Quantitative developmental analysis of homeotic changes in the inflorescence of Philodendron (Araceae)

Background and Aims: The inflorescence of Philodendron constitutes an interestingmorphological model to analyse the phenomenon of homeosis quantitativelyat the floral level. The specific goals of this study were (1)to characterize and quantify the range of homeotic transformationin Philodendron billietiae, and (2) to test the hypothesis thatthe nature of flowers surrounding atypical bisexual flowers(ABFs) channel the morphological potentialities of atypicalbisexual flowers. Methods: Inflorescences of P. billietiae at different stages of developmentwere observed using SEM. The number of appendices in male, femaleand sterile flowers were counted on 11 young inflorescences(5–6 flowers per inflorescence). The number of staminodesand carpels on ABFs were counted on 19 inflorescences (n = 143).These data were used for regression and ANOVA analyses. Results: There was an average of 4·1 stamens per male flower,9·8 carpels per female flower and 6·8 staminodesper sterile male flower. There was an average of 7·3floral appendices per atypical flower. Staminodes and carpelsare inserted on the same whorl in ABFs. A negative exponentialrelationship was found between the average number of staminodesand the number of carpels in ABFs. If only the ABFs consistingof less than six carpels are considered, there is a linear relationshipbetween the number of carpels and the average number of staminodes.The value of the slope of the regression equation indicatesthat on average, in P. billietiae, 1·36 carpels are replacedby one staminode. Conclusions: In P. billietiae, the number of appendages in female flowersimposes a constraint on the maximum total number of appendages(carpels and staminodes) that can develop on ABFs. The quantitativeanalyses indicate that the average number of different typesof floral appendages on an ABF and the number of organs involvedin a homeotic transformation are two independent phenomena.     

Arabidopsis (Brassicaceae) flower development and gynoecium patterning in wild type and Ettin mutants

Screening for mutations that alter flower development in Arabidopsis has led to the identification of two general types of genetic loci: those affecting meristem and organ identity, and those affecting growth and development independent of identity. ettin (ett) mutants belong to the latter class and exhibit pleiotropic phenotypes distinct from previously described Arabidopsis mutants. These phenotypes include increases in sepal and petal number, decreases in stamen number and anther locule number, and gross alteration of tissue patterning in the gynoecium. To determine when and how differences in ett floral meristems originate, flower development was compared between the wild type and ett mutants. ett floral meristems exhibit increases in abaxial sepal and petal primordia number without apparent increases in meristem size. Extra sepal and petal primordia develop into normal organs. In contrast, stamen and carpel primordia exhibit alterations in shape and form, subsequent to premature elongation of the terminal floral meristem. Phenotypes are allele-strength dependent. The stigma develops precociously and style differentiation is basally and abaxially misplaced in ett gynoecia. The data are discussed in the context of a model suggesting that two concentric boundaries specify the apical-basal pattern of gynoecium differentiation.     

Branch length mediates flower production and inflorescence architecture of Fouquieria splendens (ocotillo)

The capacity of individual branches to store water and fix carbon can have profound effects on inflorescence size and architecture, thus on floral display, pollination, and fecundity. Mixed regression was used to investigate the relation between branch length, a proxy for plant resources, and floral display of Fouquieria splendens (ocotillo), a woody, candelabraform shrub of wide distribution in arid North America. Long branches produced three times as many flowers as short branches, regardless of overall plant size. Long branches also had more complex panicles with more cymes and cyme types than short branches; thus, branch length also influenced inflorescence architecture. Within panicles, increasing the number of cymes by one unit added about two flowers, whereas increasing the number of cyme types by one unit added about 21 flowers. Because flower production is mediated by branch length, and because most plants have branches of various lengths, the floral display of individual plants necessarily encompasses a wide range of inflorescence size and structure.     

The fitness consequences of environmental sex reversal in fish: a quantitative review

Environmental sex reversal (ESR) occurs when environmental factors overpower genetic sex-determining factors. The phenomenon of ESR is observed widely in teleost species, where it can be induced by exposing developing fish to endocrine disrupting chemicals (EDCs). EDC-induced ESR has been exploited by the aquaculture industry, while ecological and evolutionary models are also beginning to elucidate the potential roles that sex-reversed individuals play in influencing population dynamics. However, how EDC exposure affects individual fitness remains relatively unknown. To date, many experimental studies have induced sex reversal in fish and measured fitness-as indicated by related traits such as size, survival and gonadal somatic index (GSI), but the reported results vary. Here, we meta-analytically combine the results of 78 studies of induced ESR to gain insight into the fitness of sex-reversed individuals. Overall, our results suggest that the fitness of fish exposed to EDCs is reduced at the time of exposure, with exposed individuals having a smaller size and likely a smaller GSI. Given a period of non-exposure, fish treated with EDCs can regain a size equal to those not exposed, although GSI remains compromised. Interestingly, survival does not appear to be affected by EDC treatment. The published reports that comprise our dataset are, however, based on captive fish and the general small size resulting from exposure is likely to lead to reduced survival in the wild. Additionally, reduced fitness-related parameters are likely to be due to exposure to EDCs rather than ESR itself. We suggest that theoretical models of ESR should account for the fitness-related effects that we report. Whilst we are able to shed light on the physical fitness of EDC-exposed fish, the behaviour of such individuals remains largely untested and should be the focus of future experimental manipulation.     

Positional Information,Positional Error,and Readout Precision in Morphogenesis: A Mathematical Framework

The concept of positional information is central to our understanding of how cells determine their location in a multicellular structure and thereby their developmental fates. Nevertheless, positional information has neither been defined mathematically nor quantified in a principled way. Here we provide an information-theoretic definition in the context of developmental gene expression patterns and examine the features of expression patterns that affect positional information quantitatively. We connect positional information with the concept of positional error and develop tools to directly measure information and error from experimental data. We illustrate our framework for the case of gap gene expression patterns in the early Drosophila embryo and show how information that is distributed among only four genes is sufficient to determine developmental fates with nearly single-cell resolution. Our approach can be generalized to a variety of different model systems; procedures and examples are discussed in detail.     

Sex ratio variation as a function of host size in Pseudacteon flies (Diptera: Phoridae), parasitoids of Solenopsis fire ants (Hymenoptera: Formicidae)

Some Pseudacteon (Diptera: Phoridae) flies are parasitoids of Solenopsis (Hymenoptera: Formicidae) ant workers in North and South America. Laboratory studies of sex allocation revealed a pattern of sex ratio variation as a function of host size, with more females arising from larger hosts. Environmental sex determination is a possible mechanism for the observed pattern, and examination of Pseudacteon life history reveals several traits assumed to be important in models predicting conditions under which environmental sex determination is favoured. Sex allocation patterns of Pseudacteon are compared with theoretical predictions and empirical data from better-studied hymenoptcran parasitoids that have haplodiploid sex determination. The pattern of sex ratio variation observed has important implications for biocontrol efforts of imported Solenopsis fire ants by the introduction of Pseudacteon parasitoids.     

Sex reversal and primary sex ratios in the common frog (Rana temporaria)

Sex reversal has been suggested to have profound implications for the evolution of sex chromosomes and population dynamics in ectotherms. Occasional sex reversal of genetic males has been hypothesized to prevent the evolutionary decay of nonrecombining Y chromosomes caused by the accumulation of deleterious mutations. At the same time, sex reversals can have a negative effect on population growth rate. Here, we studied phenotypic and genotypic sex in the common frog (Rana temporaria) in a subarctic environment, where strongly female‐biased sex ratios have raised the possibility of frequent sex reversals. We developed two novel sex‐linked microsatellite markers for the species and used them with a third, existing marker and a Bayesian modelling approach to study the occurrence of sex reversal and to determine primary sex ratios in egg clutches. Our results show that a significant proportion (0.09, 95% credible interval: 0.04–0.18) of adults that were genetically female expressed the male phenotype, but there was no evidence of sex reversal of genetic males that is required for counteracting the degeneration of Y chromosome. The primary sex ratios were mostly equal, but three clutches consisted only of genetic females and three others had a significant female bias. Reproduction of the sex‐reversed genetic females appears to create all‐female clutches potentially skewing the population level adult sex‐ratio consistent with field observations. However, based on a simulation model, such a bias is expected to be small and transient and thus does not fully explain the observed female‐bias in the field.     

Disentangling sex allocation in a viviparous reptile with temperature‐dependent sex determination: a multifactorial approach

Females are predicted to alter sex allocation when ecological, physiological and behavioural variables have different consequences on the fitness of male and female offspring. Traditionally, tests of sex allocation have examined single causative factors, often ignoring possible interactions between multiple factors. Here, we used a multifactorial approach to examine sex allocation in the viviparous skink, Niveoscincus ocellatus. We integrated a 16‐year observational field study with a manipulative laboratory experiment to explore whether the effects of the maternal thermal environment interact with the resources available to females for reproduction to affect sex allocation decisions. We found strong effects of temperature on sex allocation in the field, with females born in warm conditions and males in cold conditions; however, this was not replicated in the laboratory. In contrast, we found no effect of female resource availability on sex allocation, either independently, or in interaction with temperature. These results corresponded with an overall lack of an effect of resource availability on any of the life history traits that we predicted would mediate the benefits of differential sex allocation in this system, suggesting that selection for sex allocation in response to resource availability may be relatively weak. Combined, these results suggest that temperature may be the predominant factor driving sex allocation in this system.     

Effect of Aedes aegypti (Diptera:Culicidae) Age on Sex Ratios in Romanomermis culicivorax (Nematoda:Mermithidae)

First, second, third, early and late fourth-instar larvae, and pupae of Aedes aegypti were infected with Romanomerrais culicivorax and reared at 20, 25, and 30 C. An increase in the ratio of male to female nematodes was observed with increase in host age at the time of infection at each temperature. The number of pupal and late fourth-instar infections was low, but R. culicivorax continued to develop in adult A. aegypti. Since male nematodes were recovered from both male and female hosts infected as late fourth instars or pupae, the sex of the host did not influence the sex of the nematode.     

Environmental determination of sex in Apistogrammai (Cichlidae) and two other freshwater fishes (Teleostei)

Environmental sex determination by temperature could be revealed significantly in 33 Apistogramma-species and in Poecilia melanogaster . In some, but not all, Apistogramma-species pH also influences the sex ratio, whereas neither temperature nor pH affect the sex ratio of Pseudocrenilabrus multicolor victoriae . The sex in offspring of A. trifasciata is determined within a sensitive period of about 30 to at least 40 days after spawning.     

Sex determination in the larvae of the parasitic barnacle Heterosaccus lunatus: an experimental approach

Heterosaccus lunatus parasitizes the portunid crab, Charybdis callianassa, in Moreton bay, Queensland, Australia. With the host crabs maintained at 25 °C this sacculinid rhizocephalan released larval broods every 5-6 days. This study examines the effect of photoperiod on the sex proportions of successive larval broods of individual parasites. Parasitized host crabs were maintained individually in circulating 25 °C seawater within two light-tight boxes, each with a controlled light-dark cycle operating. Box A had an LD 16:8 cycle (summer) and Box B an LD 8:16 cycle (winter). Both boxes had five host crabs and when parasites released their larval broods the larvae were flushed out and retained in a filter. Each captured brood then had its larvae sized to determine the proportions of the two sexes present. H. lunatus is ideal for such study because the sex(es) of all the larval stages (four naupliar stages and the cyprid) can be unequivocally identified by size (length). The experiment was run over a 6-week period during March and April 2004 when a total of 86 larval broods were released and their sex proportion(s) determined. It is concluded that larval sex determination is effected by photoperiod, with females becoming dominant under the winter condition (LD 8:16) and males dominant under the summer condition (LD 16:8). All initial broods were male dominated, so for Box A broods the male preponderance was maintained, whilst for Box B broods there was the progressive changeover to female preponderance. Such environmental sex determination is unusual because it is oogenesis which is being controlled, whereby two ovum sizes can be produced, either singly or together, the larger being the male egg. Further work is now necessary to verify the likely hormonal control processes involved.     

Structure of the coxa and homeosis of legs in Nematocera (Insecta: Diptera)

Construction of the middle and hind coxae was investigated in 95 species of 30 nematoceran families. As a rule, the middle coxa contains a separate coxite, the mediocoxite, articulated to the sternal process. In most families, this coxite is movably articulated to the eucoxite and to the distocoxite area; the coxa is radially split twice. Some groups are characterized by a single split. The coxa in flies is restricted in its rotation owing to a partial junction either between the meron and the pleurite or between the eucoxite and the meropleurite. Hence the coxa is fastened to the thorax not only by two pivots (to the pleural ridge and the sternal process), but at the junction named above. Rotation is impossible without deformations; the role of hinges between coxites is to absorb deformations. This adaptive principle is confirmed by physical modelling. Middle coxae of limoniid tribes Eriopterini and Molophilini are compact, constructed by the template of hind coxae. On the contrary, hind coxae in all families of Mycetophiloidea and in Psychodidae s.l. are constructed like middle ones, with the separate mediocoxite, centrally suspended at the sternal process. These cases are considered as homeotic mutations, substituting one structure with a no less efficient one.     

The effect of photoperiod on flower formation in vitro in a quantitative short-day cultivar of Nicotiana tabacum

Superficial cell layers of a quantitative short-day tobacco plant ( Nicotiana tabacum L. cv. White Burley) were excised from different parts of the inflorescence (i.e. pedicels, branch internodes, rachises), and cultured in continuous darkness, continuous light or 8 h light/16 h dark daily. The flowering response in vitro of the different types of explants was investigated with respect to the effect of light on the post-evocation phases of the flowering process and explant commitment. Treatment effect was qualitatively and quantitatively influenced by explant origin. Three morphogenic features were observed: flower neoformation, caulogenesis and rhizogenesis (the latter on rachis explants only). Under all treatments, the highest flowering potential was shown by pedicels, while the highest vegetative potential was shown by rachises. Branch internodes showed an intermediate response, but with a tendency towards caulogenesis, which probably reflects their phylogenetic origin. Thus, opposite gradients of the neoformation of flowering and vegetative buds on explants were observed under all treatments. Pedicels formed new single flowers rather than inflorescences, while rachises regenerated mainly inflorescences. In darkness, flowering was limited mostly to pedicels. Vegetative bud formation was higher than floral bud regeneration in all types of explant. Continuous light enhanced the flowering response mostly in pedicel and branch internode explants. Short days enhanced flower bud formation in vitro on all types of explant. Results with respect to microsporogenesis, flower and inflorescence anomalies observed under darkness also seem to support the existence of a quantitative photoperiodic control on floral neoformation in vitro in this plant. These results suggest that in Nicotiana tabacum cv. White Burley in vivo floral induction, initiation and development are governed by the same photoperiodic requirements.     

Extrafloral nectaries in Philodendron (Araceae): distribution and structure

Extrafloral nectaries (EFNs) are involved in animal–plant interactions that lead to protection against herbivory. The presence of EFNs in Araceae is rare, besides Philodendron, there is report for only two other genera. With the aim to investigate the occurrence of EFNs in Philodendron and to describe the distribution patterns and structural organization of these glands, 75 Philodendron spp. were examined, 16 of which were selected for study by light microscopy. Three Homalomena spp. were also examined for EFNs, but these were not found. Philodendron martianum was employed as a model for additional study using scanning and transmission electron microscopy. The studied EFNs showed a high degree of structural similarity. They were present in the prophyll, leaf and spathe, becoming functional in young organs. In surface view, EFNs consisted of small areas and showed one or more stomata through which secretions were released. The secretory cells formed a globular region surrounded by ground parenchyma. In P. martianum, nectariferous parenchyma cells exhibited typical features of cells with high metabolism related to nectar secretion. These results allow us to infer that EFNs have a widespread occurrence in Philodendron, and they remain an exclusive character for this group. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180 , 229–240.