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The adaptive evolution and processing of sensory systems
Reconstructing the ancestral butterfly eye: focus on the opsins
Adriana D. Briscoe
The eyes of butterflies are remarkable, because they are nearly as diverse as the colors of wings. Much of eye diversity can be traced to alterations in the number, spectral properties and spatial distribution of the visual pigments. Visual pigments are light-sensitive molecules composed of an opsin protein and a chromophore. Most butterflies have eyes that contain visual pigments with a wavelength of peak absorbance, λmax, in the ultraviolet (UV, 300–400 nm), blue (B, 400–500 nm) and long wavelength (LW, 500–600 nm) part of the visible light spectrum, respectively, encoded by distinct UV, B and LW opsin genes. In the compound eye of butterflies, each individual ommatidium is composed of nine photoreceptor cells (R1–9) that generally express only one opsin mRNA per cell, although in some butterfly eyes there are ommatidial subtypes in which two opsins are co-expressed in the same photoreceptor cell. Based on a phylogenetic analysis of opsin cDNAs from the five butterfly families, Papilionidae, Pieridae, Nymphalidae, Lycaenidae and Riodinidae, and comparative analysis of opsin gene expression patterns from four of the five families, I propose a model for the patterning of the ancestral butterfly eye that is most closely aligned with the nymphalid eye. The R1 and R2 cells of the main retina expressed UV–UV-, UV–B- or B–B-absorbing visual pigments while the R3–9 cells expressed a LW-absorbing visual pigment. Visual systems of existing butterflies then underwent an adaptive expansion based on lineage-specific B and LW opsin gene multiplications and on alterations in the spatial expression of opsins within the eye. Understanding the molecular sophistication of butterfly eye complexity is a challenge that, if met, has broad biological implications.
eye evolution
photoreceptor
rhodopsin
visual pigment
The butterfly eye is a marvel of evolution. Butterfly vision, like that of other insects, is based on three major classes of photoreceptor, with peak sensitivity (λmax) in the ultraviolet (UV, 300–400 nm), blue (B, 400–500) and long wavelength (LW, 500–600 nm) part of the light spectrum. At the molecular level, these visual pigments are composed of a retinal-based chromophore (e.g. 11-cis-3-hydroxyretinal) (Smith and Goldsmith, 1990) attached by a Schiffbase linkage to an opsin protein. The spectral tuning of the visual pigment wavelength of peak absorbance, λmax, is achieved through the interaction of the chromophore with critical amino acid residues within the opsin. Changes in the polarity of amino acids in the chromophore-binding pocket of opsins, for example, affect the distribution of electrons in the π-electron system of the chromophore, producing a diversity of λmax values (Honig et al., 1976). In the case of the butterfly visual pigments, the three major spectral classes are encoded by ancient duplications, which produced distinct UV, B and LW opsin genes.
Unlike those of bees, the eyes of butterflies are anatomically and physiologically diverse (Peitsch et al., 1992; Briscoe and Chittka, 2001; Stavenga and Arikawa, 2006; Frentiu et al., 2007b; Frentiu et al., 2007a). This is due in part to lineage-specific opsin gene duplications, as well as to changes in the kind and distribution of lateral filtering pigments (Arikawa and Stavenga, 1997; Stavenga, 2002).
To understand this diversity, this review focuses on the eyes of four species of butterfly for which we have the most complete information, Danaus plexippus Linnaeus, Lycaena rubidus Behr, Pieris rapae crucivora Boisduval and Papilio xuthus Linnaeus, and whose eyes serve as prototypes for the butterfly families Nymphalidae, Lycaenidae, Pieridae and Papilionidae, respectively. Although the importance of non-opsin-based filtering pigments for modifying photoreceptor sensitivity to light is mentioned, it is not yet clear whether the filtering pigments in different families are homologous or not; therefore I have intentionally spotlighted the evolution of the opsins themselves because of the homologous relationship between opsin protein sequences between butterfly families and the spectral range of vision. By analyzing the physiologically determined photopigment absorbance spectra and the phylogenetic relationship of opsin gene sequences, overlaid with a character map of opsin expression patterns, I propose a novel model of the evolutionary events leading from an ancestral butterfly eye to the radiation of eye types observed today. And with an eye to the future, I emphasize a few fertile areas for further investigation.
Anatomy of the butterfly eye
Anatomically, the basic unit of the butterfly compound eye is the ommatidium. An ommatidium is composed of nine photoreceptor cells (R1–9) along with primary and secondary pigment cells. An ommatidium from the simplest eye, that of nymphalid butterflies, consists of two tiers composed, respectively, of eight photoreceptor cells (R1–8) that are elongated, and a tiny R9 cell that sits just above the basement membrane (Fig. 1A). Light passing through the cornea is focused by the crystalline cone onto the rhabdom. The fused rhabdom is composed of nine rhabdomeres, the microvillous membranes protruded by the individual photoreceptor cells, in which the visual pigments are embedded.
Diagram of an ommatidium and pattern of ultraviolet (UV), blue (B) and long wavelength (LW) opsin mRNA in the main retina of the monarch, Danaus plexippus (modified from Sauman et al., 2005). (A) Longitudinal view of an ommatidium. In nymphalids, the photoreceptor cell bodies that contribute to the fused rhabdom are organized into two tiers, composed of the R1–8 cells (tier I) and the R9 cell (tier II). Thick black lines, Ia. and Ib., indicate the approximate level from which the tangential sections of the R1–8 cells shown to the right were taken. c, cornea; cc, crystalline cone; n, photoreceptor cell nucleus; L, lamina; M, medulla. (B) Tangential views of ommatidial subtypes. Gray, purple and orange indicate the identity of the photoreceptor cells in which specific opsin mRNA expression is shown in the panels to the right. (C) Tangential section showing specific labeling of R1 and R2 cells with a digoxigenin-labeled antisense UV opsin riboprobe. Dashed circles indicate boundaries of identical individual ommatidia to those probed for B opsin mRNA in D. Scale bar, 25 μm. (D) Tangential section showing B opsin mRNA expression in an adjacent section to that shown in C. Dashed circles indicate identical ommatidia. Three subtypes of ommatidia are evident with respect to opsin expression in R1 and R2 photoreceptor cells: B–B, B–UV, UV–UV. Scale bar, 25 μm. (E) Tangential section showing LW opsin mRNA expression in all R3–8 cells (arrows). Scale bar, 25 μm.
UV and B opsin mRNA expression define three ommatidial subtypes in the main retina
The ommatidia of the main retina of the nymphalid eye are divided into three subtypes with respect to opsin mRNA expression, with R1 and R2 photoreceptor cells expressing UV–UV, UV–B or B–B, while the outer R3–8 cells express the LW opsin (Fig. 1B) (Briscoe et al., 2003). The monarch butterfly, Danaus plexippus, is a nymphalid butterfly whose eye typifies this pattern of opsin mRNA expression (Fig. 1C–E) (Sauman et al., 2005). The identity of the opsin expressed in the R9 cell of monarchs is unknown, but is likely to be the same as in another nymphalid Vanessa cardui, in which R9 expresses the LW opsin (Briscoe et al., 2003).
In addition to the ommatidia of the main retina, butterfly eyes also contain a dorsal rim area (DRA), like that found in other insects (Labhart and Meyer, 1999). In monarchs, the DRA R1–8 cells express the UV opsin exclusively (Sauman et al., 2005); these ommatidia contain rhabdoms whose microvilli are anatomically specialized to detect polarized skylight (Reppert et al., 2004). This monochromatic pattern of opsin expression in the DRA is important for efficient perception of polarized light, by avoiding interference with color information (Labhart and Meyer, 1999).
Spectral diversity of butterfly visual pigments
The wavelength of peak absorbance, λmax, of visual pigments can be directly measured using epi-microspectrophotometry, or derived from the spectral sensitivity curves of intracellular recordings of individual photoreceptor cells. In both cases, the experimental data are matched to an idealized rhodopsin template (Stavenga et al., 1993; Palacios et al., 1996), and a computational model is then applied to find the best estimate of λmax.
Based on such measurements, the spectral diversity of visual pigments in the eyes of butterflies is striking. As already alluded to, the eye of the monarch contains the smallest number of butterfly photopigments (P) with one UV (λmax=340 nm or P340), one B (P435) and one LW (P545) (Stalleicken et al., 2006; Frentiu et al., 2007a) (Fig. 2A). The lycaenid Lycaena rubidus eye contains four photopigments: one UV (P360), two B (P437 and P500) and one LW (P568) (Bernard and Remington, 1991) (Fig. 2B). The papilionid Papilio xuthus eye contains the largest number of butterfly photopigments so far described, with one UV (P360), one B (P460) and three LW (P515, P530 and P575) (Arikawa, 2003) (Fig. 2C). The eye of the pierid Pieris rapae contains four photopigments: one UV (P360), two B (P425 and P453) and one LW (P563) (Qiu and Arikawa, 2003a; Qiu and Arikawa, 2003b) (Fig. 2D).
Implications of B and LW opsin gene duplications in butterfly eye evolution
The opsin cDNAs encoding each of the visual pigments have been isolated (Kitamoto et al., 1998; Kitamoto et al., 2000; Wakakuwa et al., 2004; Arikawa et al., 2005; Sauman et al., 2005; Sison-Mangus et al., 2006; Frentiu et al., 2007a), and with these data a gene tree has been constructed for each of the three spectral classes of opsin: UV, B and LW (Fig. 3A–C, respectively). Species from all five butterfly families, Papilionidae (Papilio xuthus and P. glaucus Linnaeus), Pieridae (Pieris rapae), Nymphalidae (Danaus plexippus), Lycaenidae (Lycaena rubidus) and Riodinidae (Apodemia mormo Felder and Felder) have been included to illustrate the astonishing fact that independent duplications of either the B or LW opsin have occurred in each family, rendering the eyes of butterflies the most diverse yet characterized among insects.
Normalized absorbance spectra of butterfly visual pigments based upon the Bernard template (see Palacios et al., 1996) and wavelengths of peak absorbance (λmax) estimated from either microspectrophotometry or intracellular recordings. (A) The compound eye of the nymphalid Danaus plexippus contains three photopigments (P) with λmax values corresponding to 340 nm (gray), 435 nm (purple) and 545 nm (orange), respectively. Estimates of theλ max values of the UV (P340) and B (P435) photopigments are from electrophysiological recordings (Stalleicken et al., 2006) and for the LW (P545) photopigment, from microspectrophotometric measurements (Frentiu et al., 2007a). (B) The eye of the lycaenid Lycaena rubidus contains four photopigments: P360 (gray), P437 (purple), P500 (blue), P568 (orange). Estimates of theλ max values are from microspectrophotometric measurements (Bernard and Remington, 1991). (C) The eye of the papilionid Papilio xuthus contains five photopigments: P360, P460, P515, P530 and P575. Estimates of theλ max values are from the intracellular recordings (Arikawa, 2003). (D) The eye of the pierid Pieris rapae contains four photopigments: P360 (gray), P425 (violet), P453 (purple) and P563 (orange). Estimates of theλ max values are from intracellular recordings (Qiu and Arikawa, 2003a; Qiu and Arikawa, 2003b) (see also Wakakuwa et al., 2004; Arikawa et al., 2005).
For each of the butterfly family examples, only one UV opsin gene has been isolated, including from the riodinid Apodemia mormo, presented here for the first time (Fig. 3A). By contrast, duplicate B opsin genes have been isolated from P. rapae, encoding blue-absorbing (P453) and violet (V)-absorbing (P425) photopigments (Arikawa et al., 2005), and from L. rubidus, encoding blue-absorbing (P437) and blue–green-absorbing (P500) photopigments (Sison-Mangus et al., 2006). The bootstrap values of the relevant nodes and branching order of the tree, which nicely matches the current phylogeny of butterfly families based on independent molecular and morphological markers (see below) (Wahlberg et al., 2005), indicate that the duplication of the blue opsin gene in pierids and in lycaenids occurred independently.
The situation for the LW opsins, with respect to the frequency of gene duplication, is more complex. The eyes of pierids, nymphalids and lycaenids express only one LW opsin gene encoding photopigments of P563, P545 and P568, respectively (Wakakuwa et al., 2004; Sauman et al., 2005; Sison-Mangus et al., 2006), while the papilionid eye expresses three (P515, P530 and P575) (Kitamoto et al., 1998) and the riodinid eye expresses two LW opsin genes (P505 and P600) (Frentiu et al., 2007a) (Fig. 3C). To add more complexity, at least one additional near full-length LW opsin cDNA, PglRh4, for a total of four LW opsin genes, has been isolated from head tissue of Papilio glaucus (Briscoe, 1998; Briscoe, 2000) – an opsin that is not apparently expressed in the eye (A.D.B., unpublished data) (Lampel et al., 2005). The presence of two LW opsin genes in the genome of the silkmoth Bombyx mori, and the apparent close homology of the sphingid moth Manduca sexta opsin gene MANOP1 with one of them (Fig. 3C), suggests that M. sexta too may contain a second LW opsin, which is a homolog of the silkmoth larval `brain opsin' gene Boceropsin (Shimizu et al., 2001).
The fate of duplicate genes is typically classified into one of two categories: subfunctionalization (`division of labor'), in which one or both of the paralogs acquires a more restricted expression pattern and/or biological activity than the common ancestral gene, and neofunctionalization, in which the paralogs are free to acquire new expression patterns and/or functions (Force et al., 1999). Opsin duplication in butterflies can fall into either category (Briscoe, 2001). Importantly, duplication of butterfly opsin genes has led to the evolution of novel λmax values that ultimately impact on behavior (Kelber, 1999; Kelber and Pfaff, 1999; Kinoshita et al., 1999).
Subfunctionalization of B opsins in Pieris
In Pieris rapae, LW opsin expression is identical to that of the nymphalid eye in that its single LW opsin transcript is expressed in R3–8 (Wakakuwa et al., 2004). In addition, the expression of the UV opsin in Pieris also follows that of the nymphalid in that it is found in either UV–UV or UV–B combinations in R1 and R2. In what appears to be a complete departure from the nymphalid ground plan for the eye, however, the B–B ommatidial subtype has been completely replaced in P. rapae ventral eye by a V–V ommatidial class that is the product of a B opsin gene duplication (Fig. 4). Since the violet-absorbing visual pigment is encoded by a duplicate B opsin (Fig. 3B), and the pattern of B opsin expression in other butterflies considered so far is restricted to the R1 and R2 cells, the V–V class of ommatidium can be interpreted as an example of subfunctionalization of the ancestral B opsin domain of expression, into a more limited expression pattern.
Phylogenies of the UV, B and LW opsin genes of each of the five butterfly families including genes corresponding to the visual pigments shown in Fig. 2. The neighbor-joining method using 1st+2nd nucleotide positions and the Tamura-Nei model of evolution was used. Numbers represent the percentage of trees in which a particular node was recovered out of 500 replicates. The moths Manduca sexta (Sphingidae) and Bombyx mori (Bombycidae) are used as outgroups. (A) UV opsin gene tree. Opsin genes encoding the UV-absorbing visual pigments in Fig. 2 are indicated by bold gray. In each of the butterfly species shown, the UV visual pigment is encoded by a single-copy gene. The full-length UV opsin cDNA of A. mormo is newly presented in this study (GenBank accession no. AY587905). (B) B opsin gene tree. Opsin genes encoding the B-absorbing visual pigments in Fig. 2 are shown in bold purple. Duplications of the B opsin gene have been observed twice in independent butterfly lineages: once in the pierid P. rapae giving rise to the violet receptor (PrV) (light purple) (Arikawa et al., 2005), and once in the lycaenid L. rubidus, giving rise to a blue–green (B2)-sensitive photopigment (BRh2; blue) (Sison-Mangus et al., 2006). (C) LW opsin gene tree. Opsin genes encoding the LW-absorbing visual pigments shown in Fig. 2 are indicated by bold orange, yellow and green. LW opsin gene duplications have occurred in three of five butterfly families independently. Besides those in Papilio, LW opsin duplicate genes have been recovered from the eye of the riodinid Apodemia mormo and from the genomic DNA of the nymphalid Hermeuptychia hermes (not shown) (Frentiu et al., 2007a). Scale bars indicate substitutions per site.
Subfunctionalization and neofunctionalization of B opsins in Lycaena has led to a sexually dimorphic eye
The pattern of duplicate B opsin expression in Lycaena rubidus illustrates both neofunctionalization and subfunctionalization and leads to the situation in which male and female butterflies literally see the world through different eyes. In the compound eye of this animal, four opsin genes are expressed – one UV (UVRh), duplicate B, B1 (BRh1) and B2 (BRh2), and one LW (LWRh) opsin (Fig. 3). It is the dorsal eye of L. rubidus that is sexually dimorphic. Unlike the papilionid, pierid or nymphalid eye, in which R3–8 express one or more LW opsins, in L. rubidus males, R3–8 exclusively express the B1 (dark blue ovals) opsin, while in females, R3–8 co-express the B1 and LW (dark blue–orange ovals) opsins (Fig. 4), an example of neofunctionalization of the B1 opsin domain of expression. In addition, the R1 and R2 cells of the dorsal eye of both males and females express UV–UV almost exclusively, with a minor number of ommatidia expressing the UV–B1 or B1–B1 combination (Sison-Mangus et al., 2006). The highly territorial male L. rubidus (Bernard and Remington, 1991) probably use their dorsal eye for dichromatic color vision and the detection of moving objects, such as airborne males.
The ventral eye, by contrast, has a pattern of LW opsin expression that is more consistent with the other butterfly families examined: R3–8 of the ventral retina express the LW opsin. It is here, though, that subfunctionalization of the second blue opsin duplicate gene, B2 (light blue ovals, Fig. 4), is evident. The ventral retina contains six classes of ommatidia that differ according to the opsins expressed in R1 and R2: UV–UV, UV–B1, UV–B2, B1–B2, B1–B1 or B2–B2 (Fig. 4).
Intriguingly, while butterflies in the genus Papilio use duplicate LW opsins to see green (Kelber, 1999), the lycaenid Polyommatus icarus, uses its duplicate B2 opsin, in conjunction with its LW opsin, to see in the green part of the spectrum extending up to 560 nm (Sison-Mangus et al., 2008). This suggests that natural selection has hit upon alternative strategies for color vision in the green range in lycaenid and papilionid butterflies.
Subfunctionalization and neofunctionalization of LW opsin expression in Papilio
The basic pattern of photoreceptor cell morphology and opsin mRNA expression exemplified by the nymphalid eye is also complicated in the swallowtail butterfly Papilio by alterations in the shapes of the photoreceptor cells and by the expression of the three duplicate LW opsins. For example, the ommatidium of the papilionid eye is organized into three tiers rather than two: a distal tier in which the cell bodies of the R1–4 cells are the widest, a middle tier in which the cell bodies of the R5–8 cells are the widest, and a third tier in which the cell body of the tiny R9 cell is located (Fig. 5).
Ommatidial subtypes found in the dorsal rim area (DRA), dorsal and ventral retina of butterflies according to opsin mRNA expression in individual R1–8 photoreceptor cells (ovals). Photoreceptor cells are color coded according to the identically colored visual pigments and opsins shown in Figs 2 and 3 that are expressed in them. Question mark indicates that opsin expression is unknown in some parts of the eyes reviewed here. Boxes indicate the UV, B and LW opsin genes and lineage-specific duplicates present in each species and are organized into columns according to membership in the UV, B or LW opsin clades shown in Fig. 2. Note: the patterns of opsin mRNA expression in the riodinid Apodemia mormo are unknown and so are not included in this scheme. Left to right: D. plexippus DRA is composed of approximately 100 ommatidia organized into two to three rows that exclusively express the UV opsin (gray) in the R1–8 cells. Both the dorsal and ventral eye contain three types of ommatidia in which the R1 and R2 cells express UV–UV, UV–B or B–B opsin mRNAs and the R5–8 cells express a LW opsin (orange) (Sauman et al., 2005). P. rapae ventral eye contains four opsins and three ommatidial subtypes in which R5–8 express one LW opsin mRNA (orange) and R1 and R2 express UV–B, UV–UV or V–V (violet) opsin mRNA (Arikawa et al., 2005). It is not yet known whether other ommatidial subtypes are present in the dorsal retina or in the DRA. L. rubidus dorsal eye contains three opsins and is sexually dimorphic with respect to their expression. In males, R5–8 exclusively expresses the B1 (purple) opsin, while in females the B1 opsin is co-expressed with the LW (orange) opsin. The R1 and R2 cells of the dorsal eye are almost entirely UV–UV, with the R1 and R2 cells of a few ommatidia expressing UV–B1 and even fewer expressing B1–B1. The ventral eye of both males and females contains one additional opsin, B2, encoded by a duplicate blue opsin gene, BRh2. The R3–8 cells all express the LW opsin and R1 and R2 all express UV–B1, UV–UV, B1–B1, UV–B2, B1–B2 or B2–B2 (Sison-Mangus et al., 2006). P. xuthus contains a dorsal eye with four opsins and three distinct ommatidial subtypes: B–B (dark purple) in R1 and R2 and PxRh2 (orange) in R3–8; UV–B in R1 and R2, PxRh2 in R3 and R4, and PxRh3 (yellow) in R5–8; UV–UV (gray) in R1 and R2, PxRh2 in R3 and R4, and co-expressed PxRh2 and PxRh3 (orange–yellow). The ventral retina contains five opsins and the same three ommatidial subtypes except PxRh1 is co-expressed with PxRh2 (green–orange) in all R3 and R4 cells in the ventral eye. Squares indicate opsin genes expressed in identically colored photoreceptor cells.
In Papilio xuthus, the butterfly eye about which we know the most, three LW opsin genes (PxRh1, PxRh2 and PxRh3) are also expressed in the eye. In this butterfly, as in lycaenids, the main retina is divided into two parts: the dorsal eye and the ventral eye. Like nymphalids, the R1 and R2 cells of both the dorsal and ventral eye express UV–UV, UV–B or B–B opsin mRNAs. Unlike nymphalids, the R3 and R4 cells of the dorsal eye of P. xuthus express the PxRh2 opsin mRNA (orange ovals, Fig. 4), while in the ventral eye, the R3 and R4 cells co-express PxRh1 and PxRh2 (green–orange ovals, Fig. 4) (Kitamoto et al., 1998).
Strikingly, the expression of opsins in R5–8 is coordinated with the expression of opsins in R1 and R2 (Arikawa, 2003). In the dorsal eye, one subtype contains B–B in R1 and R2 and PxRh2 (orange ovals) in R5–8, another subtype contains UV–B in R1 and R2 and PxRh3 (yellow ovals) in R5–8 and a third subtype contains UV–UV in R1 and R2 and PxRh2 and PxRh3 (orange–yellow) in R5–8. The ventral eye contains the same three ommatidial subtypes as in the dorsal eye, except as noted above, the PxRh1 opsin is also co-expressed with the PxRh2 opsin in R3 and R4. Altogether, there are at least six classes of ommatidia with respect to opsin expression in the main retina of P. xuthus (Fig. 4).
Interestingly, the most phylogenetically basal of the P. xuthus LW opsin gene duplicates expressed in the eye, PxRh2 (Fig. 3), is also the one that has the widest expression in the R3–8 cells. And, like the LW opsin of V. cardui (Briscoe et al., 2003) and P. glaucus (Fig. 5K,L), it is also expressed in the R9 cell. This pattern is consistent with an ancestral function for PxRh2 and a subfunctionalization of the more recent duplicates, PxRh1 and PxRh3 (Briscoe, 2000), into a more limited domain of expression. It is also worth noting that PxRh3 exemplifies neofunctionalization, as it encodes P575, whose absorbance spectrum maximum is quite red shifted compared with PxRh2, which encodes P530 [see discussion in Briscoe (Briscoe, 2000)].
Diagram of an ommatidium and opsin mRNA expression in the swallowtail butterfly Papilio glaucus, newly presented here. Methods used for the in situ hybridizations are as detailed previously (Briscoe et al., 2003). (A) Longitudinal view of an ommatidium. In Papilio spp., the photoreceptor cell bodies that contribute to the fused rhabdom are organized into three tiers: I, II and III. c, cornea; cc, crystalline cone; n, photoreceptor nucleus; L, lamina; M, medulla. (B) Tangential views of an ommatidium. The most distal tier (I) contains the cell bodies of the R1 and 2 and R3 and 4 cells. The proximal tier (II) contains the cell bodies of the R5–8 cells, and the basal tier (III) contains the R9 cell body. Gray indicates the identity of the photoreceptor cells in which specific opsin mRNA expression is shown in the panels to the right. (C) Digoxigenin-labeled antisense riboprobe of UV (PglRh5) indicating the expression of this opsin mRNA in the distal tier of photoreceptor cells. This opsin is homologous to the UV opsin gene of P. xuthus, PxUV, shown in Fig. 2. (D) A semi-tangential section showing double labeling of R1 and R2 cells with antisense biotin-labeled UV (PglRh5) (brown) and digoxigenin-labeled B opsin mRNA (PglRh6) (blue). The latter is homologous to the blue (B) opsin gene of P. xuthus, PxB, shown in Fig. 2. Three subtypes of ommatidia are evident: UV–UV, UV–B and B–B. (E) UV opsin expression (red) in R1 and R2 cells using a rabbit anti-PglRh5 peptide antibody generated by the author (see Lampel et al., 2005) and visualized with a Cy3-conjugated goat anti-rabbit secondary antibody. Tangential section showing opsin staining in both the rhabdom (white arrow) and cytoplasm. Two subtypes of ommatidia are shown here, UV–UV and UV–?, although all three subtypes implied by D are present elsewhere in the section. (F) Tangential section of the ventral retina showing staining of all R3 and R4 cells with a digoxigenin-labeled antisense PglRh1 riboprobe. PglRh1 is homologous to P. xuthus PxRh1 shown in Fig. 2. (G) Tangential view of an adjacent section to F of the ventral retina showing staining of all R3 and R4 cells with digoxigenin-labeled antisense PglRh2 riboprobe, and indicating co-expression of PglRh1 and PglRh2 in these cells in the ventral retina. In the dorsal eye, only PglRh2 is present in the R3 and R4 cells (data not shown). (H) Longitudinal section of the eye indicating staining of the proximal (R5–8) tier of photoreceptor cells with a digoxigenin-labeled antisense riboprobe to PglRh3. (I) Tangential section showing strong staining of the R5–8 cells (arrow) of some ommatidia with a digoxigenin-labeled anti-sense riboprobe to PglRh3. (J) Tangential section showing strong staining of the R5–8 cells (arrow) of some ommatidia with a digoxigenin-labeled antisense riboprobe to PglRh2. (K) Dissociated ommatidium in which the R9 cell (arrow), but not the adjoining R5–8 cells, is clearly stained with a digoxigenin-labeled antisense riboprobe to PglRh2. This represents one (Type I) of three ommatidial subtypes defined by Arikawa (Arikawa, 2003). (L) Dissociated ommatidium in which both the R9 cell (arrow) and the R5–8 cells are clearly stained with a digoxigenin-labeled antisense riboprobe to PglRh2. This ommatidial subtype corresponds to either the Type II or Type III subtype (Arikawa, 2003).
Phylogeny of the butterfly families, character mapping of opsin mRNA expression patterns in individual photoreceptor cell subtypes, and proposed ancestral butterfly eye. Question mark indicates unknown opsin expression in the photoreceptor cells of the riodinid eye. (A) Character mapping of opsin mRNA expression in the R1 and R2 cells and inferred relative timing of opsin gene duplication events. Ovals indicate UV (gray) or B (purple, violet, blue) opsin expression. Boxes and ovals indicate the inferred ancestral states along specific branches of the butterfly family tree for both opsin genes and expression patterns. The R1 and R2 opsin expression patterns are not known for the Riodinidae. (B) Character mapping of opsin expression in the R3–8 cells. Ovals indicate photoreceptor cells expressing LW (orange, yellow or green) or B (purple) opsin mRNAs. Asterisk indicates ventral eye only. Red box indicates a LW opsin duplicate, the precursor to Apodemia mormo LWRh1, that is inferred to have arisen prior to the radiation of the lycaenid and riodinid families, and was subsequently lost in lycaenids. The R3–8 opsin expression patterns are not known for the Riodinidae.
Filtering pigments modify the sensitivity of photoreceptor cells to light
In addition to the visual pigments, all four butterfly eyes profiled here contain one or more combinations of yellow, orange or red filtering pigments (Arikawa et al., 1999b; Arikawa, 2003; Stavenga and Arikawa, 2006). Even more remarkable is that the eye of Pieris rapae crucivora is sexually dimorphic with respect to the expression of a blue-absorbing filtering pigment that is present only in the eye of males and is co-expressed with the violet opsin (Arikawa et al., 2005). Moreover, some butterfly eyes not reviewed here lack filtering pigments entirely (Briscoe and Bernard, 2005). Lateral filtering pigments coat the rhabdom, and filter short wavelength light, thus shifting the sensitivity of the visual pigments to the longer wavelengths. Examples of such an effect for red filtering pigments have been shown electrophysiologically in Heliconius erato (Bernhard et al., 1970; Swihart and Gordon, 1971; Struwe, 1972), Papilio xuthus (Arikawa et al., 1987; Arikawa et al., 1999a) and Pieris rapae crucivora (Qiu et al., 2002; Wakakuwa et al., 2004), and for the blue-absorbing filtering pigment of P. rapae (Arikawa et al., 2005).
The biological significance of filtering pigments is that while they do not expand the total visual range of the animal (this is dependent on the visual pigments themselves), they may expand the animal's color vision range. The red filtering pigments in the eyes of butterflies, however, do not appear to have a unified function. For instance, behavioral tests have shown that the nymphalid Heliconius erato uses a heterogeneously expressed red filtering pigment together with a single LW opsin to produce expanded color vision in the long wavelength range when foraging (Zaccardi et al., 2006), but the lycaenid Polyommatus icarus, which also has a heterogeneously expressed red filtering pigment in its eye, does not appear to have expanded color vision, at least when tested in the context of feeding (Sison-Mangus et al., 2008).
Providing a clearer role for filtering pigments in color vision will require further behavioral testing in a larger collection of butterfly species and, in some cases, such as the red receptor of Papilio, which co-expresses a red-filtering pigment with a red-absorbing visual pigment (Arikawa, 2003), it will be difficult to disentangle the impact of the filtering pigment on color vision from that of their coordinately expressed opsin.
Reconstructing the ancestral butterfly eye
The phylogeny of the butterflies provides a framework for reconstructing the pattern of opsin expression in the ancestral eye. Within the true butterflies (Papilionoidea) the current understanding of familial relationships is Papilionidae+{Pieridae+[Nymphalidae+ (Lycaenidae+Riodinidae)]} (Wahlberg et al., 2005), where papilionid and pierid butterflies represent the most basal lineages, and nymphalid, lycaenid and riodinid the most derived (Fig. 6). The inferred instances of lineage-specific opsin gene duplications (Fig. 3) together with the extant pattern of opsin mRNA expression (summarized in Fig. 4) can be mapped onto the butterfly family species tree to infer the most parsimonious ancestral state, using the sphingid moth Manduca sexta as an outgroup. Remarkably, the ancestral state is not represented by the most basal butterfly lineages but, instead, is manifested in the nymphalids.
Fig. 6A shows the most parsimonious reconstruction of the ancestral pattern of opsin mRNA expression in the R1 and R2 cells. Using this reconstruction, the ancestor of all butterfly eyes had merely one UV (gray box and ovals) and one B opsin (dark blue box and ovals) gene expressed in a non-overlapping fashion in the R1 and R2 cells – a pattern shared with the opsin expression in the main retina of sphingid moths (White et al., 2003) and bees (Spaethe and Briscoe, 2005; Wakakuwa et al., 2005) and consistent with the existence of only one UV and one B opsin gene in the genome of the silkmoth, Bombyx mori (Mita et al., 2004; Xia et al., 2004).
Subsequently, sometime after the divergence of the pierid from the papilionid and nymphalid+(lycaenid+riodinid) lineages, a B opsin gene duplication arose, presumably prior to the radiation of the Pierinae+Coliadinae subfamilies, since a homologue of the P. rapae (Pierinae) V opsin has been cloned from Colias philodice (Sison-Mangus et al., 2006), a species in one of the other major pierid subfamilies, Coliadinae. [The two more basal subfamilies, Pseudopontiinae and Dismorphiinae (Braby et al., 2006; Chew and Watt, 2006) have not yet been assayed for opsins.]
The pattern of ancestral LW opsin expression can be reconstructed similarly, and doing so suggests that the common ancestor of all butterfly eyes expressed a single LW opsin in the R3–8 cells (and also probably in the R9 cell) (Fig. 6B). Under this scenario, sometime after the split between the lineage leading to Papilio and the other butterfly families, two rounds of LW opsin gene duplication occurred, followed by subfunctionalization of the newer paralogs in Papilio. Intriguingly, the topology of the LW opsin gene tree also implies that the LW opsin duplicated prior to the radiation of the riodinid and lycaenid subfamilies, and that the homolog of the riodinid LWRh1 opsin was lost in the lineage leading to lycaenids. Subsequently, in the case of the highly divergent, sexually dimorphic pattern of opsin expression in the R3–8 cells of L. rubidus, the co-expression of a B opsin, B1 in R3–8, along with the LW opsin, can be viewed as an intermediate step along the evolution of a new function for B1 opsin in the dorsal eye of males (Sison-Mangus et al., 2006).
An eye to the future
One could imagine extending ancestral state reconstruction to other parts of the butterfly visual system to further understand the evolution of photoreceptor patterning, color vision, and even an integration of the evolution of the visual system with wing color. For instance, at present, the molecular identity, phylogenetic distribution and behavioral impact of filtering pigments in the eyes of butterflies are not well understood, but it seems likely that the ancestral butterfly eye contained heterogeneously expressed lateral filtering pigments, with secondary losses. Like the opsins, lateral filtering pigments may have evolved lineage-specific modifications and expression patterns that could have an impact on the color vision system. And, as has been pointed out, some of the pigments found in the brightly colored scales of butterfly wings are evolutionarily derived from eye pigments (Nijhout, 1991; Reed and Nagy, 2005). So it is possible that modifications of wing pigments co-evolved with the modifications of eye pigments.
Similarly, it is not known how the dorsal rim area ommatidium of the ancestral butterfly eye was patterned with respect to opsin expression. Unlike most moths, which are primarily nocturnal and may use polarized moonlight or starlight for navigation, butterflies are diurnally active and use the polarized skylight as a navigational cue (Froy et al., 2003; Reppert et al., 2004). If different opsins are expressed in the DRA ommatidia of different butterflies, this would imply that the polarized skylight cues used by butterflies are not uniform. In fact, it would not be surprising if this were the case, given the diversity of spectral classes of photoreceptors found in the DRA ommatidia of different insects (Labhart and Meyer, 1999) and the apparent ease with which domains of opsin expression in butterflies have been altered. A hint that the DRA ommatidia themselves may have evolved in Lepidoptera is evident in the observation that while the R1–8 cells in monarch DRA ommatidia express only the UV opsin (Sauman et al., 2005), in the hawkmoth, Manduca sexta, only some of the photoreceptor cells in the DRA ommatidia express the UV opsin (P357), while other photoreceptor cells express none of the other (B and L) opsins cloned (White et al., 2003).
Finally, alterations in opsin expression patterns beg the question of whether or not the neural circuitry necessary to process this information has evolved. At present, correlation between opsin expression and photoreceptor axon terminals has only been attempted in the swallowtail (Takemura et al., 2005; Takemura and Arikawa, 2006). The collateral morphology of the axons of the R5–8 cells of P. xuthus varies according to which LW opsin they express. But what about in other butterflies that do not have duplicated LW opsins?
In conclusion, it is clear that the butterfly eye is one of the most beautifully honed instruments of evolution. Bringing to bear molecular tools that are now becoming available for non-model organisms (e.g. genetic transformation, custom microarrays and RNA interference) will allow a deeper understanding of the biological events involved in butterfly eye diversity that may be applicable to a broader range of biological processes.
Thanks to Lisa Nagy for providing a supportive research environment for the Papilio glaucus work, Marilou Sison-Mangus for technical assistance, Matthew McHenry for suggesting character mapping and, especially, Steven Reppert for helpful comments on the manuscript. This work was supported by NSF grants IBN-0346765 and IOS-0646060.
© The Company of Biologists Limited 2008
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. And the bed was not scattered because bugs living in it strong held hands
Asthmatics Suffer From Complications From The Flu More Often
Asthmatics Suffer From Complications From The Flu More Often.
People with asthma visage rare risks from influenza, and a new report suggests far too few American asthma patients suffer the seasonal flu shot. "Asthmatics are at increased risk for complications from the flu," said one expert, Dr Len Horovitz, a pulmonary maestro at Lenox Hill Hospital in New York City aging womans face. "Exacerbations flare-ups of asthma are bourgeois with any viral infection, but the exacerbation from the flu is specifically severe".
The new study, led by Matthew Lozier of the US Centers for Disease Control and Prevention, looked at flu marksman uptake during the 2010-2011 flu season. The investigators found that only half of Americans with asthma got a flu sniper - a cast that was at least an improvement on the rate of 36 percent observed in the 2005-2006 flu season zetaclear. However, notwithstanding this increase, flu vaccination rates for people with asthma remain well below the federal government's Healthy People 2020 targets for flu vaccination: coverage of 80 percent for children ages 6 months to 17 years, and 90 percent for adults with asthma.
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Headache Accompanies Many Marines
Headache Accompanies Many Marines.
Active-duty Marines who tolerate a traumatic mastermind injury face significantly higher risk of post-traumatic stress disorder (PTSD), according to a new study. Other factors that gather the risk include severe pre-deployment symptoms of post-traumatic lay stress and high combat intensity, researchers report. But even after taking those factors and past brain harm into account, the study authors concluded that a new traumatic brain injury during a veteran's most brand-new deployment was the strongest predictor of PTSD symptoms after the deployment kannada. The study by Kate Yurgil, of the Veterans Affairs San Diego Healthcare System, and colleagues was published online Dec 11, 2013 in JAMA Psychiatry.
Each year, as many as 1,7 million Americans allow a damaging capacity injury, according to study background information. A traumatic brain injury occurs when the gourd violently impacts another object, or an object penetrates the skull, reaching the brain, according to the US National Institute of Neurological Disorders and Stroke coster oil ke fawaid pdf. War-related distressing brain injuries are common.
The use of improvised shaky devices (IEDs), rocket-propelled grenades and land mines in the Iraq and Afghanistan wars are the pipe contributors to deployment-related traumatic brain injuries today. More than half are caused by IEDs, the look authors noted. Previous research has suggested that experiencing a painful brain injury increases the risk of PTSD. The disorder can occur after someone experiences a shocking event.
Such events put the body and mind in a high-alert state because you feel that you or someone else is in danger. For some people, the mark related to the traumatic event doesn't go away. They may relive the affair over and over again, or they may avoid people or situations that remind them of the event. They may also feel jittery and always on alert, according to the US Department of Veterans Affairs. Many proletariat with traumatic brain injury also statement having symptoms of PTSD.
It's been unclear, however, whether the experience leading up to the injury caused the post-traumatic importance symptoms, or if the injury itself caused an increase in PTSD symptoms. The data came from a larger workroom following Marines over time. The current study looked at June 2008 to May 2012. The 1648 Marines included in the reflect on conducted interviews one month before a seven-month deployment to Iraq or Afghanistan, and a alternative interview three to six months after returning home.
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Scientists Are Exploring The Human Cerebral Cortex
Scientists Are Exploring The Human Cerebral Cortex.
Higher levels of self-professed clerical faith appear to be reflected in increased thickness of a key brain area, a uncharted study finds. Researchers at Columbia University in New York City found that the outer layer of the brain, known as the cortex, is thicker in some areas all people who place a lot of significance on religion rxlist box. The retreat involved 103 adults between the ages of 18 and 54 who were the children and grandchildren of both depressed deliberate over participants and those who were not depressed.
A team led by Lisa Miller analyzed how often the participants went to church and the destroy of importance they placed on religion. This assessment was made twice over the movement of five years neartohealth com. Using MRI technology, the cortical thickness of the participants' brains was also cadenced once.
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People consume more alcohol
People consume more alcohol.
Strong form alcohol control policies add up to a difference in efforts to help prevent binge drinking, a new study finds. Binge drinking - in a general way defined as having more than four to five alcoholic drinks in a two-hour days - is responsible for more than half of the 80000 alcohol-related deaths in the United States each year asthma ka gharelu ilaj. "If moonshine policies were a newly discovered gene, pill or vaccine, we'd be investing billions of dollars to attract them to market," study senior author Dr Tim Naimi, an associated professor of medicine at Boston University Schools of Medicine and attending doctor at Boston Medical Center (BMC), said in a BMC news release.
Naimi and his colleagues gave scores to states based on their implementation of 29 booze control policies. States with higher scheme scores were one-fourth as likely as those with lower scores to have binge drinking rates in the top 25 percent of states fav-store.net. This was realistic even after the researchers accounted for a variety of factors associated with hooch consumption, such as age, sex, race, income, geographic region, urban-rural differences, and levels of control and alcohol enforcement personnel.
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If the supporters of Ugwuonye had hoped for the granting of bail or some sort of a reprise for him today they were to be hugely disappointed.
At the Gudu High Court in the FCT, he along with two other co-defendants Messers Jekwu and Emmanuel were arraigned and ordered to be still held in prison till a final decision on their bail would be reached Related: Second Judge Withdraws From Okorocha's Certificate Of Return Suit .
That decision would be reached on January the 10th, 2019.
Ironically the lawyer for David Aiyedogbon in the murder trial is no other than Ifeanyi Ejiofor.
Once upon a time both he and Ugwuonye were Nnamdi Kalu's lawyers and one has to question his state of mind having once stood side by side with Ugwuonye in a common cause, and now trying to find him guilty and possibly send him to the gallows.
Regardless of friendship, justice i believe should prevail, no sacred cows.
This case will definitely be interesting, fingers crossed!


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Incredible Easter deals for all the family at Plymouth's top attractions
There are almost too many options to choose from
Charlotte GoslingCommercial Editor
Easter is a fantastic holiday to spend time in Plymouth, with the countryside on your doorstep, beaches galore just a stone's throw away and incredible attractions for all the family.
From action-packed afternoons filled with sporty assault course to period homes steeped in history and places to visit your favourite animals, Plymouth truly has it all.
If you're looking for some organised entertainment and want a prearranged event, check out Plymouth Live's list to the very best places to visit for a quality yet affordable day out with the family this holiday.
Super Tramp
(Image: PlymouthLive)
This Plymstock attraction boasts more than 20 trampolines, or you can try a slam dunk on the basketball court, test the Matrix with wall running, or work as a team with a game of dodgeball. The Airbag is also a great way to practise your moves in a safe environment.
Enjoy a family bounce session at the fun park for just £10 per bouncer. This is made for children aged 3-12 years and their parents/guardians who are 18 years and over. With every under five child, one adult can go free of charge to supervise during the session.
Click here for directions and visitor reviews.
Plymouth Arts Cinema
Inside the new arts cinema (Image: Jon Bishop)
Plymouth Arts Centre provides a place to enjoy contemporary art, an independent cinema and creative learning. It is based in the City Centre, at the Plymouth College of Art main campus but first opened on the Barbican in 1947. The best part is, despite a standard ticket being £9 (or a matinée £7), those 25 and under can enjoy the unique experience for just £4.
The Climbing Hangar
All the walls in The Climbing Hangar are colour coded so novices climb next to elite clamberers (Image: Penny Cross)
The Climbing Hangar is all about having fun and it offers loads of ways to find your happy at the wall. It has 13,000 square feet of climbing, training, gym and yoga space. The bouldering walls are home to absolute beginners and world champions; whoever you are, the experts will help you find your place. Adults can have fun for £8 and junior climbers - those under 14 years old - can get involved for just £8.
This is the UK's biggest aquarium with more than 4000 animals for visitors to enjoy. Children can even choose to sleep with the sharks in a unique sleepover experience by the big tank. By booking online you can save yourself 10 per cent and it comes with a complimentary annual return pass.
Pennywell Farm (Image: Richard Austin)
Pennywell Farm is a multi-award-winning attraction and the largest farm attraction in the South West. There is a different hands-on activity, show or display every half hour and four farm rides included in the admission price. Pennywell is also the home of the world-famous Pennywell miniature pigs. There is a licensed restaurant, acres of free parking and free dog kennels provided for four-legged visitors. Pennywell is open seven days a week and the perfect part is children under three can enter for free and those between the ages of four and 16 can enter for £12.95.
Saltram House
Saltram House is a George II era mansion located in Plympton and it offers a great place for walking, jogging, cycling and picnics. Here you can enjoy the famous Park Cafe, shop and visit the house as well as the garden with orangery, castle folly and chapel tea room. One adult and two children can head along to Saltram and enjoy the house and grounds for just £18.30.
Smeaton's Tower
This is the centrepiece of Plymouth's Hoe. Smeaton's Tower is one of the city's most recognisable and well-loved landmarks and you and the family can climb to the top to see the very best views in the city. The lighthouse stands at 72 feet high thanks to the original 18th century ladders. Adults can climb the tower for just £4 and children under 16 pay just £2 with under five's free.
Mayflower Museum
The museum explores the area's seafaring history and the Mayflower's 1620 voyage and includes a model ship. Set over three floors, you can immerse yourself in the city's history. Your journey of discovery begins by stepping onto the balcony on the top floor to enjoy a fantastic panoramic view of todays' busy Barbican. The best part is tickets are cheap with adult entry at just £3 and children between five and 16 costing just £1.50 each. Or save and buy a family ticket at just £8.
(Image: Paul Slater)
Grab the family and head to watch the latest blockbuster this holiday. With plenty of fantastic family films coming out over the Easter break it's important to get the best price for you and your family. If you're heading in on a Tuesday or Wednesday be sure to take advantage of Compare the Market's Meerkat Movies and Meals. All you have to do is take out insurance of any kind through them and you'll receive buy one get one free on films all year round. There are also Mini Mornings at Vue which offer kid's films at low prices. On Super Mondays, you can also enjoy big screen entertainment for a discounted price, all you have to do is sign up via the website, here.
Here you will find and indoor play and party venue providing children and their families with the opportunity to explore, learn, play and have fun. Play facilities include large climbing frames, slides, ball pools, inflatable's as well as dedicated toddler zones. There is also a café where all food is freshly prepared by on-site chefs. Babies can enter for just £1.50, one year old are £5, two to four year olds cost £7.25 and five to 12 year olds can enjoy the fun for £8.25 with all adults entering for free.
Dartmoor Zoological Park
Benjamin Mee and his family were the inspiration for the movie 'We Bought A Zoo' (Image: APEX)
This small and friendly wildlife park sits on the edge of Dartmoor and featured on a BBC TV series as well as in a Hollywood movie called We Bought A Zoo starring Matt Damon. This zoo truly has something for everyone and runs British Sign Language tours as well as close encounter sessions. Over the Easter holidays and beyond, those who are three years old and under can enter for free as well as those who are the carer for a disabled guest. A family of four costs just £38 and children under 15 can enter for just £8.95.
Clip 'n Climb Plymouth
With 44 indoor climbing walls, Clip 'n Climb Plymouth promises an exciting day out full of empowering, action-packed challenges for everybody. Test your resolve and determination on Orbital, reach dizzying heights on the UK's tallest Big Cheese, balance your way up the Stairway To Heaven or reach for the stars on Astroball – these are just a few of the challenges at Clip 'n Climb Plymouth. Family Tickets are now available saving as much as £10 for a family of four and saving £7.50 for a family of three.
Buckland Abbey
The Abbey is part museum, part house, and filled with treasures. There's no mistaking the magnificence of the Great Barn, which has remained virtually unchanged since it was built all those centuries ago. When you visit Buckland, you follow over 700 years of footsteps; from the Cistercians who built the Abbey and farmed the estate, to seafarers Grenville and Drake who changed the shape of the house and the fate of the country. A family of two adults can enjoy a day out for £30.50 or a family with one adult pay just £18.30.
Clay Art - Craft Centre
This charming pottery painting studio located on the historic Barbican, Plymouth is a friendly hub for you and your children to explore and unlock your creativity. An enormous range of quality products are available to choose from so that you can unleash your arty side. For a gift with a personal touch or a keepsake that lasts forever, this is the place to go. Prices start at £8.50 this is for a small piece, for example an egg cup or a small animal. For a basic size mug or plate £11 basic cereal bowl or similar £12 then moneyboxes start from £14 - £25. Larger items can cost from £25 - £50.
Plymouth Pavilions
Plymouth Pavilions (Image: Penny Cross / Plymouth Live)
Head along to this well-loved venue for leisure and entertainment. It includes an arena for comedy and music acts, plus an ice rink and a café. Head along for unforgettable fun with all the family whether that be to see a show, go ice skating or simply pop by for a bite to eat. Make the most of the family skate option at the ice rink where families of two adults and two children or one adult and three children can skate for £30.40.
Putters on The Hoe
Putters On The Hoe (Image: Plymouth City Council)
Putters On The Hoe is situated in Hoe Park on the top of Plymouth Hoe. This is an 18-hole grass putting course that is great for all the family and everyone of all ages and experience. You can relax before and after with snacks, hot/cold drinks and ice creams that are available to purchase from the Putters log cabin. Adults can play for just £2 and under 12s are priced at £1.50.
The Mountbatten Centre
Mountbatten Centre
The Mount Batten Watersports and Activities Centre is one of the foremost outdoor education centres in the South West – the centre has a reputation for excellence, professionalism and operates to the highest of standards. On offer are multi-activity courses, family fun days as well as RYA Instructor courses. The Mount Batten Centre also provides you with the perfect waterfront location for wining and dining as it provides stunning views over the Plymouth Cattewater. Head along to a Multi Adventure Day (MAD) for children between 7-11 years old can be enjoyed for just £36.95 per person or two can be bought for £69.95.
Barbican Theatre
Here theatre and dance will fire you up and get them excited about making, creating and exploring new moves and disciplines. The theatre is inclusive and work with anyone who wants to join them to make outstanding theatre and dance. You can get involved as a participant or as an audience member in work that inspires, challenges, entertains and educates.
(Image: DevonLive)
A superb landscaped park, with fallow deer, woodland and coastal walks as well as marvellous views, makes Mount Edgcumbe one of the most spectacular attractions in the West Country at any time of the year. The kids will love Segway-ing around the estate and taking in the stunning coastal views as they go. Or they can challenge themselves with babyyaga circus skills.
Plymouth Ski and Snowboard Centre
With three slopes to choose from, including the longest artificial slope in the South West, this is the perfect place to practise and learn new wintery skills. There is also a toboggan run and Sno-tube rides, plus a restaurant/bar. The artificial ski slopes at Alpine Park, just off the A38, are covered in perma-snow, so no matter how bright and sunny the weather, you're set for snowy joy.
The centre runs holiday clubs which provide a full day of learning and developing their chosen discipline, and your kids (ages 5-15yrs) are guaranteed to have loads of fun, get plenty of exercise and make new friends. Or enjoy a slide & ride experience (including sno-tubing and two toboggan rides) for £13 each.
(Image: John Allen)
The Plymouth Life Centre is a leisure centre in Central Park, Plymouth run by Everyone Active in partnership. This massive fitness centre has not only an Olympic 50m swimming pool but also a 12-court sports hall and an eight-rink indoor bowls hall. You can also take up a new hobby and try the exciting climbing wall. There are Easter holiday climbing taster sessions for just £5 per person between April 8 and 19.
Plympton Swimming Pool and Gym
(Image: Lucy Blake)
Plympton Pool is a 25m, four lane pool offering swimming lessons, family swims, lane swimming and aqua aerobic classes. It is set in an idyllic location overlooking the picturesque verdant pastures of the neighbouring park and cricket pitch. You can also enjoy the brand-new 30 station gym that features all the latest equipment and is sure to help you achieve your fitness goals, whatever they may be. Kids can enjoy a free taster session in the pool with an instructor this Easter.


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A bicentennial book is being compiled to coincide with the 200th anniversary of the founding of Perry County. The book will contain a history of the county from 1921 to 2021, alongside the histories of businesses, schools, churches, cemeteries, civic/social/fraternal organizations, historic sites and farms that have had a lasting impact in the county. Residents and businesses are invited to email history submissions to percohistsoc@yahoo.com; mail submissions to P.O. Box 97, Perryville, Missouri, 63775; or drop submissions off at 108 S. Progress Drive in Perryville. All submissions are due by Thursday, August 1. For information on submission format, contact (573) 768-2470.
BG's Old Tyme Deli
BG's Old Tyme Deli held their grand reopening June 1, commemorated with a ribbon-cutting ceremony, Happy Hour and dinner specials, giveaways and the launch of a new loyalty rewards program. The restaurant has implemented many changes, including a nonsmoking facility, remodeled interior and Sunday hours of business.
The Shipyard Music Festival
The Shipyard Music Festival is back at Ivers Square in Cape Girardeau for its second year, September 27 and 28, 2019. This year, the festival will feature 11 bands, co-headlined by Jukebox the Ghost and Colony House. The festival is presented by the Isle Casino Cape Girardeau and sponsored by United Airlines, River Radio, Drury Hotels, Saint Francis Healthcare System, Visit Cape, Bud Light, The Bank of Missouri and La Croix Church. Crew partners include mary jane burgers + brew, the Southeast Missourian, Horizon Screen Printing and Promotional Products, and Schaefer's Electrical Enclosures. Community partners include The Scout, Hudson Chiropractic, Cenet, Community Improvement District, Southeast Missouri State University and Shine Medical Aesthetics. Tickets are on sale at shipyardfest.com.
Marla Mills, executive director of Old Town Cape, announced plans to leave the position at the end of the summer. During Mills' 13 years as executive director of the organization, a casino and hotel opened in downtown Cape Girardeau. Other notable accomplishments include the development of the Cape Riverfront Market, facilitation of community and business workshops, creation of downtown Cape Girardeau Community Improvement District, development of a downtown tax increment financing district, Great American Main Street Award (2015), Broadway and Main Street streetscape improvements including a public restroom, Partners in Preservation grant award of more than $127,000 to revitalize Ivers Square and installation of a United States Colored Troop Memorial statue on its grounds. Old Town Cape is launching a national search for a new director, and Mills will assist in the transition.
SEMO Alliance for Disability Independence (SADI)
Maryann Gudermuth, founder and executive director of the SEMO Alliance for Disability Independence (SADI), retired May 31. Gudermuth founded SADI in Cape Girardeau in the late 1980s, securing funding for the organization in 1993. Donna Thompson takes Gudermuth's place as executive director of the organization.
Standard Democrat
Mike Jensen, publisher of the Standard Democrat in Sikeston, Missouri, retired May 31. Jensen spent nearly 50 years in the newspaper business and in 1972 helped launch the Bulletin Journal newspaper in Cape Girardeau.
The Southeast Missouri Food Bank
The Southeast Missouri Food Bank was named one of the "40 best food banks in America" in an article recently published by MSN.com. The article listed the Sikeston, Missouri-based food bank as No. 22 on its top 40 list.
Top of the Marq
Top of the Marq on the seventh floor of the historic Marquette Tower will offer panoramic views of downtown Cape Girardeau along with a cocktail bar and a "social" menu designed to appeal to a variety of tastes. It is projected to open in early July.
Information summarized from Southeast Missourian business articles and contributions from B Magazine readers. To contribute business briefs for the next issue of B Magazine, email mpohlman@rustmedia.com.


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Home / Entertainment / Angelina Jolie wins London by her elegance and refined taste
Angelina Jolie wins London by her elegance and refined taste
in Entertainment 27.11.2014
Hollywood actress Angelina Jolie appeared at the premiere of her film "Unbroken" in mind-blowing dress.
Famous Hollywood actress Angelina Jolie won her fans not only by her outfit in which she appeared at the premiere of "Unbroken" in London. The star accompanied her team as a director during the entire promotional tour, joinfo.ua informs.
It is reported that Angelina Jolie arrived to the event without her husband Brad Pitt. Although, in the media there is some information that there is discord and disagreement between the spouses.
But even minor problems did not prevent the star to be traditionally elegant. Fashion critics once said that this Jolie's dress was brilliant. The actress was dressed in a white cape by British designers Ralph & Russo.
Earlier, Angelina Jolie made a statement that she was quitting acting. From mow on her fans will not see her in the movies. The last work of the Hollywood star will be the role of the Egyptian queen Cleopatra. The actress has decided to devote herself to directing.
As previously reported, Angelina Jolie refused to invite Kim Kardashian to the premiere of "Unbroken". Although Kim denies this information.
As it is known, Jolie and Kardashian started their "competition" when the latter had lowered the actress in the list of popular show business mothers. Although the "parties" do not admit opposition between them.


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Good Omens review: David Tennant and Michael Sheen steal the show in this devilish delight
Adam StarkeyFriday 31 May 2019 9:54 am
David Tennant and Michael Sheen are a winning combo (Picture: Amazon Prime Video)
Good Omens searches for the sweet spot between heaven and hell in a countdown chase to the end of the world, where distinctly British wit collides with barking fantasy.
Based on the 1990 novel by Terry Pratchett and Neil Gaiman, this six-part miniseries carries an enormous weight both in its godly narrative and the show's origins. Adapted by Gaiman shortly after Pratchett's death back in 2015, following troubled movie plans for years, this show represents the culmination of much emotional strain and an arduous development cycle.
It's the biggest compliment to Good Omens that none of this feels like the case. This is a light, skippy, and silly dance with angels and demons, which bounds along with the creative zip and fantastical flight both authors are synonymous with.
The show revolves around angel Aziraphale (Michael Sheen) and demon Crowley (David Tennant) who, after becoming fond of living among humans on Earth from the dawn of creation, try to prevent the apocalyptic arrival of the antichrist ending life as we know it.
Aziraphale and Crowley form an unlikely bond between heaven and hell (Picture: Amazon)
While a countdown to the end of the world might scream urgency, Good Omens is lethargic and playful in tone. There's an unconventional rhythm to the show as it jumps back and forth through time, delivering comedy sketch-like scenarios one minute and chasing down the antichrist the next. It's not always successful, with some of the tangents feeling unnecessary or bloated, but the pacing is more rewarding the further you roll around in, and embrace, its obscurities.
That's because there's always an excellent cameo or surprise appearance lurking behind every corner. Good Omens has a huge cast list, ranging from lingering voices like Frances McDormand as the voice of God, Benedict Cumberbatch as Satan, or brief memorable scenes featuring League Of Gentlemen alumni Steve Pemberton, Reece Shearsmith, and Mark Gatiss. It makes the show feel like a toy box, constantly throwing new tricks at the audience which help you forgive the occasional unwieldy focus and misfires.
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Holding this all together is the binding, hysterical glue of Michael Sheen and David Tennant, who form one of the most enjoyable on-screen partnerships you'll find all year. The latter especially achieves a career-best as the nonchalant, swaggering demon – who is somewhere between Ozzy Osbourne, Bill Nighy and Marvel's Ghost Rider. They both clearly relish the witty script, and the whole show soars to its best whenever they're knocking banter between them.
While Good Omens would benefit from cutting back on less engaging story strands, like Jack Whitehall's Newton Pulsifer or antichrist Adam Young's childhood, this miniseries is a refreshingly whimsical concoction of British humour, infectious imagination, and old fashioned charms. With so much of fantasy on television embracing the darkness, Good Omens is a delectable counterweight bursting with a lust for life.
Good Omens is available to stream on Amazon Prime now.
MORE: Good Omens cast, trailer and when is it released on Amazon Prime?
MORE: Good Omens' trailer finally drops as David Tennant and Michael Sheen prepare for the end of the world
AmazonDavid TennantMichael Sheen
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As it is often said, context is everything.
"Brooklyn: The Musical" has a backstory that nearly overshadows the show itself. Its creators, Mark Schoenfeld and Barri McPherson, once collaborated decades ago before going seperate ways. More recently, McPherson, who had a comfortable life in New England, came across Schoenfeld, then a homeless street musician in Brooklyn. She took him in, and inspired by his tough life, they wrote what would become this musical.
After opening in Colorado, "Brooklyn" had a nearly full year on Broadway – October 2004 to June 2005. New York critics were not kind, but Kathleen Clarke Horrigan of Indy's Footlite Musicals saw it during its final month and fell in love. After years of hunting for a way to bring the musical to Indiana, she finally has "Brooklyn" occupying the Footlite stage.
For the most part, this show is the play-within-the-play about a Parisian girl, "Brooklyn," named for the home of the American father she never knew. After losing her mother (played by Page Brown), Brooklyn (Shelbi Berry) eventually makes her way to New York as a famous singer, with one unfinished song that only her real dad would know. Local diva Paradice (Kendra Randle) is not amused and wants this French upstart off her turf. Brooklyn accepts Paradice's challenge for a winner-take-all sing-off in hopes that this will aid her quest. But when she finds her father (Donny Torres) and learns his truth, will a happy ending to this tale be possible?
I'm leaving out a lot of details, of course, so you can discover them yourself. Dwelling on them would ruin the overall fantasia effect of the story, anyway. In the end, we truly learn who this story is about and for, which then sets the "fairy tale" as a whole in a clearer light.
The issue of homelessness permeates this story and production, but – as is true in everyday conversations – it is not directly addressed. This show won't preach to you, but does present these people's humanity, the "Heart Behind These Hands," and clues to what can bring a person down to life under a bridge. This production is also helping raise awareness and funds for the local Coalition for Homelessness Intervention and Prevention (www.chipindy.org).
Jones is a wonderful narrator with sweet voice and charisma to spare. Beautiful Berry and sassy Randle make an exellent sweet-sour yin-yang. Brown is angelic (literally) and Torres brings all the layers of his complex character. In other words, these "weeds" are a pitch-perfect bouquet of talent.
Also impressive is the look and atmosphere of the stage set by Stephen Matters, like a gritty set for "Rent" gone to seed, complete with lights and sounds (but thankfully no smells) to make you feel almost a bit unsafe. Costumes (by Curt Pickard) and props are marvels of recycling and improvisation with discarded everyday objects, oddly adding to the whimsy of some scenes.
Combine these elements with backing street people (Rayanna Bibbs, Tristan Bustos, Amy Douglas and Michael Davis) and an on-stage band led by Linda Parr, and you have one of those musicals that is as much an experience as a show. Don't be surprised if you find yourself humming an "Unfinished Lullaby" or have the words "With our tears, we water roses" tattooed to your memory.
This rare gem of an almost-forgotten musical has performances today through Sunday and Jan. 18-21 at 1847 N. Alabama. Call 317-926-6630 or visit www.footlite.org.


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Wolf King – The Path of Wrath (Review)
This is the second album from US blackened hardcore band Wolf King.
2018's Loyal to the Soil was an enjoyable slice of vicious nastiness, so I knew that checking out The Path of Wrath was definitely on the cards.
Combining dark, violent hardcore with an even stronger blackened streak, elements of sludge metal intensity, and even a touch of death metal and doom, the Wolf King of 2021 is a darker and filthier incarnation than it was previously. In fact, on The Path of Wrath it sounds as if Wolf King have truly taken their black metal influences to heart. If Loyal to the Soil was the sound of a hardcore band with a blackened coating, then The Path of Wrath is the sound of a black metal band with hardcore elements sewn into its walking corpse. This is blackened hardcore that's heavy on the blackened.
The Path of Wrath is a longer album than its predecessor, (53 minutes vs 43 minutes), but makes up for this by having a greater breadth of material than the band's debut album. As Wolf King have increased the blackened components in their sound, there's consequently more room for the exploration of atmosphere and old-school melodic black and death metal workouts. Both of which, they do rather well, (I especially like the At the Gates-isms that I can hear). The savagery of old is still here though; the band are even more ferocious actually, and their sound is rawer overall.
Wolf King's new album is a darker and well-developed expression of blackened hardcore. The band have doubled down on their extreme metal influences and created something that has their personality stamped on it more than their last album did. A strong, confident step forward into the shadows, The Path of Wrath is a very strong release.
Author wonderboxmetalPosted on March 3, 2021 Categories HardcoreTags At The Gates, Black metal, Blackened Hardcore, Death Metal, Doom, Hardcore, Melodic Black Metal, Melodic Death Metal, Metal, Sludge, Sludge metal, Wolf King
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(RT) – A baroque-styled gilded loo for university management in Russia's Urals region have raised a few eyebrows after images were leaked on social media. The eccentric toilet appeared overly posh, compared to the ones that students use.
Ornate carvings in gold-plated walls, shiny flooring and decorated ceiling, cushions and elegant lamps do not exclusively belong to the Versailles rooms, for they apparently can fit in with the university restroom in the Urals.
Commentators picked up the mood, ridiculing the university officials for their bad, or is grand taste, pointing to plane white latrines or cheap toilet paper, which obviously don't fit in with the bathroom's splendour.
The extravagant design for the management restroom is in stark contrast with the austere toilets for students and the rest of the university staff.
However, the university press office washed its hands of the matter, claiming it didn't spend "a ruble" on the makeover. The school said it was a gift from one of the graduates, and that you "don't look a gift horse in the mouth." The generous 2003 graduate, Denis Veshkurtsev, told a local news site he decided to "help his alma mater" in gratitude, adding, it was "only the beginning," with other lavatories and changing rooms coming next.


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The superstars, who've been rumored to be dating in the past, were photographed together, with RiRi donning stylish shades and Leo sporting a University of Michigan cap.
RiRi and her Oscar-winning pal weren't the only stars who came to party. In fact, The Weeknd, G-Eazy, Jermaine Dupri, Too $hort, and Michael B. Jordan also showed up and enjoyed bottle service in VIP.