Class Of Cones Research Articles

Developmental shifts in functional morphology of the retina in Atlantic tarpon,Megalops atlanticus(Elopomorpha: Teleostei) between four ecologically distinct life-history stages

The Atlantic tarpon, Megalops atlanticus, is a large piscivorous fish that supports economically important recreational fisheries in the Gulf of Mexico, Caribbean, and Florida Atlantic coast. Megalops atlanticus undergoes ontogenetic shifts in morphology, hatching in the open ocean as larvae (less than 1 cm in length), moving into hypoxic turbid mangrove marshes as juveniles (around 10 cm in length), and then moving into coastal oceanic waters as adults (over 100 cm in length). In this study, photoreceptor distributions, opsin distributions, and photoreceptor absorbance characteristics were studied with light microscopy, transmission electron microscopy, antiopsin immunofluorescence, and microspectrophotometry, respectively, at four ecologically distinct life-history stages--premetamorphic larva, settlement stage, juvenile, and adult. The purposes of this study were 1) to determine the extent to which the retina of M. atlanticus changes over the course of development and 2) to relate these retinal changes with ecological shifts between developmental stages. The new data presented here indicate that the M. atlanticus retina changes substantially in rod and cone distributions and absorbance characteristics over the course of development and that these changes correlate closely with those in habitat and behavior. We show that M. atlanticus has a rod-dominated retina at the larval stage (which is unusual for teleost larvae) and that the scotopic visual system becomes far better developed with maturity, adding a substantial tapetum and high densities of small, bundled, and stacked rod cells. We also show that there are shifts in cone and rod spectral sensitivities and an increase in the diversity of spectrally distinct cone classes, including the addition of ultraviolet cones as fish mature into adults.

Colour detection thresholds as a function of chromatic adaptation and light level

Colour threshold discrimination ellipses were measured for a number of states of chromatic adaptation and a range of luminance levels using the Colour Assessment and Diagnosis (CAD) test. An analysis of these results was carried out by examining the cone excitation signals along the cardinal axes that correspond to detection thresholds in the +L-M (reddish), -L+M (greenish), +S (bluish) and -S (yellowish) colour directions. The results reveal a strong linear relationship between the excitations induced by the adapting background field in each cone class and the corresponding changes needed for threshold detection. These findings suggest that the cone excitation change for threshold detection of colour signals is always the same for a given background excitation level (in any cone class), independent of the excitations generated in the other cone classes. These observations have been used to develop a model to predict colour detection thresholds for any specified background luminance and chromaticity within the range of values investigated in this study (e.g., luminances in the range 0.3 to 31 cd.m(-2) and chromaticities within the gamut of typical CRT displays). Predicted colour thresholds were found to be in close agreement with measured values with errors that do not, in general, exceed the measured within-subject variability.

Spectral sensitivity of single cones in rainbow trout ( Oncorhynchus mykiss): A whole-cell voltage clamp study

The UVS cone mechanism is known to light adapt at low intensities in comparison to other cones. We were interested in whether this property was related to higher sensitivity in UVS cones or to network adjustments in sensitivity. We investigated spectral sensitivity of 107 individual cone photoreceptors in rainbow trout ( Oncorhynchus mykiss) using a whole-cell voltage clamp technique. Mean time-to-peak response was 339 ± 90 ms and flash sensitivity for a 100 ms flash was 4.37 × 10 −3 ± 2.50 × 10 −3 pA photons −1 μm 2, with no significant differences between the UVS, SWS, MWS and LWS cone classes. The spectral sensitivity of the UVS, SWS and LWS cones conformed to the expression of SWS1, SWS2 and LWS opsin genes. The spectral sensitivity of MWS cones, however, showed clear evidence of co-expression of RH2a and RH2b opsin pigments. The fish used in this study bridged the ontogenetic stage where the MWS cones shift their expression from RH2b to RH2a.

The dimensionality of color vision in carriers of anomalous trichromacy

Some 12% of women are carriers of the mild, X-linked forms of color vision deficiencies called "anomalous trichromacy." Owing to random X chromosome inactivation, their retinae must contain four classes of cone rather than the normal three; and it has previously been speculated that these female carriers might be tetrachromatic, capable of discriminating spectral stimuli that are indistinguishable to the normal trichromat. However, the existing evidence is sparse and inconclusive. Here, we address the question using (a) a forced-choice version of the Rayleigh test, (b) a test using multidimensional scaling to reveal directly the dimensionality of the participants' color space, and (c) molecular genetic analyses to estimate the X-linked cone peak sensitivities of a selected sample of strong candidates for tetrachromacy. Our results suggest that most carriers of color anomaly do not exhibit four-dimensional color vision, and so we believe that anomalous trichromacy is unlikely to be maintained by an advantage to the carriers in discriminating colors. However, 1 of 24 obligate carriers of deuteranomaly exhibited tetrachromatic behavior on all our tests; this participant has three well-separated cone photopigments in the long-wave spectral region in addition to her short-wave cone. We assess the likelihood that behavioral tetrachromacy exists in the human population.

The Fundamental Solution and Strichartz Estimates for the Schrödinger Equation on Flat Euclidean Cones

We study the Schrödinger equation on a flat euclidean cone $${\mathbb{R}_+ \times \mathbb{S}^1_\rho}$$ of cross-sectional radius ρ > 0, developing asymptotics for the fundamental solution both in the regime near the cone point and at radial infinity. These asymptotic expansions remain uniform while approaching the intersection of the “geometric front,” the part of the solution coming from formal application of the method of images, and the “diffractive front” emerging from the cone tip. As an application, we prove Strichartz estimates for the Schrödinger propagator on this class of cones.

Genomic organization of duplicated short wave-sensitive and long wave-sensitive opsin genes in the green swordtail, Xiphophorus helleri

BackgroundLong wave-sensitive (LWS) opsin genes have undergone multiple lineage-specific duplication events throughout the evolution of teleost fishes. LWS repertoire expansions in live-bearing fishes (family Poeciliidae) have equipped multiple species in this family with up to four LWS genes. Given that color vision, especially attraction to orange male coloration, is important to mate choice within poeciliids, LWS opsins have been proposed as candidate genes driving sexual selection in this family. To date the genomic organization of these genes has not been described in the family Poeciliidae, and little is known about the mechanisms regulating the expression of LWS opsins in any teleost.ResultsTwo BAC clones containing the complete genomic repertoire of LWS opsin genes in the green swordtail fish, Xiphophorus helleri, were identified and sequenced. Three of the four LWS loci identified here were linked in a tandem array downstream of two tightly linked short wave-sensitive 2 (SWS2) opsin genes. The fourth LWS opsin gene, containing only a single intron, was not linked to the other three and is the product of a retrotransposition event. Genomic and phylogenetic results demonstrate that the LWS genes described here share a common evolutionary origin with those previously characterized in other poeciliids. Using qualitative RT-PCR and MSP we showed that each of the LWS and SWS2 opsins, as well as three other cone opsin genes and a single rod opsin gene, were expressed in the eyes of adult female and male X. helleri, contributing to six separate classes of adult retinal cone and rod cells with average λmax values of 365 nm, 405 nm, 459 nm, 499 nm, 534 nm and 568 nm. Comparative genomic analysis identified two candidate teleost opsin regulatory regions containing putative CRX binding sites and hormone response elements in upstream sequences of LWS gene regions of seven teleost species, including X. helleri.ConclusionsWe report the first complete genomic description of LWS and SWS2 genes in poeciliids. These data will serve as a reference for future work seeking to understand the relationship between LWS opsin genomic organization, gene expression, gene family evolution, sexual selection and speciation in this fish family.

How to project onto an isotone projection cone

The solution of the complementarity problem defined by a mapping f : R n → R n and a cone K ⊂ R n consists of finding the fixed points of the operator P K ∘ ( I - f ) , where P K is the projection onto the cone K and I stands for the identity mapping. For the class of isotone projection cones (cones admitting projections isotone with respect to the order relation they generate) and f satisfying certain monotonicity properties, the solution can be obtained by iterative processes (see G. Isac, A.B. Németh, Projection methods, isotone projection cones, and the complementarity problem, J. Math. Anal. Appl. 153(1) (1990) 258–275 and S.Z. Németh, Iterative methods for nonlinear complementarity problems on isotone projection cones, J. Math. Anal. Appl. 350(1) (2009) 340–347). These algorithms require computing at each step the projection onto the cone K . In general, computing the projection mapping onto a cone K is a difficult and computationally expensive problem. In this note it is shown that the projection of an arbitrary point onto an isotone projection cone in R n can be obtained by projecting recursively at most n - 1 times into subspaces of decreasing dimension. This emphasizes the efficiency of the algorithms mentioned above and furnishes a handy tool for some problems involving special isotone projection cones, as for example the non-negative monotone cones occurring in reconstruction problems (see e.g. Section 5.13 in J. Dattorro, Convex Optimization and Euclidean Distance Geometry, Meboo, 2005, v2009.04.11).

Avian retinal oil droplets: dietary manipulation of colour vision?

Avian vision is highly developed, with bird retinas containing rod and double-cone photoreceptors, plus four classes of single cones subserving tetrachromatic colour vision. Cones contain an oil droplet, rich in carotenoid pigments (except VS/ultraviolet-sensitive cones), that acts as a filter, substantially modifying light detected by the photoreceptor. Using dietary manipulations, we tested the effects of carotenoid availability on oil droplet absorbance properties in two species: Platycercus elegans and Taeniopygia guttata. Using microspectrophotometry, we determined whether manipulations affected oil droplet carotenoid concentration and whether changes would alter colour discrimination ability. In both species, increases in carotenoid concentration were found in carotenoid-supplemented birds, but only in the double cones. Magnitudes of effects of manipulations were often dependent on retinal location. The study provides, to our knowledge, the first experimental evidence of dietary intake over a short time period affecting carotenoid concentration of retinal oil droplets. Moreover, the allocation of carotenoids to the retina by both species is such that the change potentially preserves the spectral tuning of colour vision. Our study generates new insights into retinal regulation of carotenoid concentration of oil droplets, an area about which very little is known, with implications for our understanding of trade-offs in carotenoid allocation in birds.

Cones with bounded and unbounded bases and reflexivity

In this paper we prove two characterizations of reflexivity for a Banach space X . The first one is based on the existence in X of a closed convex cone with a nonempty interior such that all the bases generated by a strictly positive functional are bounded, while the second one is stated in terms of the nonexistence of a cone such that it has bounded and unbounded bases (both generated by strictly positive functionals) simultaneously. We call such a cone mixed based cone. We study the features of this class of cones. In particular, we show that every cone conically isomorphic to the nonnegative orthant ℓ + 1 of ℓ 1 is a mixed based cone and that every mixed based cone contains a conically isomorphic copy of ℓ + 1 . Moreover we give a detailed description of the structure of a mixed based cone. This approach allows us to prove some results concerning the embeddings of ℓ 1 and c 0 in a Banach space.

Evolution and spectral tuning of visual pigments in birds and mammals

Variation in the types and spectral characteristics of visual pigments is a common mechanism for the adaptation of the vertebrate visual system to prevailing light conditions. The extent of this diversity in mammals and birds is discussed in detail in this review, alongside an in-depth consideration of the molecular changes involved. In mammals, a nocturnal stage in early evolution is thought to underlie the reduction in the number of classes of cone visual pigment genes from four to only two, with the secondary loss of one of these genes in many monochromatic nocturnal and marine species. The trichromacy seen in many primates arises from either a polymorphism or duplication of one of these genes. In contrast, birds have retained the four ancestral cone visual pigment genes, with a generally conserved expression in either single or double cone classes. The loss of sensitivity to ultraviolet (UV) irradiation is a feature of both mammalian and avian visual evolution, with UV sensitivity retained among mammals by only a subset of rodents and marsupials. Where it is found in birds, it is not ancestral but newly acquired.

Gene therapy for red–green colour blindness in adult primates

Red-green colour blindness, which results from the absence of either the long- (L) or middle- (M) wavelength-sensitive visual photopigments, is the most common single locus genetic disorder. Here, the possibility of curing colour blindness using gene therapy was explored in experiments on adult monkeys that had been colour blind since birth. A third type of cone pigment was added to dichromatic retinas, providing the receptoral basis for trichromatic colour vision. This opened a new avenue to explore the requirements for establishing the neural circuits for a new dimension of colour sensation. Classic visual deprivation experiments1 have led to the expectation that neural connections established during development would not appropriately process an input that was not present from birth. Therefore, it was believed that treatment of congenital vision disorders would be ineffective unless administered to the very young. Here, however, addition of a third opsin in adult red-green colour-deficient primates was sufficient to produce trichromatic colour vision behaviour. Thus, trichromacy can arise from a single addition of a third cone class and it does not require an early developmental process. This provides a positive outlook for the potential of gene therapy to cure adult vision disorders.

Morphometry of scoria cones located on a volcano flank: A case study from Mt. Etna (Italy), based on high-resolution LiDAR data

By using new high-resolution (2 m) digital elevation model derived from the 2005 LiDAR survey of Mt. Etna volcano (Italy), our study measured the classical morphometrical parameters for scoria cones, i.e. W co (cone width), W cr (crater diameter), H (cone height) as well as volume, inclination of cone slope and substrate, and a number of other parameters for 135 scoria cones of Mt. Etna. Volume and age distribution of cones shows that there is no direct structural control on their emplacement in terms of Etna's rift zones. The cones are progressively smaller in size toward summit, which can be explained by the large volcano's feeding system and progressively frequent lava burial toward top. A careful analysis of H/ W co ratio (determined as 0.18 for other volcanic fields worldwide) shows that this ratio strongly depends on (1) the calculation method of H and (2) lava burial of cone. For Etnean cones, applying an improved method for calculating H relative to the dipping substrate results in a significantly lowered standard H/ W co ratio (0.137), which in turn questions the validity of the classical value of 0.18 in the case of large central volcanoes. The reduction of the ratio is not only due to methodology but also to the common lava burial. This can be expressed even better if H mean is used instead of H max ( H mean/ W co = 0.098). Using this measure, at Etna, well formed cones have higher ratios than structurally deformed (e. g. double or rifted) cones. Furthermore, although the sampled scoria cones at Etna have formed in a relatively narrow time interval (< 6500 yrs BP), there is a slight decrease in H/ W co corresponding to erosional changes detected globally ( H/ W co = 0.143, 0.135 and 0.115 for three age classes of Etna's scoria cones, corresponding to average slopes of 26.6, 23.9 and 23.7°). Because the morphometrical effect of position on a dipping substrate as well as lava burial exceeds the effect of erosion, we call attention to use caution in simply using the H/ W co ratio of scoria cones for detecting age, especially on large active volcanoes.

Longer wavelengths of light improve the growth, intake and feed efficiency of individually reared juvenile pikeperchSander lucioperca(L.)

We tested the effects of monochromatic light on the specific growth rate (SGR), feed intake and feed efficiency (FE) of juvenile pikeperch, Sander lucioperca (L.). Pikeperch were reared individually for 42 days in aquaria covered with blue, green, yellow or red gelatin filters or white paper (control; n=5). Linear regression analysis indicated a significant positive effect of longer wavelengths of light on the condition factor (CF), FE and SGR. The final weight, SGR and CF were significantly higher in fish reared under red than under white light, and FE was better under green, yellow and red light than under white light (Dunnett's post hoc test, P<0.05) while blue was comparable to white light in terms of the measured parameters. After the growth trial, the spectral sensitivity of photoreceptor cells in the retina was tested using microspectrophotometry, which revealed the presence of rods with λmax at ca. 530 nm and two cone classes, absorbing maximally at ca. 535 and 603 nm, all containing a porphyropsin-based pigment. These results suggest that the presence of mid and long wavelength-sensitive cones enhances visual sensitivity under mid-wavelength and long-wavelength environments, and thus supports the finding that longer wavelengths of incoming light can improve FE and SGR of the cultivated pikeperch.

Into the blue: Gene duplication and loss underlie color vision adaptations in a deep-sea chimaera, the elephant sharkCallorhinchus milii

The cartilaginous fishes reside at the base of the gnathostome lineage as the oldest extant group of jawed vertebrates. Recently, the genome of the elephant shark, Callorhinchus milii, a chimaerid holocephalan, has been sequenced and therefore becomes the first cartilaginous fish to be analyzed in this way. The chimaeras have been largely neglected and very little is known about the visual systems of these fishes. By searching the elephant shark genome, we have identified gene fragments encoding a rod visual pigment, Rh1, and three cone visual pigments, the middle wavelength-sensitive or Rh2 pigment, and two isoforms of the long wavelength-sensitive or LWS pigment, LWS1 and LWS2, but no evidence for the two short wavelength-sensitive cone classes, SWS1 and SWS2. Expression of these genes in the retina was confirmed by RT-PCR. Full-length coding sequences were used for in vitro expression and gave the following peak absorbances: Rh1 496 nm, Rh2 442 nm, LWS1 499 nm, and LWS2 548 nm. Unusually, therefore, for a deep-sea fish, the elephant shark possesses cone pigments and the potential for trichromacy. Compared with other vertebrates, the elephant shark Rh2 and LWS1 pigments are the shortest wavelength-shifted pigments of their respective classes known to date. The mechanisms for this are discussed and we provide experimental evidence that the elephant shark LWS1 pigment uses a novel tuning mechanism to achieve the short wavelength shift to 499 nm, which inactivates the chloride-binding site. Our findings have important implications for the present knowledge of color vision evolution in early vertebrates.

Identification And Analysis Of CNGA3 And CNGB3 From Zebrafish

Cyclic nucleotide-gated (CNG) channels are a crucial component of the phototransduction cascade in vertebrate photoreceptors. The opening and closure of these channels and consequently the influx of sodium and calcium ions into the photoreceptor outer segment is directed by the intracellular light-dependent cGMP level. Cone CNG channels are heterooligomers consisting of two A3- and two B3-subunits, which are encoded by the CNGA3 and the CNGB3 gene. In both genes mutations have been identified, which can lead to a dysfunction of the CNG channels in cone photoreceptors. In humans this results in the autosomal-recessively inherited disease achromatopsia (color blindness). In order to characterize CNG channels in zebrafish, which possess four morphologically and physiologically distinct classes of cones, we have identified two homologous candidate genes for CNGA3 and two for CNGB3 by in silico database analyses. All four genes as well as a splice variant of CNGA3-1 have been cloned and were heterologously expressed in HEK293 cells. Subsequently, the zebrafish CNG channels were functionally characterized by calcium imaging and patch-clamp measurements. The retinal expression of all four genes has been confirmed by RT-PCR. In silico analyses revealed, that the two CNGA3 candidates are located at two different locations in the zebrafish genome and are presumably a result of the whole genome duplication as it is known for several genes in zebrafish. In contrast to that, the two CNGB3 candidates are located in a tandem as a result of an additional gene duplication event. ZfCNGA3-1 and zfCNGA3-2 have 62 % identity with the human CNGA3 protein. ZfCNGB3-1 has 43 % and zfCNGB3-2 has 49 % identity with human CNGB3.

Generation of pseudo-Bessel beams at THz frequencies by use of binary axicons

In order to miniaturize and integrate conveniently in THz quasi-optical systems, binary axicons, based on binary optical ideas, are introduced in our paper and designed for generating pseudo-Bessel beams at THz frequencies. The designed binary axicons are easier to fabricate than holographic axicons, more compact and thus less lossy in the material when compared with classical cone axicons. To calculate the electromagnetic fields diffracted by binary axicons precisely, a two-dimension finite-difference time-domain (2-D FDTD) method in conjunction with Stratton- Chu formulas are employed in this paper. Applying this method, the properties of pseudo-Bessel beams produced by binary axicons are analyzed, and a brief summary is given in the end.

Alternative Theorems for Quadratic Inequality Systems and Global Quadratic Optimization

We establish alternative theorems for quadratic inequality systems. Consequently, we obtain Lagrange multiplier characterizations of global optimality for classes of nonconvex quadratic optimization problems. We present a generalization of Dine's theorem to a system of two homogeneous quadratic functions with a regular cone. The class of regular cones are cones K for which $(K\cup-K)$ is a subspace. As a consequence, we establish a generalization of the powerful S-lemma, which paves the way to obtain a complete characterization of global optimality for a general quadratic optimization model problem involving a system of equality constraints in addition to a single quadratic inequality constraint. We then present an alternative theorem for a system of three nonhomogeneous inequalities by way of establishing the convexity of the joint-range of three homogeneous quadratic functions using a regular cone. This yields Lagrange multiplier characterizations of global optimality for classes of trust-region type problems with two inequality constraints. Finally, we establish an alternative theorem for systems involving an arbitrary finite number of quadratic inequalities involving Z-matrices, which are matrices with nonpositive off diagonal elements, and present necessary and sufficient conditions for global optimality for classes of nonconvex inequality constrained quadratic optimization problems.

Cone signals for spectacle-lens compensation: Differential responses to short and long wavelengths

Chick eyes compensate for defocus imposed by spectacle lenses by making compensatory changes in eye length and choroidal thickness, a laboratory model of emmetropization. To investigate the roles of longitudinal chromatic aberration and of chromatic mechanisms in emmetropization, we examined the participation of different cone classes, and we compared the efficacy of lens compensation under monochromatic illumination with that under white light of the same illuminance to the chick eye. Chicks wore positive or negative 6 D or 8 D lenses on one eye for 3 days, under either blue (460 nm) or red (620 nm) light at 0.67 lux or under white light at 0.67 or 0.2 lux (all measures are corrected for chick photopic sensitivity). The illumination conditions were chosen to differentially stimulate either the short-wavelength and ultraviolet cones or the long-wavelength and double cones. Measurements are expressed as the relative change: the inter-ocular difference in the amount of change over the 3 days of lens wear. We find that under this low illumination the two components of lens compensation were differentially affected by the monochromatic illumination: in blue light lens compensation was mainly due to changes in eye length, whereas in red light lens compensation was mainly due to changes in choroidal thickness. In general, white light produced better lens compensation than monochromatic illumination. Negative lenses Under white light negative lenses caused an increase in eye length (60 μm) together with a decrease in choroidal thickness (−51 μm) relative to the fellow eye. Under blue light, although there was an increase in eye length (32 μm), there was no change in choroidal thickness (5 μm). In contrast, under red light there was a decrease in choroidal thickness (−62 μm) but no increase in eye length (8 μm). Relative ocular elongation was the same in white and monochromatic light. Positive lenses Under white light positive lenses caused a decrease in eye length (−142 μm) together with an increase in choroidal thickness (68 μm) relative to the fellow eye. Under blue light, there was a decrease in eye length (−64 μm), but no change in choroidal thickness (2 μm). In contrast, under red light there was an increase (90 μm) in choroidal thickness but less of a decrease (−36 μm) in eye length. Lens compensation by inhibition of ocular elongation was less effective under monochromatic illumination than under white light (white v red: p = 0.003; white v blue p = .014). The differential effects of red and blue light on the choroidal and ocular length compensatory responses suggest that they are driven by different proportions of the cone-types, implying that, although chromatic contrast is not essential for lens compensation and presumably for emmetropization as well, the retinal substrates exist for utilizing chromatic contrast in these compensatory responses. The generally better lens compensation in white than monochromatic illumination suggests that longitudinal chromatic aberration may be used in lens compensation.

Evolution of vertebrate visual pigments

The visual pigments of vertebrates evolved about 500 million years ago, before the major evolutionary step of the development of jaws. Four spectrally distinct classes of cone opsin evolved through gene duplication, followed by the rod opsin class that arose from the duplication of the middle-wave-sensitive cone opsin. All four cone classes are present in many extant teleost fish, reptiles and birds, but one or more classes have been lost in primitive fish, amphibians and mammals. Gene duplication within the cone classes, especially in teleosts, has resulted in multiple opsins being available, both temporally and spatially, during development.

Frictional contact models with local compliance: semismooth formulation

AbstractA complementarity‐based, 3‐dimensional frictional contact model with local compliance and damping was introduced in the Ph.D. thesis [45] and the paper [47] and was subsequently studied extensively in [31, 49]. In this paper, we examine a variant of this model where there is no damping in the normal contact forces but there is coupled stiffness between the normal and tangential forces via body deformations. We show that this frictional contact model admits a formulation as an ordinary differential equation with a boundedly Lipschitz continuous, albeit implicitly defined, semismooth right‐hand side with global linear growth. Several major consequences follow from such a formulation: (a) existence and uniqueness of a continuously differentiable solution trajectory originated from an arbitrary initial state, (b) finite contact forces that are semismooth functions of the system state, (c) semismooth dependence of the trajectory on the initial state, and (d) convergence of a shooting method for solving two‐point boundary problems. The derived results are valid for both a dynamic model and a quasistatic model, the latter being one in which inertia effects are ignored, and for a broad class of friction cones that include the well‐known quadratic Coulomb cone and its polygonal approximations. Part II of this work establishes the absence of Zeno states in such a frictional contact model.