspecific gene exchange. We used data from published interspecific hybridization experiments from 14 genera of angiosperms in order to test for the presence of asymmetrical barriers to gene exchange. Reproductive isolation was examined at three life-history stages: the ability of interspecific crosses to produce seeds, the viability of F1 hybrids, and the fertility of F1 hybrids. Statistically significant asymmetries in the strength of reproductive isolation between species were detected in all genera and at each of the three life-history stages. Asymmetries in seed production may be caused by a variety of mechanisms including differences in stigma/style lengths, self compatibility and differential fruit abortion. Asymmetries in post-zygotic isolation are probably caused by nuclear-cytoplasmic interactions. Asymmetrical reproductive isolation between plant taxa may have important implications for the dynamics of hybrid zones, the direction of genetic introgression and the probability of reinforcement.     

Proposed new realignments in the angiosperms

Our attempt at putatively phylogenetic classifications of Angiospermae, considering our vast ignorance of more than 120 million years of evolution of the class, must be very tentative and elastic to make full use of the new approaches and new data constantly being made available to us. Some of the realignments thus required in my system of classification are here explained. Among others, the Paeoniales are recognized and include Glaucidiaceae; various shifts of families are made within Thei–florae–Violiflorae–Malviflorae; Thymelaeaceae and Simmondsiaceae are transferred to Euphorbiales; Emblingiaceae, Gyrostemonaceae and Bataceae are added to Sapindineae; Fabineae, with Connaraceae, Surianaceae and Fabaceae, are transferred to Rutiflorae; Proteiflorae are placed near Rutiflorae, especially Fabineae; the largely Australasian–African Pittosporales are explained; Cornaceae are severely pruned; Haemodoraceae and Velloziaceae are transferred to Commelinales; Arecales, Cyclanthales and Pandanales are separated into unrelated superorders; and Typhiflorae are moved into closer proximity with Commeliniflorae.