Diversification of floral orientation in Lonicera is associated with pollinator shift and flowering phenology

Floral orientation has been suggested to be a functional floral trait that can enhance reproductive fitness through adaptation to both biotic factors (e.g., pollination) and abiotic factors (e.g., temperature). However, whether those factors drive the diversification of floral orientation needs to be explored through multiple related species. Phylogenetic relationships in Lonicera were identified by incorporating the species’ floral orientation (upward vs. downward) and pollinator types. Furthermore, transitions in floral orientation were evaluated together with pollinator shift, and seven Lonicera species were used to detect the influence of floral orientation on pollination and reproductive success. The relationship between temperature and the angle of floral orientation was investigated. Floral orientation in Lonicera showed that most of the closely related species within the same node displayed a similar pattern of floral orientation, and pollinator shift can lead to transitions between upward and downward orientation. Additionally, the angle of floral orientation increased with flowering phenology from spring to summer, and changing floral orientation significantly decreased pollination and seed production. Moreover, the average daily temperature during flowering phenology was significantly correlated with the angle of floral orientation. Our results indicate that pollinator shift could be an important event, leading to the diversification of floral orientation. Results of field investigations inferred that floral orientation should be a functional floral trait adapted to flowering phenology. Downward-facing flowers might help decrease heat loss to adapt to cold conditions. This study disentangled the influences of historical events and local adaptation on the evolution of floral orientation.     

Body condition is associated with a shorter breeding season and reduced ovulation rate in subtropical goats

To determine the effect of body condition on the characteristics of the breeding season in female goats from subtropical Mexico, does in either greater- or lesser-body condition (n=20/group) were monitored for the expression of estrous behavior, ovulation and ovulation rate between June and April. The commencement of estrus and ovulation occurred earlier (P<0.05) in does in greater than lesser body condition. The cessation of estrus and ovulation was later (P<0.05) in female goats with greater than those with lesser body condition. Does in greater body condition had more (P<0.001) normal estrous cycles than those in lesser body condition. As consequence, does from the greater body condition group had fewer short (P<0.001) or long (P<0.05) estrous cycles than those of in the lesser body condition group. The ovulation rate was greater (P<0.01) in the greater (1.9+/-0.1) than lesser body condition does (1.6+/-0.1). In conclusion, female goats in lesser body condition have a shorter breeding season, more abnormal estrous cycles, and fewer ovulations than does in greater body condition.     

Parasitic mite and trematode abundance are associated with reduced reproductive output and physiological condition of freshwater mussels

Although historically understudied, parasites may play an important role in freshwater invertebrate population ecology and evolution. We quantified abundance of parasitic mites and trematodes in the freshwater mussel Pyganodon grandis, in a southeastern Alabama stream (USA), to assess parasite impact on adult mussel physiological condition and reproductive output. We used stepwise multiple regression analyses to assess the effects of mussel size and parasites on reproduction and condition. Multiple regression analysis found no multivariate models that predicted reproductive output or physiological condition. However, univariate models revealed that increased parasite densities predict reduced mussel reproductive output and physiological condition. These findings suggest that parasites may have important negative consequences for freshwater mussels. We hypothesize that elevated parasite loads may reduce mussel fitness in impounded or nutrient-enriched streams with high densities of intermediate hosts (chironomid midges).     

Tree size and its relationship with flowering phenology and reproductive output in Wild Nutmeg trees

Reproductive strategies, sexual selection, and their relationship with the phenotype of individuals are topics widely studied in animals, but this information is less abundant for plants. Variability in flowering phenology among individuals has direct impact on their fitness, but how reproductive phenology is affected by the size of the individuals needs further study. We quantified the flowering intensity, length, and reproductive synchronization of two sympatric dioecious Wild Nutmeg tree species (Virola, Myristicaceae) in the Brazilian Atlantic forest, and analyzed its relationships with tree size. Two distinct strategies in flowering timing and intensity were found between species (annual versus biennial flowering), and among individuals in the annual flowering species (extended versus peak flowering). Only for the annual flowering species the reproductive output is related to tree size and large trees present proportionally higher flower coverage, and lower synchronization than smaller ones. Flowering is massive and highly synchronized in the biennial species. Sex ratios are not different from 1:1 in the two species, and in the two segregated reproductive subgroups in the biennial flowering species. The biennial flowering at individual level is a novelty among reproductive patterns in plants, separating the population in two reproductive subgroups. A proportional increase in the reproductive output with size exists only for the annual flowering species. A biennial flowering can allow resource storage favouring massive flowering for all the individuals diluting their relationship with size.     

Agonistic reciprocity is associated with reduced male reproductive success within haremic social networks

While individual variation in social behaviour is ubiquitous and causes social groups to differ in structure, how these structural differences affect fitness remains largely unknown. We used social network analysis of replicate bluebanded goby (Lythrypnus dalli) harems to identify the reproductive correlates of social network structure. In stable groups, we quantified agonistic behaviour, reproduction and steroid hormones, which can both affect and respond to social/reproductive cues. We identified distinct, optimal social structures associated with different reproductive measures. Male hatching success (HS) was negatively associated with agonistic reciprocity, a network structure that describes whether subordinates ‘reciprocated’ agonism received from dominants. Egg laying was associated with the individual network positions of the male and dominant female. Thus, males face a trade-off between promoting structures that facilitate egg laying versus HS. Whether this reproductive conflict is avoidable remains to be determined. We also identified different social and/or reproductive roles for 11-ketotestosterone, 17β-oestradiol and cortisol, suggesting that specific neuroendocrine mechanisms may underlie connections between network structure and fitness. This is one of the first investigations of the reproductive and neuroendocrine correlates of social behaviour and network structure in replicate, naturalistic social groups and supports network structure as an important target for natural selection.     

Effects of season and population size on pollination and reproductive output in a Mediterranean shrub

• Pollinator guilds may change throughout extended flowering periods, affecting plant reproductive output, especially in seasonal climates. We hypothesised a seasonal shift in pollinator guild and an autumn reduction in pollinator abundance, especially in small and sparse populations.
• We recorded pollinator identity, abundance and behaviour in relation to flower density from plant to population throughout the extended flowering of Ononis tridentata. We evaluated female reproductive output by recording pollination success and pre‐dispersal seed predation in eight populations of contrasting size and density. Offspring quality was also characterised through seed weight and germination.
• A diverse guild of insects visited O. tridentata in spring, while only Apis mellifera was observed in autumn. Visitation frequency did not vary seasonally, but the number of flowers per foraging bout was lower, and seeds were heavier and had a higher germination rate in autumn. Plant and neighbourhood flowering display were not related to pollinator visitation frequency or behaviour. However, the rate of fertilised ovules, seed set and autumn flowering display size were positively related to population density.
• The maintenance of pollination in autumn enhances the reproductive performance of O. tridentata due to higher quality of autumn seed, and to a large reduction in seed predator pressure. We also suggest that observed changes in pollinator behaviour could be one of the processes behind seasonal variation in seed performance, since geitonogamous crosses were less likely to occur in autumn.

     

early bolting in short days: an Arabidopsis mutation that causes early flowering and partially suppresses the floral phenotype of leafy

The time of flowering in Arabidopsis is controlled by multiple endogenous and environmental signals. Some of these signals promote the onset of flowering, whereas others repress it. We describe here the isolation and characterization of two allelic mutations that cause early flowering and define a new locus, EARLY BOLTING IN SHORT DAYS (EBS). Acceleration of flowering time in the ebs mutants is especially conspicuous under short-day photoperiods and results from a reduction of the adult vegetative phase of the plants. In addition to the early flowering phenotype, ebs mutants show a reduction in seed dormancy, plant size, and fertility. Double mutant analysis with gibberellin-deficient mutants indicates that both the early-flowering and the precocious-germination phenotypes require gibberellin biosynthesis. Analysis of the genetic interactions among ebs and several mutations causing late flowering shows that the ft mutant phenotype is epistatic over the early flowering of ebs mutants, suggesting that the precocious flowering of ebs requires the FT gene product. Finally, the ebs mutation causes an increase in the level of expression of the floral homeotic genes APETALA3 (AP3), PISTILLATA (PI), and AGAMOUS (AG) and partially rescues the mutant floral phenotype of leafy-6 (lfy-6) mutants. These results suggest that EBS participates as a negative regulator in developmental processes such as germination, flowering induction, and flower organ specification.     

Seed ecology of a Mediterranean perennial herb with an exceptionally extended flowering and fruiting season

Lobularia maritima is a Mediterranean short-lived herb with a flowering and fruiting season that lasts for ten months. Previous studies have shown that recruitment in periods other than autumn of the flowering season has few demographic implications; that is contributes little to the population growth rate. Since environmental conditions in periods other than autumn would allow recruitment, we examined to what extent the seed ecology of L. maritima accounts for recruitment shortage for the greater part of the year. To this end, we studied the effects of selfing and outcrossing on seed production and germination, within- and between-year variation in seed mass and germination, seed characteristics in the soil seed bank throughout the year, and the effect of temperature as a factor controlling seed germination. Results indicate that selfing does not decrease recruitment, and thus the observed changes in visitation rate and pollinator composition throughout the year cannot account for differences in recruitment. Germinability decreases throughout the year, suggesting a possible cost in reproduction associated with extended flowering. L. maritima has a transient seed bank whose seeds also experience a decrease in their germination throughout the year. Finally, temperature affects seed germination patterns, indicating the existence of quiescence mechanisms that prevent germination in the months prior to the summer drought. Overall, the results obtained support and, at least partly, explain the recruitment patterns of L. maritima observed in the field.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 273–280.     

Effects of shift in flowering time on the reproductive output of Xanthium canadense in a seasonal environment

We studied the effects of a change in flowering date on the reproductive output of a short-day annual plant, Xanthium canadense. The flowering date was changed by photoperiodic manipulation to 1 month earlier or later than the natural flowering date. Plants with the natural flowering date attained the highest reproductive output. For those flowering 1 month earlier or later, the reproductive output was decreased by 42% or 23%, respectively. The reproductive output was analyzed as the product of the biomass production during the reproductive period and its allocation to the reproductive organs. Although delay in flowering increased biomass production, it decreased its fractional allocation to the reproductive organs. The highest reproductive output in the natural flowering plants resulted from a compromise between these two effects of flowering. Plants flowering earlier had higher translocation rates to the reproductive organs and accelerated plant senescence. Later flowering caused a reduction in biomass translocation to the reproductive organs and thus extended the reproductive period. These experimental results are discussed in relation to the cost of reproduction and the optimal time for flowering that maximizes the final reproductive output. It is suggested that the natural flowering time maximized the reproductive output while minimizing the cost of reproduction. Received: 11 September 1997 / Accepted: 12 December 1997     

Phytochrome control of flowering is temperature sensitive and correlates with expression of the floral integrator FT

In Arabidopsis flowering is accelerated by reduced red:far-red (R:FR) ratio which signals the presence of neighbouring vegetation. Hastened flowering is one component of the shade-avoidance syndrome of responses, which alter many aspects of development in response to the threat of potential competition. Of the red/far-red-absorbing photoreceptors it is phyB that plays the most prominent role in shade-avoidance, although other related phytochromes act redundantly with phyB. It is well established that the phyB mutant has a constitutively early flowering phenotype. However, we have shown that the early flowering phenotype of phyB is temperature-dependent. We have established that this temperature-sensitive flowering response defines a pathway that appears to be independent of the autonomous-FLC pathway. Furthermore, we have demonstrated that the phytochromes control the expression of the floral promoter FT. We have also shown that other phyB-controlled responses, including petiole elongation, are not sensitive to the same temperature change. This suggests that discrete pathways control flowering and petiole elongation, components of the shade-avoidance response. This work provides an insight into the phytochrome and temperature interactions that maintain flowering control.     

Selection on flowering time and floral display in an alpine and a lowland population of Arabidopsis lyrata

To determine whether population differentiation in flowering time is consistent with differences in current selection, we quantified phenotypic selection acting through female reproductive success on flowering phenology and floral display in two Scandinavian populations of the outcrossing, perennial herb Arabidopsis lyrata in two years. One population was located in an alpine environment strongly affected by grazing, whereas the other was close to sea level and only moderately affected by herbivory. Multiple regression models indicated directional selection for early end of flowering in one year in the lowland population, and directional selection for early start of flowering in one year in the alpine population. As expected, there was selection for more inflorescences in the lowland population. However, in the alpine population, plants with many inflorescences were selectively grazed and the number of inflorescences produced was negatively related to female fitness in one year and not significantly related to female fitness in the second year. The results are consistent with the hypothesis that genetic differentiation in flowering phenology between the study populations is adaptive, and indicate that interactions with selective grazers may strongly influence selection on floral display in A. lyrata.     

Temporal variation in the female components of reproductive success over the extended flowering season of a Mediterranean perennial herb

We examined the reproductive success of the perennial herb Lobularia maritima during its extended flowering and fruiting season. The within- and between-year variability of the female components of reproductive success (from flower, fruit and seed production to seed survival, seed germination and seedling establishment) were analysed during four flowering seasons. All the components of reproductive success studied showed a significant within- and between-year variation. September was the period of the year with the maximum values of flower and fruit production, and the highest germination and establishment rates. Nevertheless, seed losses due to both predispersal and postdispersal seed predation during this period were also the highest, seriously reducing seed output in this period. On the other hand, in those periods in which seed production was low, i.e. January and May, the percentage of seeds lost to seed predators was the lowest. Reproductive success in each period of the flowering season was estimated using a simple demographic model, in which the information concerning all the components already calculated was integrated. The two variables used to estimate reproductive success in each period, i.e. the number of new individuals produced per plant and the probability of a seed becoming an adult plant, showed relatively small differences over the year. These results suggest a counter-balance of the different components of reproductive success in this species, with favourable and unfavourable periods for the different components being compensated during its extended flowering season.     

early in short days 4, a mutation in Arabidopsis that causes early flowering and reduces the mRNA abundance of the floral repressor FLC

The plant shoot is derived from the apical meristem, a group of stem cells formed during embryogenesis. Lateral organs form on the shoot of an adult plant from primordia that arise on the flanks of the shoot apical meristem. Environmental stimuli such as light, temperature and nutrient availability often influence the shape and identity of the organs that develop from these primordia. In particular, the transition from forming vegetative lateral organs to producing flowers often occurs in response to environmental cues. This transition requires increased expression in primordia of genes that confer floral identity, such as the Arabidopsis gene LEAFY. We describe a novel mutant, early in short days 4 (esd4), that dramatically accelerates the transition from vegetative growth to flowering in Arabidopsis: The effect of the mutation is strongest under short photoperiods, which delay flowering of Arabidopsis: The mutant has additional phenotypes, including premature termination of the shoot and an alteration of phyllotaxy along the stem, suggesting that ESD4 has a broader role in plant development. Genetic analysis indicates that ESD4 is most closely associated with the autonomous floral promotion pathway, one of the well-characterized pathways proposed to promote flowering of Arabidopsis: Furthermore, mRNA levels of a floral repressor (FLC), which acts within this pathway, are reduced by esd4, and the expression of flowering-time genes repressed by FLC is increased in the presence of the esd4 mutation. Although the reduction in FLC mRNA abundance is likely to contribute to the esd4 phenotype, our data suggest that esd4 also promotes flowering independently of FLC. The role of ESD4 in the regulation of flowering is discussed with reference to current models on the regulation of flowering in Arabidopsis.     

Paclobutrazol or uniconazol applied early in the previous season promotes flowering of field-grown Rhododendron and Kalmia

Field-grown large leaf Rhododendron and Kalmia latifolia were treated with one of three concentrations of paclobutrazol or uniconazol in April, June, or August in the second year from propagation. The elongation of stems was measured as was the number of flower buds initiated. Spray applications in April or June increased flowering at the lowest concentrations tested; 4 mg · L^–1 paclobutrazol and 1.5 mg · L^–1 uniconazol. Flowering was induced in cultivars that normally did not flower, and the number of flower buds per plant was increased in cultivars that normally flowered. All treatments that induced flowering also reduced stem elongation. Spray application in August failed to enhance flowering. At the concentrations tested, uniconazol was more effective than paclobutrazol in increasing the number of flower buds and reducing stem elongation of Rhododendron. For Kalmia, there was less response to the concentration of growth retardant, and the two chemicals enhanced flowering equally.     

Fecundity and reproductive output of the caridean shrimp Periclimenes paivai associated with scyphozoan jellyfish

Periclimenes paivai is notable for its symbiotic lifestyle with jellyfish. The objective of this study was to investigate the fecundity and reproductive output of females, and specifically to determine whether females protected in their host have a greater reproductive success than other shrimp species. All anatomical parts of the medusae were analyzed and the P. paivai individuals were separated. Among 83 females analyzed, 65 presented embryos in the first developmental stage, and 18 were in the final stage of development. The mean fecundity was 229.08 ± 120.04 in initial-stage embryos and 191 ± 114.76 in final-stage embryos. We observed a positive relationship between the embryo number and the female size; however, the embryo number was not related to the embryonic stage, which indicates that females did not lose their embryos during the incubation period. The weight of brood mass during production is closely related to the female weight according to the results of linear regression, as fecundity increased with female size. In its symbiosis, P. paivai can increase the amount of energy used to produce embryos (mean reproductive output = 10.38%) and can elevate the reproductive success.     

Later chronotype is associated with higher hemoglobin A1c in prediabetes patients

The circadian system is known to play a role in glucose metabolism. Chronotype reflects the interindividual variability in the phase of entrainment. Those with later chronotype typically prefer later times in the day for different activities such as sleep or meals. Later chronotype has been shown to be associated with metabolic syndrome, increased diabetes risk and poorer glycemic control in type 2 diabetes patients. In addition, “social jetlag”, a form of circadian misalignment due to a mismatch between social rhythms and the circadian clock, has been shown to be associated with insulin resistance. Other sleep disturbances (insufficient sleep, poor sleep quality and sleep apnea) have also been shown to affect glucose metabolism. In this study, we explored whether there was a relationship between chronotype, social jetlag and hemoglobin A1c (HbA1c) levels in prediabetes patients, independent of other sleep disturbances. A cross-sectional study was conducted at the Department of Family Medicine, Ramathibodi Hospital, Bangkok, from October 2014 to March 2016 in 1014 non-shift working adults with prediabetes. Mid-sleep time on free day adjusted for sleep debt (MSFsc) was used as an indicator of chronotype. Social jetlag was calculated based on the absolute difference between mid-sleep time on weekdays and weekends. The most recent HbA1c values and lipid levels were retrieved from clinical laboratory databases. Univariate analyses revealed that later MSFsc (p = 0.028) but not social jetlag (p = 0.48) was significantly associated with higher HbA1c levels. Multivariate linear regression analysis was applied to determine whether an independent association between MSFsc and HbA1c level existed. After adjusting for age, sex, alcohol use, body mass index (BMI), social jetlag, sleep duration, sleep quality and sleep apnea risk, later MSFsc was significantly associated with higher HbA1c level (B = 0.019, 95% CI: 0.00001, 0.038, p = 0.049). The effect size of one hour later MSFsc on HbA1c (standardized coefficient = 0.065) was approximately 74% of that of the effect of one unit (kg/m²) increase in BMI (standardized coefficient = 0.087). In summary, later chronotype is associated with higher HbA1c levels in patients with prediabetes, independent of social jetlag and other sleep disturbances. Further research regarding the potential role of chronotype in diabetes prevention should be explored.     

Temporal changes in floral resource availability and flower visitation in a butterfly

Foraging affects survival and reproductive success in animals, including flower-visiting insects. Plant-derived floral food resources (i.e. nectar and pollen) may be rapidly changing in space and time and pollinators may need to quickly switch to new resources. Butterflies are suitable model organisms to investigate foraging behaviour of insect pollinators, because they can be easily monitored under natural conditions. We studied flower visitation patterns in the Clouded Apollo butterfly Parnassius mnemosyne in relation to the abundance of available floral resources. We recorded flower visitation daily in individually marked butterflies, listed flowering species and estimated flower abundance categories every 3 days in a single meadow, during five consecutive flight periods. Butterflies visited 35 nectar plants from the 71 species available. Few nectar plants were frequently visited (visit ratios for the annually most visited species: 37–60%), many were scarcely visited and no visits were observed on several abundant species. Flower abundance and visit ratio varied among years and within flight periods. The number of visits increased with flower abundance in the seven most frequently visited plant species, but not in the occasionally visited ones. Beside their choosiness, Parnassius mnemosyne butterflies were able to adjust foraging behaviour to rapidly changing resource distributions. Diet selectivity in adults might increase the vulnerability of this species. However, visitation plasticity may mitigate the effect of the lack of some nectar plants, as complementary resources can be used as alternatives.     

Adenosine kinase deficiency is associated with developmental abnormalities and reduced transmethylation

Adenosine (Ado) kinase (ADK; ATP:Ado 5' phosphotransferase, EC 2.7.1.20) catalyzes the salvage synthesis of adenine monophosphate from Ado and ATP. In Arabidopsis, ADK is encoded by two cDNAs that share 89% nucleotide identity and are constitutively, yet differentially, expressed in leaves, stems, roots, and flowers. To investigate the role of ADK in plant metabolism, lines deficient in this enzyme activity have been created by sense and antisense expression of the ADK1 cDNA. The levels of ADK activity in these lines range from 7% to 70% of the activity found in wild-type Arabidopsis. Transgenic plants with 50% or more of the wild-type activity have a normal morphology. In contrast, plants with less than 10% ADK activity are small with rounded, wavy leaves and a compact, bushy appearance. Because of the lack of elongation of the primary shoot, the siliques extend in a cluster from the rosette. Fertility is decreased because the stamen filaments do not elongate normally; hypocotyl and root elongation are reduced also. The hydrolysis of S-adenosyl-L-homo-cysteine (SAH) produced from S-adenosyl-L-methionine (SAM)-dependent methylation reactions is a key source of Ado in plants. The lack of Ado salvage in the ADK-deficient lines leads to an increase in the SAH level and results in the inhibition of SAM-dependent transmethylation. There is a direct correlation between ADK activity and the level of methylesterified pectin in seed mucilage, as monitored by staining with ruthenium red, immunofluorescence labeling, or direct assay. These results indicate that Ado must be steadily removed by ADK to prevent feedback inhibition of SAH hydrolase and maintain SAM utilization and recycling.