SciCombinator

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Concept: Plant sexuality

319

Recent advances in molecular phylogenetics and a series of important palaeobotanical discoveries have revolutionized our understanding of angiosperm diversification. Yet, the origin and early evolution of their most characteristic feature, the flower, remains poorly understood. In particular, the structure of the ancestral flower of all living angiosperms is still uncertain. Here we report model-based reconstructions for ancestral flowers at the deepest nodes in the phylogeny of angiosperms, using the largest data set of floral traits ever assembled. We reconstruct the ancestral angiosperm flower as bisexual and radially symmetric, with more than two whorls of three separate perianth organs each (undifferentiated tepals), more than two whorls of three separate stamens each, and more than five spirally arranged separate carpels. Although uncertainty remains for some of the characters, our reconstruction allows us to propose a new plausible scenario for the early diversification of flowers, leading to new testable hypotheses for future research on angiosperms.

Concepts: Pollination, Flowering plant, Flower, Pollen, Plant sexuality, Petal, Sepal, Asteraceae

168

The evolution of mating systems, which exhibit an extraordinary diversity in flowering plants, is of central interest in plant biology. Herkogamy, the spatial separation of sexual organs within flowers, is a widespread floral mechanism that is thought to be an adaptive trait reducing self-pollination in hermaphroditic plants. In contrast with previous studies of herkogamy that focused on plants with relatively large floral displays, we here characterized herkogamy in , a model plant with a strong selfing syndrome. Developmental features, reproductive consequences, and genetic architecture of herkogamy were exploited using naturally variable accessions, under both greenhouse and natural conditions. Our results demonstrate that the degree of herkogamy can strongly influence the mating patterns of : approach herkogamy can effectively promote outcrossing, no herkogamy is also capable of enhancing the opportunity for outcrossing, and reverse herkogamy facilitates efficient self-pollination. In addition, we found that the expression of herkogamy in was environment-dependent and regulated by multiple quantitative trait loci. This study reveals how minor modifications in floral morphology may cause dramatic changes in plant mating patterns, provides new insights into the function of herkogamy, and suggests the way for dissecting the genetic basis of this important character in a model plant.

Concepts: Genetics, Biology, Model organism, Arabidopsis thaliana, Pollination, Flowering plant, Flower, Plant sexuality

168

Circumnutation, the helical movement of growing organ tips, is ubiquitous in land plants. The mechanisms underlying circumnutation have been debated since Darwin’s time. Experiments in space and mutant analyses have revealed that internal oscillatory (tropism-independent) movement and gravitropic response are involved in circumnutation. Female flower buds of tape grass (Vallisneria asiatica var. biwaensis) circumnutate on the water surface. Our observations and experiments with an artificial model indicated that gravitropism is barely involved in circumnutation. Instead, we show that helical intercalary growth at the base of peduncle plays the primary role in all movements in Vallisneria. This growth pattern produces torsional bud rotation, and gravity and buoyancy forces have a physical effect on the direction of peduncle elongation, resulting in bud circumnutation on the water surface. In contrast to other water-pollinated hydrophilous plants, circumnutation in Vallisneria enables female flowers to actively collect male flowers from a larger surface area of water.

Concepts: Plant, Bud, Buoyancy, Root, Flower, Plant stem, Plant sexuality, Surface area

29

Bumblebees use invisible temperature patterns on flowers to make foraging decisions.

Concepts: Pollination, Flower, Pollinator decline, Pollination syndrome, Plant sexuality, Bee

28

Asparagus officinalis is most extensively studied species within the genus Asparagus, which is well known as garden asparagus. This species is dioecious with unisexual flowers, which means that generative propagation gives roughly equal number of male and female plants. Male plants are high yielders and preferred commercially over female plants. Tissue culture techniques could efficiently promote vegetative propagation of male plants and pave the way for efficient plant breeding.This chapter describes an efficient micropropagation protocol for developing rapid growing in vitro Asparagus shoot cultures. The source of explants, inoculation, and shoot proliferation, followed by shoot propagation, rooting, and acclimatization is described. The optimal medium for Asparagus micropropagation described in this chapter is composed of MS macro- and microelements and a combination of auxins and cytokinins. Plant growth regulators NAA, kinetin, and BA were used in various concentrations. Three different media representing the whole micropropagation protocol of Asparagus are described; medium for shoot initiation, medium for shoot multiplication, and medium for root formation. By in vitro propagation of Asparagus, root initiation is difficult, but can be promoted by adding growth retardant ancymidol which also greatly promotes shoot development and suppresses callus formation.

Concepts: Plant, Root, Seed, Plant sexuality, Plant reproduction, Vegetative reproduction, Perennial plant, Asparagus

26

To analyze the antioxidant prenylflavonoids in different parts of Macaranga tanarius (M. tanarius) (Euphorbiaceae) including the leaf, petiole, stem, leaflet, flower and fruit (only in female plant), and to evaluate their antioxidant properties.

Concepts: Plant, Fruit, Leaf, Plant morphology, Flower, Plant sexuality, Macaranga, Macaranga tanarius

24

Sex determination is as important for the fitness of plants as it is for animals, but its mechanisms appear to vary much more among plants than among animals, and the expression of gender in plants differs in important respects from that in most animals. In this Minireview, I provide an overview of the broad variety of ways in which plants determine sex. I suggest that several important peculiarities of plant sex determination can be¬†understood by recognising that: plants show an alternation of generations between sporophytic and gametophytic phases (either of which may take control of sex determination); plants are modular in structure and lack a germ line (allowing for a quantitative expression of gender that is not common in animals); and separate sexes in plants have ultimately evolved from hermaphroditic ancestors. Most theorising about sex determination in plants has focused on dioecious species, but we have much to learn from monecious or hermaphroditic species, where sex is determined at the level of modules, tissues or cells. Because of the fundamental modularity of plant development and potentially important evolutionary links between monoecy and dioecy, it may be useful to relax the distinction often made between ‘developmental sex determination’ (which underpins the development of male versus female flowers in monoecious species) and ‘genetic sex determination’ (which underpins the separation of males and females in dioecious species, often mediated by a genetic polymorphism and sex chromosomes). I also argue for relaxing the distinction between sex determination involving a genetic polymorphism and that involving responses to environmental or hormonal cues, because non-genetic cues might easily be converted into genetic switches.

Concepts: Male, Evolution, Female, Chromosome, Plant, Sex, Flower, Plant sexuality

24

Cost efficient foraging is of especial importance for animals like hawkmoths or hummingbirds that are feeding ‘on the wing’, making their foraging energetically demanding. The economic decisions made by these animals have a strong influence on the plants they pollinate and floral volatiles are often guiding these decisions. Here we show that the hawkmoth Manduca sexta exhibits an innate preference for volatiles of those Nicotiana flowers, which match the length of the moth’s proboscis. This preference becomes apparent already at the initial inflight encounter, with the odour plume. Free-flight respiration analyses combined with nectar calorimetry revealed a significant caloric gain per invested flight energy only for preferred-matching-flowers. Our data therefore support Darwin’s initial hypothesis on the coevolution of flower length and moth proboscis. We demonstrate that this interaction is mediated by an adaptive and hardwired olfactory preference of the moth for flowers offering the highest net-energy reward.

Concepts: Fruit, Pollination, Motivation, Flower, Pollination syndrome, Lepidoptera, Pollen, Plant sexuality

19

Pollen transport by water-flow (hydrophily) is a typical, and almost exclusive, adaptation of plants to life in the marine environment. It is thought that, unlike terrestrial environments, animals are not involved in pollination in the sea. The male flowers of the tropical marine angiosperm Thalassia testudinum open-up and release pollen in mucilage at night when invertebrate fauna is active. Here we present experimental evidence that, in the absence of water-flow, these invertebrates visit the flowers, carry and transfer mucilage mass with embedded pollen from the male flowers to the stigmas of the female flowers. Pollen tubes are formed on the stigmas, indicating that pollination is successful. Thus, T. testudinum has mixed abiotic-biotic pollination. We propose a zoobenthophilous pollination syndrome (pollen transfer in the benthic zone by invertebrate animals) which shares many characteristics with hydrophily, but flowers are expected to open-up during the night.

Concepts: Insect, Lake, Oceanography, Pollination, Flower, Pollen, Plant sexuality, Pelagic zone

13

Sex determination in maize involves the production of staminate and pistillate florets from an initially bisexual floral meristem. Pistil elimination in staminate florets requires jasmonic acid signaling, and functional pistils are protected by the action of the silkless 1 (sk1) gene. The sk1 gene was identified and found to encode a previously uncharacterized family 1 uridine diphosphate glycosyltransferase that localized to the plant peroxisomes. Constitutive expression of an sk1 transgene protected all pistils in the plant, causing complete feminization, a gain-of-function phenotype that operates by blocking the accumulation of jasmonates. The segregation of an sk1 transgene was used to effectively control the production of pistillate and staminate inflorescences in maize plants.

Concepts: Evolution, Plant, Reproductive system, Plant morphology, Flower, Pollen, Plant sexuality, Gynoecium