Double fertilization in flowering plants: 1898–2008

The present article gives a brief survey of results of studies in the area of plant embryology directly associated with the discovery made by S.G. Navashin in 1898 of double fertilization in vivo and in vitro. These studies utilized methods of electronic and fluorescence microscopy, cytophotometry, and cultures of isolated ovules, sperm, and the embryo sac central cell. Questions related to the origin of the female gametophyte of flowering plants, double fertilization, and the endosperm are considered. It is emphasized that progress in this field is associated chiefly with the study of molecular processes that regulate the development and functioning of the female gametophyte and the sporophyte on early stages of ontogenesis.     

How many nuclei make an embryo sac in flowering plants?

Research on early-divergent angiosperms, including Amborella, the putative sister to all other extant angiosperms, is increasingly used as a yardstick to infer the nature of the hypothetical ancestral angiosperm. Some traits are relatively diverse (and hence relatively labile) in this phylogenetic grade, compared with the more derived eudicot clade, in which developmental patterns have become increasingly canalized. One of the many mysteries surrounding the origin of the angiosperms is the evolutionary origin of the Polygonum-type embryo sac (monosporic, eight-nucleate and seven-celled) that occurs in the majority of flowering plants. Observations on the megagametophyte of Amborella are conflicting, but a recent report of a supernumerary synergid in this genus raises the question of whether the Polygonum-type embryo sac is derived by duplication of a four-nucleate structure or by reduction from a multicellular structure.     

Trait selection in flowering plants: how does sexual selection contribute?

By highlighting and merging the frameworks of sexual selectionenvisioned by Arnold (1994) and Murphy (1998), we discuss howsexual selection can occur in plants even though individualsdo not directly interact. We review studies on traits that influencepollen export and receipt in a variety of hermaphroditic andgynodioecious plants with the underlying premise that pollinationdynamics influences mate acquisition. Most of the studies reviewedfound that phenotypes that enhance pollen export are in harmonywith those that enhance pollen receipt suggesting that in manycases pollinator visitation rates limit both male and femalefunction. In contrast, fewer traits were under opposing selection;but when they were, the traits most often were associated withenhancing the specific aspects of a given sex function. Ourreview helps clarify and illustrate why sexual selection canbe a component of trait evolution in hermaphrodite plants.     

Male sterility in flowering plants: are plant growth substances involved?

Male sterility in flowering plants is of tremendous importance not only in molecular and developmental studies of stamen and pollen grains and evolutionary studies on the origin of dioecy, but also in its commercial application in hybrid seed production. This paper reviews the literature on the possible involvement of plant growth substances (PGSs) in male sterility, and in normal stamen and pollen development. Different experimental approaches on a number of male sterile systems and normal plants have shown that nearly all PGSs, i.e., gibberellins, cytokinins, auxin, abscisic acid, and ethylene, directly or indirectly influence the expression of male sterility. Analyses of endogenous PGSs have revealed that in male sterile plants the level and/or metabolism of more than one PGS is affected. These studies support the suggestion that it is the relative ratio of various PGSs, rather than any one substance, that is critical for normal stamen and pollen development. It is also proposed that gene-regulated male sterility is likely mediated through an altered balance of endogenous PGSs in developing flowers and stamens.     

Phylogeny of flowering plants by the chloroplast genome sequences: in search of a “lucky gene”

One of the most complicated remaining problems of molecular-phylogenetic analysis is choosing an appropriate genome region. In an ideal case, such a region should have two specific properties: (i) results of analysis using this region should be similar to the results of multigene analysis using the maximal number of regions; (ii) this region should be arranged compactly and be significantly shorter than the multigene set. The second condition is necessary to facilitate sequencing and extension of taxons under analysis, the number of which is also crucial for molecular phylogenetic analysis. Such regions have been revealed for some groups of animals and have been designated as "lucky genes". We have carried out a computational experiment on analysis of 41 complete chloroplast genomes of flowering plants aimed at searching for a "lucky gene" for reconstruction of their phylogeny. It is shown that the phylogenetic tree inferred from a combination of translated nucleotide sequences of genes encoding subunits of plastid RNA polymerase is closest to the tree constructed using all protein coding sites of the chloroplast genome. The only node for which a contradiction is observed is unstable according to the different type analyses. For all the other genes or their combinations, the coincidence is significantly worse. The RNA polymerase genes are compactly arranged in the genome and are fourfold shorter than the total length of protein coding genes used for phylogenetic analysis. The combination of all necessary features makes this group of genes main candidates for the role of "lucky gene" in studying phylogeny of flowering plants.     

What is going on with the hormonal control of flowering in plants?

Molecular genetic studies using Arabidopsis thaliana as a model system have overwhelmingly revealed many important molecular mechanisms underlying the control of various biological events, including floral induction in plants. The major genetic pathways of flowering have been characterized in-depth, and include the photoperiod, vernalization, autonomous and gibberellin pathways. In recent years, novel flowering pathways are increasingly being identified. These include age, thermosensory, sugar, stress and hormonal signals to control floral transition. Among them, hormonal control of flowering except the gibberellin pathway is not formally considered a major flowering pathway per se, due to relatively weak and often pleiotropic genetic effects, complex phenotypic variations, including some controversial ones. However, a number of recent studies have suggested that various stress signals may be mediated by hormonal regulation of flowering. In view of molecular diversity in plant kingdoms, this review begins with an assessment of photoperiodic flowering, not in A. thaliana, but in rice (Oryza sativa); rice is a staple crop for human consumption worldwide, and is a model system of short-day plants, cereals and breeding crops. The rice flowering pathway is then compared with that of A. thaliana. This review then aims to update our knowledge on hormonal control of flowering, and integrate it into the entire flowering gene network.     

Genetic diversity in cultivated plants—loss or stability?

Human activities like urbanisation, the replacement of traditional agriculture systems by modern industrial methods or the introduction of modern high-yielding varieties may pose a danger to the biological diversity. Using microsatellite markers, we analysed samples of cultivated wheat (Triticum aestivum L.) collected over an interval of 40–50 years in four comparable geographical regions of Europe and Asia. No significant differences in both the total number of alleles per locus and in the PIC values were detected when the material collected in the repeated collection missions in all four regions were compared. About two-thirds of the alleles were common to both collection periods, while one-third represented collection mission-specific alleles. These findings demonstrate that an allele flow took place during the adaptation of traditional agriculture to modern systems, whereas the level of genetic diversity was not significantly influenced.Communicated by G. Wenzel     

New insights into cell–cell communications during seed development in flowering plants

The evolution of seeds is a major reason why flowering plants are a dominant life form on Earth. The developing seed is composed of two fertilization products, the embryo and endosperm, which are surrounded by a maternally derived seed coat. Accumulating evidence indicates that efficient communication among all three seed components is required to ensure coordinated seed development. Cell communication within plant seeds has drawn much attention in recent years. In this study, we review current knowledge of cross-talk among the endosperm, embryo, and seed coat during seed development, and highlight recent advances in this field.     

How many flowering plants are pollinated by animals?

It is clear that the majority of flowering plants are pollinated by insects and other animals, with a minority utilising abiotic pollen vectors, mainly wind. However there is no accurate published calculation of the proportion of the ca 352 000 species of angiosperms that interact with pollinators. Widely cited figures range from 67% to 96% but these have not been based on firm data. We estimated the number and proportion of flowering plants that are pollinated by animals using published and unpublished community‐level surveys of plant pollination systems that recorded whether each species present was pollinated by animals or wind. The proportion of animal‐pollinated species rises from a mean of 78% in temperate‐zone communities to 94% in tropical communities. By correcting for the latitudinal diversity trend in flowering plants, we estimate the global number and proportion of animal pollinated angiosperms as 308 006, which is 87.5% of the estimated species‐level diversity of flowering plants. Given current concerns about the decline in pollinators and the possible resulting impacts on both natural communities and agricultural crops, such estimates are vital to both ecologists and policy makers. Further research is required to assess in detail the absolute dependency of these plants on their pollinators, and how this varies with latitude and community type, but there is no doubt that plant–pollinator interactions play a significant role in maintaining the functional integrity of most terrestrial ecosystems.     

How many species of flowering plants are there?

We estimate the probable number of flowering plants. First, we apply a model that explicitly incorporates taxonomic effort over time to estimate the number of as-yet-unknown species. Second, we ask taxonomic experts their opinions on how many species are likely to be missing, on a family-by-family basis. The results are broadly comparable. We show that the current number of species should grow by between 10 and 20 per cent. There are, however, interesting discrepancies between expert and model estimates for some families, suggesting that our model does not always completely capture patterns of taxonomic activity. The as-yet-unknown species are probably similar to those taxonomists have described recently—overwhelmingly rare and local, and disproportionately in biodiversity hotspots, where there are high levels of habitat destruction.     

Do nonasterid holoparasitic flowering plants have plastid genomes?

Past work involving the plastid genome (plastome) of holoparasitic plants has been confined to Scrophulariaceae (or Orobanchaceae) which have truncated plastomes owing to loss of photosynthetic and other genes. Nonasterid holoparasites from Balanophoraceae (Corynaea), Hydnoraceae (Hydnora) and Cytinaceae (Cytinus) were tested for the presence of plastid genes and a plastome. Using PCR, plastid 16S rDNA was successfully amplified and sequenced from the above three holoparasites. The sequence of Cytinus showed 121 single base substitutions relative to Nicotiana (8% of the molecule) whereas higher sequence divergence was observed in Hydnora and Corynaea (287 and 513 changes, respectively). Secondary structural models for these 16S rRNAs show that most changes are compensatory, thus suggesting they are functional. Probes constructed for 16S rDNA and for four plastid-encoded ribosomal protein genes (rps2, rps4, rps7 and rpl16) were used in Southern blots of digested genomic DNA from the three holoparasites. Positive hybridizations were obtained using each of the five probes only for Cytinus. For SmaI digests, all plastid gene probes hybridized to a common fragment ca. 20 kb in length in this species. Taken together, these data provide preliminary evidence suggestive of the retention of highly diverged and truncated plastid genome in Cytinus. The greater sequence divergence for 16S rDNA and the negative hybridization results for Hydnora and Corynaea suggests two possibilities: the loss of typically conserved elements of their plastomes or the complete absence of a plastome.     

How do plants know when other plants are flowering? Resource depletion,pollen limitation and mast‐seeding in a perennial wildflower

Mast-seeding is the synchronous and periodic reproduction by plant populations. This phenomenon has been widely studied from a community-level perspective, but we know extremely little about how plants are able to synchronize reproduction. Here, we present the first experimental test of proximate mechanisms of mast-seeding, by preventing reproduction in an iteroparous, mast-seeding wildflower. Through a series of experiments, we show that mobile carbohydrate stores (NSC) control alternate-year flowering by individual plants; seed set depletes NSC which prevents flowering the following year. Plants are synchronized by density-dependent pollen limitation; when plants flower asynchronously, they set fewer seeds, which prevents NSC depletion. Therefore, these individual plants flower in subsequent years and become synchronized. Because mast-seeding is a consequence of physiological controls of reproduction, differences in plant resource acquisition and allocation could dramatically change patterns of seed production, and changes in plant consumers and pollinators could change selection on physiological and developmental pathways.     

The best of both worlds? A review of delayed selfing in flowering plants

Premise of Study

In a seminal body of theory, Lloyd showed that the fitness consequences of selfing will depend on its timing in anthesis. Selfing that occurs after opportunities for outcrossing or pollen dispersal can provide reproductive assurance when pollinators are limited and is expected to incur little cost, even when inbreeding depression is high. As a result, delayed selfing is often interpreted as a “best‐of‐both‐worlds” mating system that combines the advantages of selfing and outcrossing.

Methods

We surveyed 65 empirical studies of delayed selfing, recording floral mechanisms and examining information on inbreeding depression, autofertility, and other parameters to test the support for delayed selfing as a best‐of‐both‐worlds strategy.

Key Results

Phylogenetic distribution of the diverse floral mechanisms suggests that some basic floral structures may predispose plant taxa to evolve delayed selfing. Delayed selfing appears to serve as a best‐of‐both‐worlds strategy in some but not all species. While the capacity for autonomous selfing is often high, it is lower, in some cases, than in related species with earlier modes of selfing. In other delayed‐selfers, low inbreeding depression and reduced investment in corollas and pollen suggest limited benefits from outcrossing.

Conclusions

Despite a growing literature on the subject, experimental evidence for delayed selfing is limited and major gaps in knowledge remain, particularly with respect to the stability of delayed selfing and the conditions that may favor transitions between delayed and earlier selfing. Finally, we suggest a potential role of delayed selfing in facilitating transitions from self‐incompatibility to selfing.     

Death without sex—the ‘problem of the small’ and selection for reproductive economy in flowering plants

Most of the resident plants within vegetation fail to leave descendants because of death without sex—i.e. sexual reproduction fails (zero fecundity), primarily because of relatively small plant size. I propose that this ‘problem of the small’ represents one of the principal driving forces of evolution by natural selection, and that the main product of this selection is ‘reproductive economy’, manifested by several plant traits that are widely distributed among angiosperms: sexual maturity at a relatively young age and small size, relatively small seed size, selfing (including through mixed mating), and of particular interest here, clonality. In non-clonal species, an offspring develops from a zygote into a single ‘rooted unit’, i.e. a distinct vascular transition point between live shoot and root tissue. Clonal species can produce an indeterminate number of these rooted unit offspring asexually, all as products of a single zygote. Clonality is a common strategy in angiosperms because it confers a potential two-fold fitness benefit—especially in relatively small species—by promoting longevity of the zygote product, while at the same time providing a fecundity supplement (through asexual multiplication of rooted units), thereby allowing offspring production economically, i.e. without requiring large adult size, and without even requiring the fertilization of ovules. The primary fitness benefit from clonality, therefore, is that the somatic product of a zygote can effectively avoid an intrinsic limitation predicted for all non-clonal plants: the trade-off between longevity and the potential rate of offspring/descendant production. These major fitness benefits of clonality are explored in considering why clonality is less common in larger species, why the largest species (trees) generally do not have the longest-lived zygote product, and in re-assessing traditional and recent views concerning the loss of sex in clonal plants, the predicted trade-off between the size and number of clonal offspring, and the predicted trade-off between sexual and asexual reproduction.     

γ-Tocopherol biokinetics and transformation in humans

Background: The uptake and biotransformation of γ-tocopherol (γ-T) in humans is largely unknown. Using a stable isotope method we investigated these aspects of γ-T biology in healthy volunteers and their response to γ-T supplementation.

Methods: A single bolus of 100 mg of deuterium labeled γ-T acetate (d²-γ-TAC, 94% isotopic purity) was administered with a standard meal to 21 healthy subjects. Blood and urine (first morning void) were collected at baseline and a range of time points between 6 and 240 h post-supplemetation. The concentrations of d² and d⁰-γ-T in plasma and its major metabolite 2,7,8-trimethyl-2-(b-carboxyethyl)-6-hydroxychroman (-γ-CEHC) in plasma and urine were measured by GC-MS. In two subjects, the total urine volume was collected for 72 h post-supplementation. The effects of γ-T supplementation on α-T concentrations in plasma and α-T and γ-T metabolite formation were also assessed by HPLC or GC-MS analysis.

Results: At baseline, mean plasma α-T concentration was approximately 15 times higher than γ-T (28.3 vs. 1.9 µmol/l). In contrast, plasma γ-CEHC concentration (0.191 µmol/l) was 12 fold greater than α-CEHC (0.016 µmol/l) while in urine it was 3.5 fold lower (0.82 and 2.87 µmol, respectively) suggesting that the clearance of α-CEHC from plasma was more than 40 times that of γ-CEHC. After d²-γ-TAC administration, the d² forms of γ-T and γ-CEHC in plasma and urine increased, but with marked inter-individual variability, while the d⁰ species were hardly affected. Mean total concentrations of γ-T and γ-CEHC in plasma and urine peaked, respectively, between 0–9, 6–12 and 9–24 h post-supplementation with increases over baseline levels of 6–14 fold. All these parameters returned to baseline by 72 h. Following challenge, the total urinary excretion of d²-γ-T equivalents was approximately 7 mg. Baseline levels of γ-T correlated positively with the post-supplementation rise of (d⁰ + d²) – γ – T and γ-CEHC levels in plasma, but correlated negatively with urinary levels of (d⁰ + d²)-γ-CEHC. Supplementation with 100 mg γ-TAC had minimal influence on plasma concentrations of α-T and α-T-related metabolite formation and excretion.

Conclusions: Ingestion of 100mg of γ-TAC transiently increases plasma concentrations of γ-T as it undergoes sustained catabolism to CEHC without markedly influencing the pre-existing plasma pool of γ-T nor the concentration and metabolism of α-T. These pathways appear tightly regulated, most probably to keep high steady-state blood ratios α-T to γ-T and γ-CEHC to α-CEHC.     

Preparing threatened plants for translocation: does home soil addition and nutrient loading improve growth and flowering?

Plant Ecology - Translocation of threatened plants is increasingly being used as a conservation or mitigation action. The success of this practice is mixed and methods to increase likelihood of...     

Genetic perspectives on “Lion Conservation Units” in Eastern and Southern Africa

Current understanding of genetic variation in lions (Panthera leo) is inadequate to guide many management decisions necessary for conservation of the species. We studied sequence variation in the mitochondrial cytochrome-b (cyt-b) gene of 75 lions and nuclear variation at 11 microsatellite loci of 480 lions across 8 range states (Cameroon, Uganda, Kenya, Zambia, Zimbabwe, South Africa, Botswana, and Namibia) and 13 Lion Conservation Units (LCUs) plus two other unassigned sites (Cameroon and Zimbabwe). A total of 11 cyt-b haplotypes were found, whose variation follows an isolation-by-distance model. In combination with previously known sequences, the haplotypes document the close relationship, derived position, and limited variability of Asian and West and Central African lions relative to other extant lions. Both phylogenetic analyses and substitution networks identify two clades in Eastern and Southern Africa—one restricted to Namibia and South Africa and the other more widespread across the region. However, these analyses are equivocal on which of these is closest to the ancestor of modern lions. Microsatellite analyses showed high levels of variation within and among populations, subdivision among most LCUs, and evidence of isolation by distance. While rates of gene flow are generally low, admixture among lions in northern Botswana, Caprivi Strip (Namibia) and Zambia is apparent from STRUCTURE analyses. Conservation management plans should incorporate information on genetic variability and gene flow in delimiting management units and in guiding translocations of lions to minimize inbreeding and to control problem animals.     

The birds, the bees, and the virtual flowers: can pollinator behavior drive ecological speciation in flowering plants?

Biologists have long assumed that pollinator behavior is an important force in angiosperm speciation, yet there is surprisingly little direct evidence that floral preferences in pollinators can drive floral divergence and the evolution of reproductive (ethological) isolation between incipient plant species. In this study, we expose computer-generated plant populations with a wide variation in flower color to selection by live and virtual hummingbirds and bumblebees and track evolutionary changes in flower color over multiple generations. Flower color, which was derived from the known genetic architecture and phenotypic variance of naturally occurring plant species pollinated by both groups, evolved in simulations through a genetic algorithm in which pollinator preference determined changes in flower color between generations. The observed preferences of live hummingbirds and bumblebees were strong enough to cause adaptive divergence in flower color between plant populations but did not lead to ethological isolation. However, stronger preferences assigned to virtual pollinators in sympatric and allopatric scenarios rapidly produced ethological isolation. Pollinators can thus drive ecological speciation in flowering plants, but more rigorous and comprehensive behavioral studies are required to specify conditions that produce sufficient preference levels in pollinators.