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PubMed2005

Genome size, quantitative genetics and the genomic basis for flower size evolution in Silene latifolia.

BACKGROUND AND AIMS: The overall goal of this paper is to construct an overview of the genetic basis for flower size evolution in Silene latifolia. It aims to examine the relationship between the molecular bases for flower size and the underlying assumption of quantitative genetics theory that quantitative variation is ultimately due to the impact of a number of structural genes. SCOPE: Previous work is reviewed on the quantitative genetics and potential for response to selection on flower size, and the relationship between flower size and nuclear DNA content in S. latifolia. These earlier findings provide a framework within which to consider more recent analyses of a joint quantitative trait loci (QTL) analysis of flower size and DNA content in this species. KEY RESULTS: Flower size is a character that fits the classical quantitative genetics model of inheritance very nicely. However, an earlier finding that flower size is correlated with nuclear DNA content suggested that quantitative aspects of genome composition rather than allelic substitution at structural loci might play a major role in the evolution of flower size. The present results reported here show that QTL for flower size are correlated with QTL for DNA content, further corroborating an earlier result and providing additional support for the conclusion that localized variations in DNA content underlie evolutionary changes in flower size. CONCLUSIONS: The search image for QTL should be broadened to include overall aspects of genome regulation. As we prepare to enter the much-heralded post-genomic era, we also need to revisit our overall models of the relationship between genotype and phenotype to encompass aspects of genome structure and composition beyond structural genes.

Biological Evolution↗

PubMed2004

Flower size variation in Rosmarinus officinalis: individuals, populations and habitats.

BACKGROUND AND AIMS: Flowers are relatively invariant organs within species, but quantitative variation often exists among conspecifics. These variations represent the raw material that natural selection can magnify, eventually resulting in morphological divergence and diversification. This paper investigates floral variability in Rosmarinus officinalis, a Mediterranean shrub. METHODS: Nine populations were selected in three major southern Spanish habitats (coast, lowland and mountains) along an elevation gradient. Flower samples from randomly chosen plants were collected from each population, and a total of 641 flowers from 237 shrubs were weighed while still fresh to the nearest 0.1 mg. Leaves from the same plants were also measured. Variations among habitats, sites and plants were explored with general linear model ANOVA. Leaf-flower covariation was also investigated. KEY RESULTS: Most (58%) mass in flowers was accounted for by the corolla, whose linear dimensions correlated directly with flower mass. Averaged over plants, the mass of a flower varied between 12 mg and 38 mg. Habitat, site (within habitat) and shrub identity had significant effects on mass variance. Flowers from the coast were the smallest (17 mg) and those from the mountains the largest (25 mg on average). A pattern of continuously increasing flower size with elevation emerged which was largely uncoupled from the geographical pattern of leaf size variation. CONCLUSIONS: As regards flower size, a great potential to local differentiation exists in Rosmarinus. Observed divergences accord with a regime of large-bodied pollinator selection in the mountains, but also with resource-cost hypotheses on floral evolution that postulate that reduced corollas are advantageous under prevailingly stressful conditions.

Climate↗

PubMed2005

The pollination ecology of Paraboea rufescens (Gesneriaceae): a buzz-pollinated tropical herb with mirror-image flowers.

BACKGROUND AND AIMS: Gesneriaceae is a pantropical plant family with over 3000 species. A great variety of pollination mechanisms have been reported for the neotropical members of the family, but the details of buzz-pollination and enantiostyly for the family have not been described. We investigated the floral biology and pollination ecology of Paraboea rufescens in Xishuangbanna, south-west China, considering three aspects: (1) the type of enantiostyly exhibited; (2) whether the species is self-compatible; and (3) whether pollinator behaviour could enhance the precision of pollen transfer between flowers of contrasting stylar orientation. METHODS: Flowering phenology was monitored once a month during vegetative growth, and once a week during flowering both in the field and under cultivation. Pollination manipulations and pollinator observation in the field were conducted. KEY RESULTS: Anthesis occurred early during the morning, and flowers remained open for 1-5 d, depending on weather conditions. Controlled pollinations revealed that P. rufescens is self-compatible, and exhibited inbreeding depression in seed set. Plants were pollinator limited in natural populations. The similar stylar deflection among flowers within a plant limits autonomous self-pollination as well as pollination between flowers. Two species of bumble bees (Bombus spp.), Amegila malaccensis and Nomia sp. effectively pollinated P. rufescens. These pollinators visited flowers in search of pollen with almost the same frequency. None of the pollinators appeared to discriminate between left- or right-handed flowers. CONCLUSIONS: Paraboea rufescens exhibits monomorphic enantiostylous flowers and a buzz-pollination syndrome. Floral morphology in P. rufescens and pollinator foraging behaviour seems likely to reduce self-pollination and pollinations between flowers of the same stylar deflection.

Animals↗

PubMed2003

Revisiting phase transition during flowering in Arabidopsis.

Single-phase transition during flowering has been suggested by Hempel and Feldman (1994) [Planta 192: 276]. When early flowering ecotypes of Arabidopsis were microscopically observed, a long day signal simultaneously induced the acropetal (bottom to top) production of flower primordia and the basipetal (top to bottom) differentiation of paraclades (axillary flowering shoots) from the axils of pre-existing leaf primordia. However, this model could not account for the production of an extra number of secondary shoots in the TERMINAL FLOWER 1 overexpressor line or AGL20 overexpressor line in Columbia background with a functional allele of FRIGIDA. We report here that Columbia with a functional allele of FRIGIDA under long days and Columbia under short days show an inflorescence-producing phase between the vegetative and the flower-producing phases, supporting two-step phase transition during flowering. In addition, a late-flowering mutant, fwa shows an inflorescence phase but fca, fy and fve follow a single-phase transition, suggesting flowering time mutations have different effects on phase transition during flowering.

Arabidopsis↗

PubMed2005

Isolation and characterization of flower-specific transcripts in Acacia mangium.

Acacia mangium Willd. is a legume tree species native to subtropical and tropical regions of Asia and Australia. Many features of its flower development are common to other legume tree species. To identify genes involved in its floral development, we constructed a subtractive flower cDNA library against vegetative tissues. The 1123 expressed sequence tags (ESTs) represented 576 unique genes. Macroarray analysis further identified 147 of these genes as specific to the early, late or whole flowering process. Eight percent of these flower-specific genes encode MADS-domain-containing transcription factors and MYB proteins. Four percent encode other transcription factors and 10% encode regulatory proteins such as G proteins, kinases and phosphatases. Flower-specific transcripts for gibberellic acid (GA) synthesis and GA-induced proteins, as well as other stress- and pathogenesis-related genes (9%), implicate their involvement in A. mangium flower development. Eighteen percent of the flower-specific genes encode hypothetical proteins and 18% encode proteins of unknown functions. The RNA blot hybridization confirmed and detailed the expression patterns of selected genes. Functions of the A. mangium flower-specific genes are discussed based on comparison with their Arabidopsis homologues, most of which have been implicated in Arabidopsis floral development. Our work suggests general conservation of floral development in A. mangium and Arabidopsis. Further characterization of the conserved and different flower-specific genes will delineate the flowering process of this important legume tree species and facilitate genetic modification of its reproduction.

Acacia↗

PubMed2003

Evolution of complex flowering strategies: an age- and size-structured integral projection model.

We explore the evolution of delayed age- and size-dependent flowering in the monocarpic perennial Carlina vulgaris, by extending the recently developed integral projection approach to include demographic rates that depend on size and age. The parameterized model has excellent descriptive properties both in terms of the population size and in terms of the distributions of sizes within each age class. In Carlina the probability of flowering depends on both plant size and age. We use the parameterized model to predict this relationship, using the evolutionarily stable strategy (ESS) approach. Despite accurately predicting the mean size of flowering individuals, the model predicts a step-function relationship between the probability of flowering and plant size, which has no age component. When the variance of the flowering-threshold distribution is constrained to the observed value, the ESS flowering function contains an age component, but underpredicts the mean flowering size. An analytical approximation is used to explore the effect of variation in the flowering strategy on the ESS predictions. Elasticity analysis is used to partition the agespecific contributions to the finite rate of increase (lambda) of the survival-growth and fecundity components of the model. We calculate the adaptive landscape that defines the ESS and generate a fitness landscape for invading phenotypes in the presence of the observed flowering strategy. The implications of these results for the patterns of genetic diversity in the flowering strategy and for testing evolutionary models are discussed. Results proving the existence of a dominant eigenvalue and its associated eigenvectors in general size- and age-dependent integral projection models are presented.

Adaptation, Biological↗

PubMed2004

PAF1-complex-mediated histone methylation of FLOWERING LOCUS C chromatin is required for the vernalization-responsive, winter-annual habit in Arabidopsis.

The winter-annual habit (which typically involves a requirement for exposure to the cold of winter to flower in the spring) in Arabidopsis thaliana is mainly due to the repression of flowering by relatively high levels of FLC expression. Exposure to prolonged cold attenuates FLC expression through a process known as vernalization and thus permits flowering to occur in the spring. Here we show that the elevated FLC expression characteristic of nonvernalized winter annuals requires two genes, EARLY FLOWERING 7 (ELF7) and EARLY FLOWERING 8 (ELF8), that are homologs of components of the PAF1 complex of Saccharomyces cerevisiae. Furthermore, ELF7 and ELF8 are also required for the expression of other genes in the FLC clade of flowering repressors such as MAF2 and FLM. FLC, FLM, and MAF2 are involved in multiple flowering pathways that account for the broad effects of elf7 and elf8 mutations on flowering behavior. ELF7 and ELF8 are required for the enhancement of histone 3 trimethylation at Lys 4 in FLC chromatin. This modification of FLC chromatin appears to be required to elevate FLC expression to levels that can delay flowering in plants that have not been vernalized. A model of the role of ELF7, ELF8, and other previously described genes in the modification of the chromatin of flowering repressors is presented.

Acclimatization↗

PubMed2002

Evidence for an evolutionarily conserved interaction between cell wall biosynthesis and flowering in maize and sorghum.

BACKGROUND: Factors that affect flowering vary among different plant species, and in the grasses in particular the exact mechanism behind this transition is not fully understood. The brown midrib (bm) mutants of maize (Zea mays L.), which have altered cell wall composition, have different flowering dynamics compared to their wild-type counterparts. This is indicative of a link between cell wall biogenesis and flowering. In order to test whether this relationship also exists in other grasses, the flowering dynamics in sorghum (Sorghum bicolor (L.) Moench) were investigated. Sorghum is evolutionarily closely related to maize, and a set of brown midrib (bmr) mutants similar to the maize bm mutants is available, making sorghum a suitable choice for study in this context. RESULTS: We compared the flowering time (time to half-bloom) of several different bmr sorghum lines and their wild-type counterparts. This revealed that the relationship between cell wall composition and flowering was conserved in sorghum. Specifically, the mutant bmr7 flowered significantly earlier than the corresponding wild-type control, whereas the mutants bmr2, bmr4, bmr6, bmr12, and bmr19 flowered later than their wild-type controls. CONCLUSION: The change in flowering dynamics in several of the brown midrib sorghum lines provides evidence for an evolutionarily conserved mechanism that links cell wall biosynthesis to flowering dynamics. The availability of the sorghum bmr mutants expands the germplasm available to investigate this relationship in further detail.

Body Height↗

PubMed2006

Patterns of MADS-box gene expression mark flower-type development in Gerbera hybrida (Asteraceae).

BACKGROUND: The inflorescence of the cut-flower crop Gerbera hybrida (Asteraceae) consists of two principal flower types, ray and disc, which form a tightly packed head, or capitulum. Despite great interest in plant morphological evolution and the tractability of the gerbera system, very little is known regarding genetic mechanisms involved in flower type specification. Here, we provide comparative staging of ray and disc flower development and microarray screening for differentially expressed genes, accomplished via microdissection of hundreds of coordinately developing flower primordia. RESULTS: Using a 9K gerbera cDNA microarray we identified a number of genes with putative specificity to individual flower types. Intrestingly, several of these encode homologs of MADS-box transcription factors otherwise known to regulate flower organ development. From these and previously obtained data, we hypothesize the functions and protein-protein interactions of several gerbera MADS-box factors. CONCLUSION: Our RNA expression results suggest that flower-type specific MADS protein complexes may play a central role in differential development of ray and disc flowers across the gerbera capitulum, and that some commonality is shared with known protein functions in floral organ determination. These findings support the intriguing conjecture that the gerbera flowering head is more than a mere floral analog at the level of gene regulation.

Asteraceae↗

PubMed2007

Flower tracking in hawkmoths: behavior and energetics.

As hovering feeders, hawkmoths cope with flower motions by tracking those motions to maintain contact with the nectary. This study examined the tracking, feeding and energetic performance of Manduca sexta feeding from flowers moving at varied frequencies and in different directions. In general we found that tracking performance decreased as frequency increased; M. sexta tracked flowers moving at 1 Hz best. While feeding rates were highest for stationary flowers, they remained relatively constant for all tested frequencies of flower motion. Calculations of net energy gain showed that energy expenditure to track flowers is minimal compared to energy intake; therefore, patterns of net energy gain mimicked patterns of feeding rate. The direction effects of flower motion were greater than the frequency effects. While M. sexta appeared equally capable of tracking flowers moving in the horizontal and vertical motion axes, they demonstrated poor ability to track flowers moving in the looming axis. Additionally, both feeding rates and net energy gain were lower for looming axis flower motions.

Animals↗

PubMed2005

FRIGIDA-independent variation in flowering time of natural Arabidopsis thaliana accessions.

FRIGIDA (FRI) and FLOWERING LOCUS C (FLC) are two genes that, unless plants are vernalized, greatly delay flowering time in Arabidopsis thaliana. Natural loss-of-function mutations in FRI cause the early flowering growth habits of many A. thaliana accessions. To quantify the variation among wild accessions due to FRI, and to identify additional genetic loci in wild accessions that influence flowering time, we surveyed the flowering times of 145 accessions in long-day photoperiods, with and without a 30-day vernalization treatment, and genotyped them for two common natural lesions in FRI. FRI is disrupted in at least 84 of the accessions, accounting for only approximately 40% of the flowering-time variation in long days. During efforts to dissect the causes for variation that are independent of known dysfunctional FRI alleles, we found new loss-of-function alleles in FLC, as well as late-flowering alleles that do not map to FRI or FLC. An FLC nonsense mutation was found in the early flowering Van-0 accession, which has otherwise functional FRI. In contrast, Lz-0 flowers late because of high levels of FLC expression, even though it has a deletion in FRI. Finally, eXtreme array mapping identified genomic regions linked to the vernalization-independent, late-flowering habit of Bur-0, which has an alternatively spliced FLC allele that behaves as a null allele.

Alternative Splicing↗

PubMed1997

Spontaneous flower constancy and learning in honey bees as a function of colour

When presented with an artificial flower patch of blue and yellow pedicellate flowers, individual honey bees, Apis mellifera L., became constant to one of the two flower colours, rarely even sampling the alternative colour. Some bees visited only blue flowers while others visited only yellow flowers. This paper describes the onset of constancy for bees that had had no experience with the experimental apparatus. In 3020 visits, bees failed to land on or drink from the flower colour on which they first landed only 17 times. This behaviour was not modified by quality or quantity of reward, training to the experimental site, group effects or presence of odour during trials. However, when we trained bees to a target painted with two colours and then forced them to sample monomorphic flower patches in sequence, all bees visited the only colour present: yellow or blue. When we subsequently offered these same bees yellow and blue flowers simultaneously (rewarded choices), they became constant. Eleven of 23 bees showed constancy to the less rewarding flower morph without even sampling the alternative. Those bees failed to sample even though they had previously been forced to visit the alternative flower morph, which offered a reward with twice the calories/volume. Constancy is thus spontaneous in honey bees, but it can be hidden by some experimental protocols designed to study learning.1997The Association for the Study of Animal Behaviour

Journal Article↗

PubMed2006

Spatial encoding by bumblebees (Bombus impatiens) of a reward within an artificial flower array.

We presented bumblebees a spatial memory task similar to that used with other species (e.g., cats, dogs, and pigeons). In some conditions we allowed for presence of scent marks in addition to placing local and global spatial cues in conflict. Bumblebees (Bombus impatiens) were presented an array of artificial flowers within a flight cage, one flower offering reward (S+), while the others were empty (S-). Bees were tested with empty flowers. In Experiment 1, flowers were either moved at the time of testing or not. Bees returned to the flower in the same absolute position of the S+ (the flower-array-independent (FAI) position), even if it was in the wrong position relative to the S- and even when new flower covers prevented the use of possible scent marks. New flower covers (i.e., without possible scent marks) had the effect of lowering the frequency of probing behavior. In Experiment 2, the colony was moved between training and testing. Again, bees chose the flower in the FAI position of the S+, and not the flower that would be chosen using strictly memory for a flight vector. Together, these experiments show that to locate the S+ bees did not rely on scent marks nor the positions of the S-, though the S- were prominent objects close to the goal. Also, bees selected environmental features to remember the position of the S+ instead of relying upon a purely egocentric point of view. Similarities with honeybees and vertebrates are discussed, as well as possible encoding mechanisms.

Animals↗

PubMed1999

A novel calcium-binding protein is expressed in Brassica pistils and anthers late in flower development.

We describe the cloning and characterization of PCP, a novel calcium-binding protein that is expressed predominantly in the pistils and anthers of Brassica flowers late in flower development. A PCP cDNA - isolated from a subtracted cDNA library enriched in transcripts present in the pistil late in flower development - potentially encodes a 175 amino acid protein with a calculated molecular weight of 19.1 kDa. Other than limited homology to a repetitive C-terminal polyacidic region of PCP, none of the sequences in the GenBank database shares identity to PCP. This unique protein was purified from an Escherichia coli expression system and shown to bind calcium in a specific manner, both in a protein blot assay and by equilibrium dialysis. PCP binds 29 mol of calcium per mol of PCP protein with an apparent affinity constant of 3.2 x 10(2)/M, values consistent with the presence of a high capacity/low-affinity calcium-binding domain. PCP-specific mRNAs are detected predominantly in the stigma and style of pistils excised from open flowers; much lower levels of expression are seen in anthers of open flowers and in root and leaf tissue. Expression in the pistil steadily increases during flower development and peaks at flower opening. A PCP-specific antibody first detects the protein in pistils at one day prior to flowering, with higher levels of the protein seen in the pistils of open flowers. A low level of the protein is present in anthers of open flowers; however, PCP is not detected in either root or leaf extracts. The pattern of PCP expression is consistent with a possible role for PCP in pollen-pistil interactions or in pistil development. The results are also discussed in light of the central role calcium maintains in pollen tube growth and fertilization.

Amino Acid Sequence↗

PubMed2001

CONSTANS mediates between the circadian clock and the control of flowering in Arabidopsis.

Flowering is often triggered by exposing plants to appropriate day lengths. This response requires an endogenous timer called the circadian clock to measure the duration of the day or night. This timer also controls daily rhythms in gene expression and behavioural patterns such as leaf movements. Several Arabidopsis mutations affect both circadian processes and flowering time; but how the effect of these mutations on the circadian clock is related to their influence on flowering remains unknown. Here we show that expression of CONSTANS (CO), a gene that accelerates flowering in response to long days, is modulated by the circadian clock and day length. Expression of a CO target gene, called FLOWERING LOCUS T (FT), is restricted to a similar time of day as expression of CO. Three mutations that affect circadian rhythms and flowering time alter CO and FT expression in ways that are consistent with their effects on flowering. In addition, the late flowering phenotype of such mutants is corrected by overexpressing CO. Thus, CO acts between the circadian clock and the control of flowering, suggesting mechanisms by which day length regulates flowering time.

Arabidopsis↗

PubMed2000

A MADS domain gene involved in the transition to flowering in Arabidopsis.

Flowering time in many plants is triggered by environmental factors that lead to uniform flowering in plant populations, ensuring higher reproductive success. So far, several genes have been identified that are involved in flowering time control. AGL20 (AGAMOUS LIKE 20) is a MADS domain gene from Arabidopsis that is activated in shoot apical meristems during the transition to flowering. By transposon tagging we have identified late flowering agl20 mutants, showing that AGL20 is involved in flowering time control. In previously described late flowering mutants of the long-day and constitutive pathways of floral induction the expression of AGL20 is down-regulated, demonstrating that AGL20 acts downstream to the mutated genes. Moreover, we can show that AGL20 is also regulated by the gibberellin (GA) pathway, indicating that AGL20 integrates signals of different pathways of floral induction and might be a central component for the induction of flowering. In addition, the constitutive expression of AGL20 in Arabidopsis is sufficient for photoperiod independent flowering and the over-expression of the orthologous gene from mustard, MADSA, in the classical short-day tobacco Maryland Mammoth bypasses the strict photoperiodic control of flowering.

Amino Acid Sequence↗

PubMed2001

Control of flowering time by FLC orthologues in Brassica napus.

FLOWERING LOCUS C (FLC) in Arabidopsis encodes a dosage dependent repressor of flowering. We isolated five FLC-related sequences from Brassica napus (BnFLC1-5). Expression of each of the five sequences in Arabidopsis delayed flowering significantly, with the delay in flowering time ranging from 3 weeks to more than 7 months, relative to the flowering time of 3 weeks in untransformed Ler. In the reciprocal experiment, expression of Arabidopsis FLC (AtFLC) in an early flowering B. napus cultivar delayed flowering by 2-6 weeks, confirming the requirement of this gene for floral repression. In B. napus, we show that late flowering and responsiveness to vernalization correlate with the level of BnFLC mRNA expression. The different BnFLC genes show differential expression in leaves, stems and shoot tips, but expression is not detectable in roots. Vernalization dramatically reduces the level of BnFLC transcript and restores early flowering in the winter cultivar Colombus. We conclude that BnFLC genes confer winter requirement in B. napus and account for the major vernalization-responsive flowering time differences in the different cultivars of B. napus in a manner analogous to that of AtFLC in Arabidopsis ecotypes.

Amino Acid Sequence↗

PubMed2001

Polycomb repression of flowering during early plant development.

All plants flower late in their life cycle. For example, in Arabidopsis, the shoot undergoes a transition and produces reproductive flowers after the adult phase of vegetative growth. Much is known about genetic and environmental processes that control flowering time in mature plants. However, little is understood about the mechanisms that prevent plants from flowering much earlier during embryo and seedling development. Arabidopsis embryonic flower (emf1 and emf2) mutants flower soon after germination, suggesting that a floral repression mechanism is established in wild-type plants that prevents flowering until maturity. Here, we show that polycomb group proteins play a central role in repressing flowering early in the plant life cycle. We found that mutations in the Fertilization Independent Endosperm (FIE) polycomb gene caused the seedling shoot to produce flower-like structures and organs. Flower-like structures were also generated from the hypocotyl and root, organs not associated with reproduction. Expression of floral induction and homeotic genes was derepressed in mutant embryos and seedlings. These results suggest that FIE-mediated polycomb complexes are an essential component of a floral repression mechanism established early during plant development.

Arabidopsis↗