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Pattern formation and eyespot determination in butterfly wings.

Butterfly wings display pattern elements of many types and colors. To identify the molecular processes underlying the generation of these patterns, several butterfly cognates of Drosophila appendage patterning genes have been cloned and their expression patterns have been analyzed. Butterfly wing patterns are organized by two spatial coordinate systems. One system specifies positional information with respect to the entire wing field and is conserved between fruit flies and butterflies. A second system, superimposed on the general system and involving several of the same genes, operates within each wing subdivision to elaborate discrete pattern elements. Eyespots, which form from discrete developmental organizers, are marked by Distal-less gene expression. These circular pattern elements appear to be generated by a process similar to, and perhaps evolved from, proximodistal pattern formation in insect appendages.

Amino Acid Sequence↗

Ecological correlates of polyphenism and gregarious roosting in the grass yellow butterfly Eurema elathea (Pieridae).

Eurema elathea adults were census weekly (1992-1994) in six night-roosts around a forest fragment on a farm, and in two roosts in the urban area of Uberlândia, Minas Gerais, Brazil. Males were grouped in six phenotypic classes. These were based on a range between having a conspicuous wide black bar at the dorsal forewing inner margin (wet season dark morphs) and the absence of that bar (dry season light morphs). The body mass and wing area of co-occurring morphs were compared: differents morphs showed similar means. The abundance of butterflies and morph frequencies varied in close relation to humidity (rainfall). Individuals were infrequent and monomorphically dark in the wet season while light morphs predominated in dry periods when population peaked. A lower fraction of recaptured individuals and higher recruitment were recorded compared to other night-roosting butterflies. Dispersal potential was similar between the sexes and varied seasonally with a more sedentary population in dry periods. The maximum residence time recorded was 91 days for a female and 84 days for a male. The fraction of individuals that moved from one roosting site to another was similar in both sexes and male morphs, but significantly higher on the farm than in the urban area. Also, a significantly higher fraction (21.3%) of marked butterflies was recaptured in the urban area than on the farm (15.6%), suggesting a behavioral modification for sedentariness in the urban individuals. The selective forces shaping a gregarious roosting habit in E. elathea and other butterflies are discussed and a protocooperational strategy for saving energy is proposed.

Animals↗

Nonrandom movement behavior at habitat boundaries in two butterfly species: implications for dispersal.

We observed meadow brown (Maniola jurtina) and gatekeeper (Pyronia tithonus) butterflies at habitat boundaries and observed spontaneous movements out of suitable habitat in order to investigate such movements in relation to dispersal. We found that butterflies of both species were aware of the position of a highly permeable habitat boundary without needing to cross it. Nevertheless, a considerable proportion of butterflies close to the boundary left their habitat (25-43%). Butterflies that crossed the boundary, and moved substantial distances into unsuitable habitat (up to 350 m in M. jurtina and 70 m in P. tithonus), usually returned to their original habitat patch (98-100%). Movement trajectories, at least in M. jurtina, were significantly different from, and more directed and systematic than, a correlated random walk. Approximately 70-80% of spontaneous movements into unsuitable habitat in both species were "foray" loops comparable to those described in mammals and birds. We conclude that, since migrants seemed to have considerable control over leaving their patch and over their subsequent movement trajectories, chance encounter rates with habitat boundaries, and indeed habitat leaving rates, might be less crucial in determining dispersal rates than is usually assumed. In addition, random dispersal trajectories should not be taken for granted in population or evolution models.

Animals↗

Building phenological models from presence/absence data for a butterfly fauna.

Species phenology is increasingly being used to explore the effects of climate change and other environmental stressors. Long-term monitoring data sets are essential for understanding both patterns manifest by individual species and more complex patterns evident at the community level. This study used records of 78 butterfly species observed on 626 days across 27 years at a site in northern California, USA, to build quadratic logistic regression models of the observation probability of each species for each day of the year. Daily species probabilities were summed to develop a potential aggregate species richness (PASR) model, indicating expected daily species richness. Daily positive and negative contributions to PASR were calculated, which can be used to target optimum sampling time frames. Residuals to PASR indicate a rate of decline of 0.12 species per year over the course of the study. When PASR was calculated for wet and dry years, wet years were found to delay group phenology by up to 17 days and reduce the maximum annual expected species from 32.36 to 30. Three tests to determine how well the PASR model reflected the butterfly fauna dynamics were all positive: We correlated probabilities developed with species presence/absence data to observed abundance by species, tested species' predicted phenological patterns against known biological characteristics, and compared the PASR curve to a spline-fitted curve calculated from the original species richness observations. Modeling individual species' flight windows was possible from presence/absence data, an approach that could be used on other similar records for butterfly communities with seasonal phenologies, and for common species with far fewer dates than used here. It also provided a method to assess sample frequency guidelines for other butterfly monitoring programs.

Animals↗

Tungstate-induced color-pattern modifications of butterfly wings are independent of stress response and ecdysteroid effect.

Systemic injections of sodium tungstate, a protein-tyrosine phosphatase (PTPase) inhibitor, to pupae immediately after pupation have been shown to efficiently produce characteristic color-pattern modifications on the wings of many species of butterflies. Here we demonstrated that the tungstate-induced modification pattern was entirely different from other chemically-induced ones in a species of nymphalid butterfly Junonia (Precis) orithya. In this species, the systemic injections of tungstate produced characteristic expansion of black area and shrinkage of white area together with the move of parafocal elements toward the wing base. Overall, pattern boundaries became obscure. In contrast, an entirely different modification pattern, overall darkening of wings, was observed by the injections of stress-inducing chemicals, thapsigargin, ionomycin, or geldanamycin, to pupae under the rearing conditions for the adult summer form. On the ventral wings, this darkening was due to an increase of the proportion of peppered dark scales, which was reminiscent of the natural fall form of this species. Under the same rearing conditions, the injections of ecdysteroid, which is a well-known hormone being responsible for the seasonal polyphenism of nymphalid butterflies, yielded overall expansion of orange area especially around eyespots. Taken together, we conclude that the tungstate-induced modifications are clearly distinguishable from those of stress response and ecdysteroid effect. This conclusion then suggests that the putative PTPase signaling pathway that is sensitive to tungstate uniquely contributes to the wing-wide color-pattern development in butterflies.

Animals↗

Territorial defence in speckled wood butterflies: do the hottest males always win?

Males of the butterfly Pararge aegeria defend sun spot territories on the forest floor as a strategy to locate females: the defence involves a spiral flight between two competing males. The success of any particular fight has been interpreted as following the simple rule 'resident wins, intruder loses' as predicted by game theory models. Here we use field experiments to show that temperature has a marked effect on the territorial defence of these ectothermic insects. The spiral flights were true contests, not conventional signals, so the flight endurance of the butterflies was an important factor in determining the result of the contest. Butterflies that were warmer could fly for longer and were more likely to win a contest; therefore temperature was an important component of contest asymmetry, and resident butterflies that increased their body temperatures by basking in sun spots were more likely to win than the intruding males which might have cooled down during flight around the site. The rule that the resident always wins is not disputed; however, the causal factors underpinning the rule are questioned. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Colour characterization of a Morpho butterfly wing-scale using a high accuracy nonstandard finite-difference time-domain method.

In certain species of moths and butterflies iridescent colours arise from subwavelength diffractive structures. The optical properties of such a structure depend strongly on wavelength, incidence angle and state of polarization of illuminating radiation and on the viewing angle. Such structures can be analyzed only by solving Maxwell's equations, but since analytical solutions exist for only a few simple, highly symmetric structures numerical methods must be employed. We investigated the optical properties of butterfly wings in two dimensions by simulating a scale structure using a high accuracy version of nonstandard finite-difference time-domain algorithm. The simulated structure is a computer-generated model of a certain quasi-periodic arrangement of tree-like structures observed in the transmission electron micrograph (TEM) image of a transverse cross-section of a single scale from Morpho butterfly wings. We assumed that the structure is made of a slightly lossy dielectric material. We checked the accuracy and validity of our approach, by computing scattered field intensities due to an infinite cylinder and compared the results with analytical calculations using Mie theory. Next we deduced the wavelength dependence of a real refractive index and an absorption coefficient for the ground scales on the wings of Morpho sulkowskyi butterfly by computing the reflectivity and transmissivity spectrum of a scale at normal incidence, and comparing with experimental measurements. Finally, we calculated the tristimulus values and corresponding colour coordinates for various viewing directions from the scale's far-field reflectivity and transmissivity spectra to characterize its colour rendering abilities.

Journal Article↗

Tetranuclear and Octanuclear Manganese Carboxylate Clusters: Preparation and Reactivity of (NBu(n)(4))[Mn(4)O(2)(O(2)CPh)(9)(H(2)O)] and Synthesis of (NBu(n)(4))(2)[Mn(8)O(4)(O(2)CPh)(12)(Et(2)mal)(2)(H(2)O)(2)] with a "Linked-Butterfly" Structure.

The reaction of Mn(O(2)CPh)(2).2H(2)O and PhCO(2)H in EtOH/MeCN with NBu(n)(4)MnO(4) gives (NBu(n)(4))[Mn(4)O(2)(O(2)CPh)(9)(H(2)O)] (4) in high yield (85-95%). Complex 4 crystallizes in monoclinic space group P2(1)/c with the following unit cell parameters at -129 degrees C: a = 17.394(3) Å, b = 19.040(3) Å, c = 25.660(5) Å, beta = 103.51(1) degrees, V = 8262.7 Å(3), Z = 4; the structure was refined on F to R (R(w)) = 9.11% (9.26%) using 4590 unique reflections with F > 2.33sigma(F). The anion of 4 consists of a [Mn(4)(&mgr;(3)-O)(2)](8+) core with a "butterfly" disposition of four Mn(III) atoms. In addition to seven bridging PhCO(2)(-) groups, there is a chelating PhCO(2)(-) group at one "wingtip" Mn atom and terminal PhCO(2)(-) and H(2)O groups at the other. Complex 4 is an excellent steppingstone to other [Mn(4)O(2)]-containing species. Treatment of 4 with 2,2-diethylmalonate (2 equiv) leads to isolation of (NBu(n)(4))(2)[Mn(8)O(4)(O(2)CPh)(12)(Et(2)mal)(2)(H(2)O)(2)] (5) in 45% yield after recrystallization. Complex 5 is mixed-valent (2Mn(II),6Mn(III)) and contains an [Mn(8)O(4)](14+) core that consists of two [Mn(4)O(2)](7+) (Mn(II),3Mn(III)) butterfly units linked together by one of the &mgr;(3)-O(2)(-) ions in each unit bridging to one of the body Mn atoms in the other unit, and thus converting to &mgr;(4)-O(2)(-) modes. The Mn(II) ions are in wingtip positions. The Et(2)mal(2)(-) groups each bridge two wingtip Mn atoms from different butterfly units, providing additional linkage between the halves of the molecule. Complex 5.4CH(2)Cl(2) crystallizes in monoclinic space group P2(1)/c with the following unit cell parameters at -165 degrees C: a = 16.247(5) Å, b = 27.190(8) Å, c = 17.715(5) Å, beta = 113.95(1) degrees, V = 7152.0 Å(3), Z = 4; the structure was refined on F to R (R(w)) = 8.36 (8.61%) using 4133 unique reflections with F > 3sigma(F). The reaction of 4 with 2 equiv of bpy or picolinic acid (picH) yields the known complex Mn(4)O(2)(O(2)CPh)(7)(bpy)(2) (2), containing Mn(II),3Mn(III), or (NBu(n)(4))[Mn(4)O(2)(O(2)CPh)(7)(pic)(2)] (6), containing 4Mn(III). Treatment of 4 with dibenzoylmethane (dbmH, 2 equiv) gives the mono-chelate product (NBu(n)(4))[Mn(4)O(2)(O(2)CPh)(8)(dbm)] (7); ligation of a second chelate group requires treatment of 7 with Na(dbm), which yields (NBu(n)(4))[Mn(4)O(2)(O(2)CPh)(7)(dbm)(2)] (8). Complexes 7 and 8 both contain a [Mn(4)O(2)](8+) (4Mn(III)) butterfly unit. Complex 7 contains chelating dbm(-) and chelating PhCO(2)(-) at the two wingtip positions, whereas 8 contains two chelating dbm(-) groups at these positions, as in 2 and 6. Complex 7.2CH(2)Cl(2) crystallizes in monoclinic space group P2(1) with the following unit cell parameters at -170 degrees C: a = 18.169(3) Å, b = 19.678(4) Å, c = 25.036(4) Å, beta = 101.49(1) degrees, V = 8771.7 Å(3), Z = 4; the structure was refined on F to R (R(w)) = 7.36% (7.59%) using 10 782 unique reflections with F > 3sigma(F). Variable-temperature magnetic susceptibility studies have been carried out on powdered samples of complexes 2 and 5 in a 10.0 kG field in the 5.0-320.0 K range. The effective magnetic moment (&mgr;(eff)) for 2 gradually decreases from 8.61 &mgr;(B) per molecule at 320.0 K to 5.71 &mgr;(B) at 13.0 K and then increases slightly to 5.91 &mgr;(B) at 5.0 K. For 5, &mgr;(eff) gradually decreases from 10.54 &mgr;(B) per molecule at 320.0 K to 8.42 &mgr;(B) at 40.0 K, followed by a more rapid decrease to 6.02 &mgr;(B) at 5.0 K. On the basis of the crystal structure of 5 showing the single Mn(II) ion in each [Mn(4)O(2)](7+) subcore to be at a wingtip position, the Mn(II) ion in 2 was concluded to be at a wingtip position also. Employing the reasonable approximation that J(w)(b)(Mn(II)/Mn(III)) = J(w)(b)(Mn(III)/M(III)), where J(w)(b) is the magnetic exchange interaction between wingtip (w) and body (b) Mn ions of the indicated oxidation state, a theoretical chi(M) vs T expression was derived and used to fit the experimental molar magnetic susceptibility (chi(M)) vs T data. The obtained fitting parameters were J(w)(b) = -3.9 cm(-)(1), J(b)(b) = -9.2 cm(-)(1), and g = 1.80. These values suggest a S(T) = (5)/(2) ground state spin for 2, which was confirmed by magnetization vs field measurements in the 0.5-50.0 kG magnetic field range and 2.0-30.0 K temperature range. For complex 5, since the two bonds connecting the two [Mn(4)O(2)](7+) units are Jahn-Teller elongated and weak, it was assumed that complex 5 could be treated, to a first approximation, as consisting of weakly-interacting halves; the magnetic susceptibility data for 5 at temperatures >/=40 K were therefore fit to the same theoretical expression as used for 2, and the fitting parameters were J(w)(b) = -14.0 cm(-)(1) and J(b)(b) = -30.5 cm(-)(1), with g = 1.93 (held constant). These values suggest an S(T) = (5)/(2) ground state spin for each [Mn(4)O(2)](7+) unit of 5, as found for 2. The interactions between the subunits are difficult to incorporate into this model, and the true ground state spin value of the entire Mn(8) anion was therefore determined by magnetization vs field studies, which showed the ground state of 5 to be S(T) = 3. The results of the studies on 2 and 5 are considered with respect to spin frustration effects within the [Mn(4)O(2)](7+) units. Complexes 2 and 5 are EPR-active and -silent, respectively, consistent with their S(T) = (5)/(2) and S(T) = 3 ground states, respectively.

Journal Article↗

Relationships between energetic, stroke determinants, and velocity in butterfly.

The purpose of this study was to identify the relationship between the bioenergetical and the biomechanical variables (stroke parameters), through a range of swimming velocities, in butterfly stroke. Three male and one female butterflier of international level were submitted to an incremental set of 200-m butterfly swims. The starting velocity was 1.18 m . s (-1) for the males and 1.03 m . s (-1) for the female swimmer. Thereafter, the velocity was increased by 0.05 m . s (-1) after each swim until exhaustion. Cardio-pulmonary and gas exchange parameters were measured breath by breath for each swim to analyze oxygen consumption and other energetic parameters by portable metabolic cart (K4b (2), Cosmed, Rome, Italy). A respiratory snorkel and valve system with low hydrodynamic resistance was used to measure pulmonary ventilation and to collect breathing air samples. Blood samples from the ear lobe were collected before and after each swim to analyze blood lactate concentration (YSI 1500 L, Yellow Springs, US). Total energy expenditure (E (tot)), energetic cost (EC), stroke frequency (SF), stroke length (SL), mean swimming velocity (V), and stroke index (SI) were calculated for each lap and average for each 200-m stage. Correlation coefficients between E (tot) and V, EC, and SF, as well as between EC and SI were statistically significant. For the relation between EC and SL, only one regression equation presented a correlation coefficient with statistical significance. Relations between SF and V, as well as between SI and V were significant in all of the swimmers. Only two individual regression equations presented statistically significant correlation coefficient values for the relation established between V and the SL. As a conclusion, the present sample of swims demonstrated large inter individual variations concerning the relationships between bioenergetic and biomechanical variables in butterfly stroke. Practitioners should be encouraged to analyze the relationships between V, SF, and SL individually to detect the deflection point in SL in function of swimming velocity to further determine appropriate training intensities when trying to improve EC.

Adolescent↗

Buried tension sutures: force-tension comparisons of pulley, double butterfly, mattress, and simple interrupted suture patterns.

OBJECTIVES: To compare suture patterns (simple interrupted, modified pulley, horizontal mattress, double butterfly) by use of a novel in vitro model that quantifies force required for closure against tension. STUDY DESIGN: In vitro study. SAMPLE POPULATION: Suture constructs. METHODS: An in vitro suturing frame was constructed with extension springs to provide a standard resistance against movement of 2 panels of loops toward each other. Four different suture patterns were applied to the frame in randomized trials. The frame was affixed to a tensiometer that measured the force required for each suture pattern to close a fixed distance (3.81 cm) at a constant rate (5.08 cm/min) against the tension (0.6 kg/cm) of the extension springs. The closure distance and tension were selected subjectively to mimic tension encountered clinically during wound reconstruction oncologic surgical procedures. RESULTS: The modified pulley suture required the least force to close, followed by the double butterfly, simple interrupted, and horizontal mattress patterns. CONCLUSIONS: The modified pulley suture and double butterfly suture patterns require less force to close a given distance under tension than either simple interrupted or horizontal mattress suture patterns. CLINICAL RELEVANCE: Application of tension relieving suture techniques for fascia, subcutaneous, and buried-knot subdermal suturing should be considered to facilitate direct closure of wounds under tension. Modified pulley and double butterfly suture patterns offer some mechanical advantage by requiring less force to achieve closure compared with other suture patterns that might be used in reconstructive surgery.

Animals↗

Rationalizing venepuncture pain: comparison of lignocaine injection, Butterfly (21 gauge and 23 gauge) and Venflon (20 gauge).

Two hundred and seventy-eight patients scheduled for all types of surgery and premedicated with diazepam and metoclopramide were randomly allocated to one of four groups to compare the relative pain of an injection of 0.25 ml of lignocaine 1% via a 25 gauge needle with the pain of the siting of a 21 gauge Butterfly (Abbott), 23 gauge Butterfly or a 20 gauge Venflon (Vigo Spectramed). The injection of lignocaine and insertion of the 23 gauge Butterfly were associated with the least complaints of pain and least observed responses to pain. The 21 gauge Butterfly and 20 gauge Venflon were associated with complaints of greater pain and more pain responses. We conclude that a pre-cannulation injection of lignocaine causes minimal discomfort and is the most appropriate means of reducing the discomfort of venous cannulation when not using skin penetrating analgesic creams.

Adolescent↗

Subcutaneous infusion: non-metal cannulae vs metal butterfly needles.

This review aimed to evaluate the effectiveness of non-metal cannulae compared to metal butterfly needles in maintaining subcutaneous infusion sites in patients receiving palliative care. The Cochrane Library, Medline, Pre-Medline, Embase, CINAHL, Amed and Cancerlit were searched for relevant studies. Controlled trials comparing non-metal cannulae with metal butterfly needles for giving subcutaneous infusion to palliative care patients were included. The outcome considered was site duration in terms of hours of patency or until change was required. Four trials met the inclusion criteria although overall quality was poor due to low follow-up. Studies examined either Teflon or Vialon-coated catheters. All studies showed non-metal cannulae to be superior to metal. In individual studies estimates in mean increase in duration of the site range from 21 to 159 hours. It seems that non-metal cannulae are more effective in maintaining the duration of subcutaneous infusion sites than butterfly needles. Both types of non-metal catheter showed clear benefits. This review has not examined other outcomes but in general adverse effects lead to the removal of the catheter and so would be reflected in the outcome of considered. Although historically non-metal cannulae have been considerably more expensive there is now little difference between metal and Teflon-coated catheters. This review recommends the use of non-metal cannulae in preference to butterfly needles.

Clinical Trials as Topic↗

First-trimester sonographic diagnosis of holoprosencephaly: value of the "butterfly" sign.

OBJECTIVE: To study the value of choroid plexus dysmorphology as a screening tool for the first-trimester sonographic diagnosis of holoprosencephaly in a high-risk population. METHODS: A total of 378 consecutive pregnancies undergoing chorionic villus sampling between 11 and 14 weeks' gestation were scanned before the procedure, following the recommendations of the Fetal Medicine Foundation (London, England). A cross-sectional view of the fetal brain, including the visualization of both choroid plexuses (the "butterfly" sign), was obtained in all cases. RESULTS: There were 3 cases in which the butterfly sign was not identified. In these cases, the first-trimester diagnosis of holoprosencephaly was confirmed by the presence of a single monoventricular cavity and fused thalami. Two of these fetuses had features of facial dysmorphism at the time of presentation and 2 had extracranial anomalies, including a cystic hygroma in 1 and a small omphalocele and polydactyly in another. Chromosomal analysis showed trisomy 13 in 2 cases and a ring chromosome 13 in the other. CONCLUSIONS: This series suggests that failure to identify the butterfly sign is a warning sign of holoprosencephaly in the first trimester. Systematic identification of the butterfly sign at the time of sonographic assessment of nuchal translucency provides a valuable tool for the early screening of holoprosencephaly.

Abnormalities, Multiple↗

[Butterfly-shaped dystrophy of the macula with absent pre-beta-fraction of the electropherogram of the serum lipoproteins (author's transl)].

A case of butterfly-shaped dystrophy of the macula is described. Clinical and electrophysiological data are compared with those of 11 cases described in the literature. An aberrant electropherogram of the lipoproteins examined at two different laboratories was found to be present. The serum lipoproteins showed total lack of the pre-beta-fraction. Two children of the patient were examined, too. They showed minimal macular changes which however did not justify the diagnosis of butterfly-shaped dystrophy. Their pre-beta-lipoprotein fractions of the electropherogram were below the Finish standards. The proband is the first case of butterfly-shaped macular dystrophy described from Finland and therefore it was not possible to control the laboratory findings on other cases. The absence of the pre-beta-lipoprotein fraction might be an important feature of butterfly-shaped dystrophy of the macula.

Adolescent↗

Overestimation of neonatal PO2 by collection of arterial blood gas values with the butterfly infusion set.

The butterfly, or scalp vein infusion set, is a widely used and convenient way to collect arterial blood gas samples in neonates. Analyzing arterial blood gas values by using this technique in a clinical situation, we obtained elevated values of PO2 in blood samples collected and measured from butterfly tubing in comparison with tuberculin syringe controls. Diffusion of gas through the polyvinylchloride tubing was suggested and proved by a study of pairs of butterfly and tuberculin syringes filled with venous blood from the blood bank. In conclusion, the butterfly gives falsely elevated PO2 values as a result of gas diffusion and is not a reliable technique for arterial blood gas sampling in neonates.

Blood Specimen Collection↗

Not all butterfly eyes are created equal: rhodopsin absorption spectra, molecular identification, and localization of ultraviolet-, blue-, and green-sensitive rhodopsin-encoding mRNAs in the retina of Vanessa cardui.

Surveys of spectral sensitivities, visual pigment spectra, and opsin gene sequences have indicated that all butterfly eyes contain ultraviolet-, blue-, and green-sensitive rhodopsins. Some species also contain a fourth or fifth type, related in amino acid sequence to green-sensitive insect rhodopsins, but red shifted in absorbance. By combining electron microscopy, epi-microspectrophotometry, and polymerase chain reaction cloning, we found that the compound eye of Vanessa cardui has the typical ultrastructural features of the butterfly retina but contains only the three common insect rhodopsins. We estimated lambda-max values and relative densities of the rhodopsins in the Vanessa retina (0.72, P530; 0.12, P470; and 0.15, P360) from microspectrophotometric measurements and calculations based on a computational model of reflectance spectra. We isolated three opsin-encoding cDNA fragments that were identified with P530, P470, and P360 by homology to the well-characterized insect rhodopsin families. The retinal mosaic was mapped by opsin mRNA in situ hybridization and found to contain three kinds of ommatidia with respect to their patterns of short wavelength rhodopsin expression. In some ommatidia, P360 or P470 was expressed in R1 and R2 opposed receptor cells; in others, one cell expressed P360, whereas its complement expressed P470. P530 was expressed in the other seven cells of all ommatidia. P470-expressing cells were abundant in the ventral retina but nearly absent dorsally. Our results indicated that there are major differences between the color vision systems of nymphalid and papilionid butterflies: the nymphalid Vanessa has a simpler, trichromatic, system than do the tetrachromatic papilionids that have been studied.

Amino Acid Sequence↗

Ultrastructure of the extraocular photoreceptor in the genitalia of a butterfly, Papilio xuthus.

This paper describes the ultrastructure of a sensory neuron found in the extraocular photoreceptive site on the butterfly genitalia. Our previous studies have shown that there are two pairs of the photoreceptive sites in a butterfly (four per individual). Each photoreceptive site is recognizable by a transparent area in the pigmented cuticle of the genitalia. From the nerve that extends from the transparent cuticle to the last abdominal ganglion, a sustained train of single unit spikes can be recorded in response to a light flash. The single unit spikes disappear when the transparent cuticle is covered, thus indicating that a single photoreceptor exists close to it. Here, we examined complete serial semithin sections of plastic-embedded photoreceptive sites of both male and female, and observed an ovoid structure close to the transparent cuticle. The structure contained the cell body of a sensory neuron whose axon was in the nerve branch from which the photoresponse had been recorded. Further electron microscopy revealed that the cell body extended a few distal processes that protrude tubular membrane from the tip, forming a structure resembling the phaosome (from Greek; phaos = light, some = body) which was first described in the earthworm dermal photoreceptors. The sensory neuron was also found in a surgically isolated nerve-photoreceptor preparation that responded to the light. We therefore propose that the phaosome-containing sensory neuron is the butterfly genital photoreceptor.

Animals↗

Elements of butterfly wing patterns.

The color patterns on the wings of butterflies are unique among animal color patterns in that the elements that make up the overall pattern are individuated. Unlike the spots and stripes of vertebrate color patterns, the elements of butterfly wing patterns have identities that can be traced from species to species, and typically across genera and families. Because of this identity it is possible to recognize homologies among pattern elements and to study their evolution and diversification. Individuated pattern elements evolved from non-individuated precursors by compartmentalization of the wing into areas that became developmentally autonomous with respect to color pattern formation. Developmental compartmentalization led to the evolution of serially repeated elements and the emergence of serial homology. In these compartments, serial homologues were able to acquire site-specific developmental regulation and this, in turn, allowed them to diverge morphologically. Compartmentalization of the wing also reduced the developmental correlation among pattern elements. The release from this developmental constraint, we believe, enabled the great evolutionary radiation of butterfly wing patterns. During pattern evolution, the same set of individual pattern elements is arranged in novel ways to produce species-specific patterns, including such adaptations as mimicry and camouflage.

Adaptation, Physiological↗